Philippine Land Mammals (Part 2 of 2) by Filipiniana Online

298

PHILIPPINE LAND MAMMALS

width slightly below middle of inner margin; muzzle broad and conical, nearly naked; feet rather large; wings attach to base of toes; postcalcaneal lobe distinct, not deep; extreme tip of tail projecting. Skull and teeth.-Upper incisors sloping considerably inwards and forwards, bifid; outer incisor on each side slender and shorter than outer cusp of inner incisor; lower incisors not crowded in tooth row; second upper premolar separated by a short interval from canine; first small internal premolar almost concealed by canine. Measurements of Pipistrellus tenuis (Temnninck). mm.

Head and body Tail Head Ear Tragus Forearm Thumb Third finger Fifth finger Tibia Foot

419 25.4 13.97 11.26 4.57 29.2 7.1 50.8 38.1 12.7 6.35

Remarks.-The species is included on Hoffman's record for Mindanao. The type locality is Java and Sumatra. PIPISTRELLUS ABRAMUS (Temminck)

Vesperlilio abramus TElMMINCK, Monog. Mamm. 2 (1841) 232. Vesperugo abramus DOBSON, Monog. Asiat. Chirop. (1876) 97; Cat. Chirop. Brit. Mus. (1878) 226; BLANFORD, Journ. Asiat. Soc. Bengal 57: 266; Fauna Brit. India, Mamm. (1888-91) 313, fig. 73.

Description of the species.-{From No. 342, E. H. Taylor collection; collected in Manila, May, 1920, by Charles S. Banks.) Ears moderate, rather rounded at top, anterior margin straight to near base; posterior edge rather curved, with two slight notches; a prominent lobe below tragus; tragus curved slightly forward, inner margin concave, outer rounded; tip somewhat rounded; basal lobe small with a slight notch above it; muzzle blunt, swollen by glands with a broad median depression; forehead and crown of head thickly furred; anterior part of muzzle covered with fine short hair, appearing nearly naked in adults; feet small, toes much longer than sole; wing membrane attaches

299

PIPISTRELLUS

on outer edge of base of outer toe; a distinct, rather elongate postcalcaneal lobe; penis very long;* extreme tip of tail extends beyond membrane. Color.-Fur brown to grayish brown; slightly more reddish brown below; basal portion of hairs darker; membranes dark brownish black. Teeth.-Upper incisors bifid, small external cusp placed slightly posteriorly; outer incisors nearly as long as inner, exceeding outer cusp of latter in length; second upper premolar separated from canine by slight interval. Measurements of Pipistrellus abrannrus (Temminck). E. H. Taylor collection. No. 342'. male.

mm.

Head and body Tail Height of ear Tragus height Forearm First finger and claw Second finger Third finger: Metacarpal First phalanx Second and third phalanges Founth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx Foot and claws Tibia Calcar

47 31 11

3.7 34.2 6 30

31 12

18 30

9.5 29 8 7 7 12.5

Remarks.-The identification of this form was first made by H. E. Anthony, American Museum of Natural History, according to Dr. C. S. Banks, who presented me with this specimen. The species is apparently not unc()mmon in Manila; there were fourteen specimens in the lot collected by Doctor Banks.

* Said to be proportionally longer than in any other known species. Blanford, loco cit.

300

PHILIPPINE LAND MAMMALS PIPISTRELLUS IMBRICATUS

(Horsfield)

Vespertilio imbricatus HORSFIELD, Zool. Res. in Java (1824). Vespertilio macro tis TEMMINCK, Monog. Mamm. 2 (1827-41) 218. V esperugo imb?'icatus DOBSON, Cat. Asiat. Chirop. (1876) 93 (Java). Pipist?'ellus imbt'icatus THOMAS, Trans. Zool. Soc. London 14 (1898) 385 (Manila); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 17; Proc. U. S. Nat. Mus. 46 (1913) 310 (Manila, Luzon).

Description of the species.-Ear as long as head or slightly longer, broadly rounded off above; basal lobe of inner margin rounded, ascending portion slightly convex; upper third of outer margin flattened, middle third slightly convex, faintly ~marginate opposite base of tragus, and terminating close to angle of mouth in a distinct lobe; tragus peculiarly shaped and characteristic of this species; in general form, regularly crescent shaped, with a large triangular lobule at base of outer margin; inner margin regularly and rather deeply concave, outer margin correspondingly convex, meeting inner margin at an acute point directed inwards; greatest width of tragus slightly below middle of inner margin; glandular prominences between eyes and nostrils well developed, causing a slight hollow on muzzle above; wing membranes attaGh to base of toes; postcalcaneal lobe small, indistinct; tip of tail projecting; toes rather more than half the length of foot, armed with feeble claws; fur of body covers proximal third of humerus and half of femur, its extent on wing membranes limited to a narrow portion along sides of body; beneath, humerus and femur similarly covered, but a wider portion of wing membrane occupied, fur extending as far as a line drawn from elbow to knee joint; upper surface of interfemoral covered as far as end of third caudal vertebra; beneath, fur of abdomen covering root of tail only, remaining portion clothed with short thinly spread hair. Color.-Above dark brown, beneath similar, the fur tipped with ashy color. Teeth.-Upper incisors long, bifid, slanting considerably inwards; outer incisors unicuspidate, acutely pointed, half the length of inner incisors, but less than half their transverse diameter; lower incisors slightly crowded, equal in size, distinctly trifid; second upper premolar very close to canine; first upper premolar very small, its summit scarcely elevated above gum, crushed between canine and second premolar, not visible from without; first lower premolar acutely pointed, equal to threefourths second premolar.

301

PTERYGISTES

Measurements of Pipistrellus imbricatus (Horsfield). mm.

Head and body, female in alcohol Tail Head Ear Tragus Forearm Thumb Second finger Fourth finger Tibia Foot and claws

45.6 38.1 14 15.2 5.58 by 3.04 34.3 7 58.4 44.4' 14 6.3

Remarks.-The type locality is Java. Thomas reports a specimen "picked up in a dying state on the side-walk in Manila." Hollister reports seven specimens from Manila taken by Steere. Gi!nus PTERYGISTES Kaup

Pterygistes KAm', Skizzirte Entw-Gesch. u. N aturl. Syst. d. Europ. Thierw. 1 (1829) 99; MILLER, Ann. & Mag. Nat. Hist. VI 20 (1897) 384; Bull. U. S. Nat. Mus. 57 (1907) 207. Vesperugo KEYSERLINO and BLASIUS, Wiegmann's Archiv. fiir Naturgesch. 5/ 1 1 (1839) 312 (part.). Noctulinia GRAY, Ann. & Mag. Nat. Hist. 10 (1892) 258 (part.); ALLEN, Proc. U. S. Nat. Mus. (1893) 30. Pan~~go KOLEMATI, Algero. deutsch. naJturhisrt., Zeitung Dresden, neue folge 2 (1856) 13l. Vesperugo DOBSON, Cat. Ohirop. Brit. Mus. (1878) 183. Nyctalus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 310.

In general the genus is like Pipistrellus with which it agrees in dental formula. Fifth finger much shortened, scarcely longer than metacarpal of fourth and third; skull with nares extending unusually far back, halfway to interorbital constriction, and with large anterior palatal emargination; teeth as in Pipistr'ellus, but outer upper incisor deeply concave and with a large anterior and small posterior secondary cusp; canine and last upper premolar always strongly in contact; anterior premolar small, quite invisible from outer side; canines without trace of secondary cusps; first and second upper molars with small but rather distinct hypocones; third upper molar, with about half the crown area of first, its metacone and three commissures well developed; lower molars normal in all respects. (After Miller). The principal external character, as mentioned above, is the shortened fifth finger. In the related genus Pipistrellus the fifth

302

PHILIPPINE LAND MAMMALS

finger exceeds the combined length of metacarpal and first phalanx of fourth or third digit. 路 . 2-2 1-1 , 2-2 3-3 Dent a 1 f ormu Ia. 1. 3-3' c. 1-1' pm. 2-2' m. 3-3 = 34. Only the species Pterygistes stenopterus (Dobson) has been reported from the Philippines. PTERYGISTES STENOPTERUS (Dobson)

Vesperugo stenopterus DOBSON, Proc. Zool. Soc. London (1875) 470; Cat. Asiat. Chirop. (1876) 91. Nyctalus stenopterus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 310 (Zamboanga).

Description of the species.-Crown of head slightly elevated, almost level with face line; muzzle broad, flat; labial glandular prominences largely developed, upper lip symmetrically thickened; ears short, triangula:r. ; distance between base of inner margin and termination of outer margin near angle of mouth, directly in a line below eye, equal to distance from base of inner margin to summit of ear; integument forming conch very thick, especially lower half of outer side; tragus short, broad above, narrow opposite base of inner margin, with triangular lobe at base of outer margin; thumb rather long; feet wholly free from wing membrane, which attaches to tibia a short distance above ankle; postcalcaneal lobe small, termination of calcar indistinct; tip of tail free; muzzle in front of eyes, above and below, almost naked; ear conch with short hairs on upper portion; on dorsal surface, base of interfemoral membrane alone covered and below fur on body extends upon interfemoral membrane, along sides of body; antebrachial ~embrane with few hairs; wing membrane covered as far as a line drawn from elbow to knee joint, and a band of hair passing outward to carpus posterior to forearm; legs and feet appear almost naked. Color.-Fur short; above dark brown; below nearly the same; wing membranes dark brown or black. T eeth.-Inner incisors, on each side, long, with a second posteroexternal cusp; outer incisor lying close to hiner, short, without secondary cusp, not equaling height of short cusp of inner incisor; second premolar separated by a short diastema from canine through which first small premolar is visible from without; lower incisors crowded; outer incisors largest; first and second premolars equal in vertical extent, but first nearly twice second in transverse diameter. (After Dobson.)

303

TYLONYCTERIS

Measurements of Pterygistes stenopterus (Dobson). mm.

Head and body Tail Head Ear Tragus' Forearm Thumb Second finger Fourth finger Tibia Foot and claws

58.7 42.5 18.7 15.2 X 10 5.8 X 3.8 38.7 8.1 66 31.9 14 11.4

Remarks.-The type locality is Sarawak, Borneo. Hollister records 'a specimen from Zamboanga, Mindanao. The measurements given here are from Dobson. Hollister (loc. cit.) has placed this species in the genus Nyctalus.* Genus TYLONYCTERIS Peters

Tylonycteris PETERS, Monatsb. Akad. Wiss. Berlin (1872) 703; MILLER, Proc. Acad. Nat. Sci. Philadelphia (1898) 321; Bull. U. S. Nat. Mus. 57 (1907) 212. Vesperus DOBSON, Cat. Chiro-p. Bl' it. Mus. (1878) 184 (subgenus of Vesperugo).

Small bats with entire head greatly broadened and flattened and a conspicuous fleshy pad on ball of thumb and sole of foot; wing perfectly normal; tail well developed, extending to edge of wide interfemoral membrane; tragus well developed, simple;

* Palmer, T. S., Index Generum Mammalium, N. Am. Fauna 23 (January 23, 1904) 464. 'NYCTALUS (subgenus) BowmcR, 1825. Chiroptera, Pteropodidae. Excursions in Madeira and Porto Santo, 36, 1825. TYPE: Nyctalus verrucosus Bowdich, from Madeira. "It forms a new subgenus between pharopus (Pteropus) and cepha-lotes." (Bowdich.) Nyctalus: VIIK'TaXOS (= Jlv~'TaXos), drowsy in allusion to its crepuscular habits. NYCTALUS (subgenus of Vespertilio) LESSON, 1842. Chiro-ptera, Vespertilionidae. Nouv. Tableau Regne Animal, Mamm., 27, 1842. SPECIES, 4: Vespertilio te'T11lTfllinckii Horsfield, from Java; V. belangeri I. Geoffroy, from Po-ndicherry, India; Nycticejus heathii Horsfield, from Madras, India; and N. alecto Gervais, from Manila, Philippine Islands. Name preoccupied by Nyctalus Bowdich, 1825, a genus of Pteropodidae.

304

PHILIPPINE LAND MAMMALS

muzzle with no leaflike growth; ears not connected by membrane and with a well-defined anterior basal lobe on anterior border. Skull broad, depth of brain case through audital bulla barely one-half mastoid breadth; upper surface flat, sloping gradually forward to nares; rostrum short and broad, its length scarcely equal to lachrymal breadth, its depth at front edge of orbit less than half breadth at same region; maxillaries not concealing tooth rows when viewed from above; a distinct blunt proj ection over anterior upper edge of orbit, perhaps representing an incipient postorbital process; teeth n.ormal, without special peculiarities except that upper incisors are conspicuously bicuspidate, crowns longer than broad and nearly as long as high, outer cusp projecting noticeably inward, greatest width of tooth at level of its point; upper canine with well-developed secondary cusp on posterior cutting edge, unconnected with cingulum and extending nearly to middle of main shaft. 1-1 1-1 3-3 . 2- 2 Dental formula: 1. 3-l c. 1-1' pm. 2-2' m. 3-3 = 32. A single wide-ranging species, Tylonycteris pachypus Temminck, is known. It is distributed from Tenasserim throughout the Malay Archipelago to the Philippines. TYLONYCTERIS

P ACHYPUS

(Temminck)

Vespe?,tilio pachypus TEMMINCK, Monog. Mamm. 2 (1841) 217, figs. 4-6 (Java and Sumatra). Vespe?'us pachY1JUS WAGNER, Suppl. Schreb. Saugeth. 5 (1855) 741; DOBSON, Proc. Asiat. Soc. Bengal (1871) 212. Scotophilus fulvidus BLYTH, Joum. Asiat. Soc. Bengal 28 (1859) 293. Tylonycteris pachypus PmERs, Monatsb. Akad. Wiss. Berlin (1872) 704; (southem Luzon); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 17; Proc. U. S. Nat. Mus. 46 (1913) (Luzon, Palawan). ? Tylonycteris meyeni PETERS, Monatsb. Akad. Wiss. Berlin (1872) 705 (southern Luzon). Vespentgo pachypus DOBSON, Cat. Asiat. Chirop. (1876) 115 (India, Tenasserim, Andaman Islands, Sumatra, Java, Philippine Islands).

Description of the species.-Crown of head flat, not raised above broad flattened muzzle; nostrils not projecting, directed forwards and slightly downwards; ears shorter than head, triangular, with rather broadly rounded tips; outer margin straight, terminating towards angle of mouth in a small lobe separated by a shallow notch; tragus short, slightly narrowed upwards and rounded off above, reaching greatest width opposite base of inner margin; at base of outer margin a small triangular lobule;

305

SCOTOPHILUS

undersurface of base of thumbs and soles of feet with broad fleshy pads; in some specimens pads extend along interior surface of thumb almost to base of claw, which is small and acutely pointed; fleshy sole of foot forms almost a circular disk, especially towards toes, under which its round margin proj ects slightly; toes short, not exceeding half the length of foot and armed with short claws; wings rather short; wing membranes attach at base of toes; tail projecting by extreme tip only; calcar short and feeble; fur dense, fine, moderately long, scarcely extending upon membranes. Color.-Generally bright reddish brown above, paler beneath. Teeth.-Upper incisors short, second and shorter outer cusp of inner incisor exceeding slightly in vertical extent the unicuspid outer incisor; lower incisors trifid, not crowded together. Measu?'ements of Tylonycteris pachypus (Te??VIninck). mm.

43.7 32.5 12 5 27.5 5.6 50 35 11.4 6.3

Head and body Tail Ear Tragus Forearm First finger Third finger Fifth finger Tibia Foot and claws

Remarks.-The type locality is given as Java and Sumatra. In the Philippines the species has been reported from Cebu, Luzon, Mindanao, Negros, and Palawan. It is widely distributed, occurring throughout the Malay Archipelago and southern Asia to India. Genus SCOTOPHILUS Leach

Sea top hilus LEACH, Trans. Linn. Soc. London 13 (1821) 69; DOBSON, Cat. Chirop. Brit. Mus. (1878) 256. Paehyotus GRAY, Zool. Misc. 1 (1831) 38 (part); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 219, pIs. 1 and 11, fig. 2 (Miller believes ScotophiZa Hubner, 1816, antedates SeotophiZus). Nyetieejus TElMMINCK, Monogr. Mamm. 2 (1835-41) 148.

Like related genera, externally; skull with no special peculiarities; in largest species a high backward-projecting occipital crest; upper incisors simple, unicuspid, nearly half as high as canines with which they are in contact or nearly so; outer 269453-20

30.6

PHILIPPINE LAND MAMMALS

lower incisors largest; upper premolars without cusps on inner side; canines simple; with cingulum but with no secondary cusps; first and second upper molars with main cusps close together and so displaced outward that protocone of first lower molar is near middle of crown; mesostyle particularly reduced and sometimes absent; third upper molar with protocone, paracone, parastyle, and two commissures of which the second is short; lower molars with hypoconid and entoconid reduced in size, so that second triangle in each tooth is smaller than first and with cusps noticeably lower; in third lower molar second triangle greatly reduc~d. (After Miller.) . 1-1 1-1 1-1 3-3 Dental formula: 1. 3-3' c. 1-1' pm. 2-2' m. 3-3 =30. Ten or more species are known, of which only Scotophilus temminckii (Horsfield) has been recorded from the Philippines. SCOTOPHILUS TEMMINCKII (Horsfield)

VespertiUo te'l'/UYYl,inckii HORSFIELD, Zooi. Res. in Java (1824). Vespertilio belangeri 1. GEOFFROY, Belang. Voy. aux Indes Orient (1834) 87 and 92, pl. 3.

Vespertilio noctulinus 1. GEOFFROY, Belang. Voy. aux Indes Orient (1834) 92, pI. 3. Scotophilus temminckii GRAY, Mag. Zooi. and Bot. 2 (1838) 497; PETERS, Monatsb. Akad. Wiss. Bedin (1866) 679; DOBSON, Proc. Zooi. Soc. London (1875) 370; Cat. Asiat. Chirop. (1876) 120 (figs. a and b); Cat. Chirop. Brit. Mus. (1878) 25& (Negros); HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 17; Proc. U. S. Nat. Mus. 46 (1913) 310 (Luzon, Catanduanes, Ticao, Sibuyan, Panay, Guimaras, Negros, Mindanao, and Palawan). Nyctdcejus temminckii TEMMINCK, Monogr. Mammal 2 (183&-41) 149; WAGNER, Suppl. Schreb. Siiugeth. 1 (1841) 541 and 5 (1855) 764; CANTOR, Journ. Asiat. Soc. Bengal 15 (1846) 185; BLYTH, Journ. Asiat. Soc. Bengal (185f) 157; HORSFIELD, Cat. Mamm. Mus. East Ind. Go. (1851) 37; BLYTH, Cat. Mamm. Mus. Asiat. Soc. Bengal (1863) 路 31. Nycticejus luteus BLYTH, Journ. Asiat. Soc. Bengal 22 (1851) 157; 21 (1852) 346; WAGNER, Suppl. Schreb. Siiugeth. 5 (1855) 765. Nycticejus flaveolu8 BLYTH, Horsfield, Catt. Mamm. Mus. East Ind. Co. (1851) 37.

Description of the species.-(From No. 325, E. H. Taylor collection; collected at Manila, P. I., April, 1922, by E. H. Taylor.) Muzzle thick, obtusely conical; head slightly raised above muzzle line; glandular prominences, between eyes and nose, small,

SCOTOPHILUS

307:

not causing a depression on muzzle above; ears short, narrow, and rounded at tips; inner basal lobe somewhat rounded; horizontal part of margin straight, forming a right angle with ascending portion; lower half of ascending part of inner margin straight or even faintly concave, upper half convex, tip not projecting outwards; from summit of ear, outer margin slanting slightly outwards and downwards for a short distance, then straight and slightly deflected outwards as far as a point opposite base of tragus, where it is deeply emarginate, and beyond which it terminates in a short convex lobe; tragus elongate, narrowed toward tip, point rounded; it curves forwards and inwards; lower third of outer margin straight, with a distinct small horizontal lobule at base, upper two-thirds convex; inner margin concave; from base of inner margin a narrow prominent ridge passes outward and slightly upward across anterior surface of tragus to outer margin; wing membranes attach to side of foot near base of toes; postcalcaneal lobe narrow; feet moderately large, toes about as long as metatarsals; penis large, distal portion much widened ; extremity of tail projecting; fur above 'almost wholly confined to body, scarcely extending in any direction upon wing membranes; wing membranes beneath, covered. with fine hairs as far as a line drawn from elbow to knee joint; interfemoral membrane naked, except a few scattered hairs at base of tail; fur above elongate, practically unicolored. Calor.-Dresden brown (Ridgway) above, individual hairs slightly lighter near base; below similar, but distinctly lighter brown; wing membranes much darker brown than fur; underside of tail membrane lighter than other portions of membranes. Skull and teeth.-Skull with brain case raised but little above level of rostrum, with prominent crests; occipital and sagittal crests forming at their junction a thick projecting process from which the skull slopes evenly downward and forward to end of nasal bones in front; occiput concave with prominent occipital crest; facial bones shortened in front of infraorbital foramina which are large and well defined; bony palate narrow behind last molar, extending backwards as far as middle of zygomatic arches; a small median projection on posterior edge of palate; basioccipital between cochlere broad; paroccipital and mastoid processes well developed, prominent.

308

PHILIPPINE LAND MAMMALS

Measu'rements of Scotophilus temminckii (Horsfield). mm.

Head and body Head Tail Ear above notch Tragus Forearm Thumb Second finger Metacarpal Third finger Metacarp.al Fourth finger Metacarpal Fifth finger Metacarpal Tibia Foot and claw Skull: Greatest length Zygomatic width Depth of brain case and bullre Palate from nO(tch Upper tooth roW' Lower tooth row Mandible

74 24 39 13.5 7.5 5004'

9 53 48 85 50 72 49 62 45 22 11

19.8 13 10 7.1 7.7 8

Remarks.-This species, which ranges from Afghanistan to the Philippines, is variable in size and color. Philippine specimens have smaller ears, a shorter tail, and a decidedly shorter forearm than the specimen whose measurements are given by Dobson. The species is widespread in the Philippines. I have numerous specimens in my collection taken in Manila. Specimens were captured in a media agua on a house in Manila, where there was a colony numbering several hundred individuals. The variation in color is rather great. Most of the specimens observed are either dark Dresden brown or reddish cnestnut-buff. Apparently, the color does not depend upon age or sex, as the two colors are present in both young and old individuals and in both sexes. Specimens kept in alcohol become greatly faded in a few years. In specimens obtained at Iwahig Penal Colony, Palawan, the teeth are slightly smaller. This is noticeable in the thickness of the incisors and canines, and the actual length of the molars in the lower jaw. The posterior portion of the mandible is larger.

MOLOSSIDÂŁ

309

The type locality is Java. In the Philippines the species is known from Luzon, Negros, Panay, Palawan, Catanduanes. Ticao, Sibuyan, and Mindanao. This species is probably the one most commonly seen about Manila at twilight. Family MOLOSSIDJE Molossi PETERS, Monatsb. Akad. Wiss. Berlin (1865) 258; FITZINGER, Sitz-Ber. Akad. Wiss. Wien. 61 (1870) 458. Molossidre GILL, Arrangement of the Families of Mammals, Smithsonian Misc. CoIl. (1872) 17; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 241, pI. 14, figs. 1-5. GymnU1'idre AMEGHINO, Actas de la Acad. Nac. Ciencias. Argentine and Cordoba (1889) 351.

Most authors have treated this group as a division of either the Vespertilionidre or the Emballonuridre. Humerus with trochiter much larger than trochin, the difference in size more noticeable than in the Vespertilionidre; trochin articulating with scapula by a surface nearly as large as glenoid fossa; epitrochlea short but with very conspicuous spinous process; capitellum almost directly in line with nearly straight shaft; ulna less reduced than in the Vespertilionidre, the very slender shaft about half as long as radius; second finger with well-developed metacarpal and one rudimentary phalanx; third finger with three phalanges, of which the first is flexed on upper side of metacarpal when wing is at rest, and third is cartilaginous except occasionally at extreme base where a distinct joint is formed with middle phalanx; fifth finger scarcely longer than metacarpal of first; shoulder girdle normal except that seventh cervical vertebra is fused with first dorsal; foot short and broad but of normal structure; fibula complete, bowed outward from tibia, its diameter about half that of latter, entering conspicuously into mechanical scheme of the short, stout leg; pelvis normal, boundaries of sacral vertebrre clearly defined; lumbar vertebI'ire not anchylosed; skull without postorbital processes; premaxillaries with or without nasal branches, when present forming two palatal foramina; when absent allowing formation of one; posterior orifice of anteorbital canal not enlarged; teeth normal; ears variable in form and size, sometimes joined across forehead, tragus much reduced, antitragus usually very large, anterior border of auricle without basal lobe; muzzle obliquely truncate, usually sprinkled with short, modified hairs with spoon-shaped tips, nostrils usually opening on a special pad, upper surface of pad often set with

310

PHILIPP]NE LAND MAMMALS

fine horny excrescences; wing narrow; fifth finger much short,e ned; 'membranes thick and .leathery; uropatagium short, tail projecting conspicuously beyond its free edge. (After Miller.) About fifteen genera are recognized .as belonging to the family, containing approximately ninety species. These are distributed over the warmer parts of both hemispheres, but probably absent from South America. The -genera Philippinopter'US, Cheiromeles, and Chrerephon are known from the Philippines. The records here given for the first two genera are the first for the Philippines. K ey to the Philippine genera of Mol08Sid83. Ears joined together anteriorly; body heavily furred; lips wrinkled; no neck gland; smaller. b 1 . Two upper premolars ___ ____ ___ ___ __ ____ ____ __._. Ch83rephon Dobson (p. 310). b 2. One upper premolar _______ .____. __.__ ____ __ Philippinopterus g. nov. (p. 3>14). a'. Ears free; body nearly naked; lips smoOlth; neck gland present; larger. Cheiromeles HOl'sfield (p. 320). a

Genus CHlEREPHON Dobson

Ch83rephon DOBSON, Journ. Asiatic Soc. Bengal 43 pt. 2 (1874) 144; .ANDERSON, Ann. Mus. Civ. de Stor. Nat. Genova 3 3 (1907) 35; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 244; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 20. Nyctinomus DOBSON, Cat. Chirop. Brit. Mus. (1878) 420 (part.).

Ears connected by a low band across crown; tragus very small ; tail long, free from interfemoral membrane for about half its length; upper lip thick, deeply grooved by vertical wrinkles; wing membrane attaches to lower end of tibia; fur relatively short and dense. Teeth and sk7ull.-Upper incisors simple, well developed, about half as high as canines, far apart at base, strongly converging at tip, separated from canine by a space equal to their greatest diameter; lower incisors equal, bifid, their crowns in contact with each other; two outer in contact with canines; canines strong and well developed, cingulum distinct but not forming a true secondary cusp except occasionally at base of tooth; first upper premolars minute but in tooth row; second upper premolar not touching canine, well developed, with a large anterointernal cusp; molars normal, first and second with well-developed hypocone. Skull with rounded or somewhat flattened, moderately wide brain case about one and one-third times as long as rostrum; premaxillary bones complete, their palatal branches isolating

311

CILEREPHON

two small palatal foramina; when closing of emargination is complete a slight anterior median notch is sometimes developed, but this never extends behind incisors . . 1-1 1-1 2-2, 3-3 Dental formula: 1. 2- 2' c. 1-1' pm. 2-2 m. 3- 3 = 30. Ch;;erephon and Nyctinomus resemble each other closely in external characters, but differ in the structure of the palate. In Ch;;erephon the maxillaries are completely developed. The genus includes all species, formerly associated with Nyctinomus, without obvious palatal emargination. From Philippinopterus it differs in havipg two upper premolars instead of one. CHlEREPHON LUZONUS HoUister

Chawephon luzonus HOLLISTER, Proc. U. S. Nat. Mus. ? Ch83rephon plicatu8 (auct.).

(1913) 312.

Description of the, species.-Head not noticeably flattened; eye small, well back of anterior attachment of ear; nostril at tip of snout; snout obliquely truncate; nostrils directed forward, somewhat outward and downward; distance between nostrils equal to their distance from mowth; a serrated keel forms the upper edge of nostrils, behind which is a distinct groove; below, between nostrils is a slight keel, beneath which is a transverse groove that practically surrounds nose; upper lips with elongate glandular plicre or wrinkles which overlap lower jaw; lower jaw short, opening on ventral side of head; chin with broad glandular areas bordering lips, separated by a median groove, extending to a rounded median papilla; lips and chin covered with very short bristly hair rising from rounded papillre ; forward tip of "nose," around and between nostrils, naked; snout with numerous short spoon-tipped modified hairs; ears extraordinarily complicated, brought forward and joined together at a point nearer tip of muzzle than eye; extending outward on either side in a plane with top of head, forward part of ear somewhat inflated; anterior margin of ear extending outward from connecting membrane, forming a lobe directly anterior to antitragus; latter forming a strong rounded free lobe; posterior (or upper) margin of ear folded back and wrinkled, rounded in general outline; a few transverse ridges well defined in outer part of ear, and an indication of a slight lobe rising from inner surface of ear near base, set off sharply from antitragus by a deep groove; tragus very small, directed upward, upper part forming almost a square, top sharply trun-

312

PHILIPPINE LAND MAMMALS

cate; tragus connected with antitragus by a low membrane; a deep groove below eye; eye almost concealed by anterior flap of ear; laid forward, ear extends slightly beyond tip of muzzle; wing membrane not ample, membrane attaching near middle of tibia; fifth finger short, metacarpal about half as long as third; second finger with phalanx scarcely discernible; narrow membrane between second and third finger scarcely extending beyond. distal end of metacarpal of third; phalanges of third and fourth fingers fold,. first in, then out, and the outer phalanx turns up; thumb short, with a distinct rounded callous at base; antebrachial membrane of wing reaches base of thumb as a very narrow margin; interfemoral membrane "much restricted; calcar short; no postcalcaneal lobe; tail about as long as hind limb, inclosed in interfemoral membrane for half its length; feet of moderate size, two outer toes slightly shorter, much thickened by a lateFoinferior glandular area thickly covered with hairs of varying lengths; median toes with very long hairs only at base of claws; longest hairs curved abruptly at end in many specimens; outer toe somewhat opposed to other toes. Fur on head extends forward on ears to tip of membrane connecting ears; less than one-half of outer surface of ears covered with fur; wing membrane unfurred above and below, except on parts adjacent to body; very short minute hairs cover entire surface above and below, longer on basal portion, sparse and shorter on distal portion. Color.-Fur brown above, hairs much lighter near base, fur on top of head darker; hair below, on neck, buff, tipped with gray, on belly gray only slightly lighter at base; wing membranes lavender brown. Teeth and skuU.-Upper incisors caniniform, separated widely at base, converging toward tips but never touching, separated from canines by a distinct diastema; a slightly developed cingulum; lower incisors forming a continuous semicircular row, touching each other as well as canines, strongly bifid, not reaching as high as cingulum of canines; cingulum of upper canine low; that of lower canine much widened, forming a small anterointernal cusp and a slight posterointernal cusp; lower canines rather narrowly separated; first upper premoJar small, directly in tooth row, not crowded; skull rather flattened above with a low sagittal crest; a distinct supraorbital ridge terminating anteriorly in a distinct groove on rostrum between supraorbital regions.

313

CHlEREPHON

Measurements of Chll3rephon luzonus Hollister. [Measurements in millimeters.] E. H . Taylor collection No. and sex. 514 ,

female.

481 ,

556

481,

479,

female.

' male.

female.

8 .5 39

99 62 37 18 15 3 .8 9. 5 42 8.5 39

97 63 34 18 15 3.8 9.2 44 8.5 42

98 60 38 17" 15 3.8 9.5 42 9 41

Third finger: MetacarpaL ___________ [ First phalanx _ _ _______ _ Second phalanx_ ___ ___ _ Third phalanx .. ____ .__ _

41 16 .5 15. 5 7

40 . 5 17 16". 5 6.5

Fourth finger: MetacarpaL_ __ ____ ____ First phalanx- _________ Second phalanx- __ ___ __ Fifth finger . __ _____________ Tibia _______ ______________ Foot. ________ ____ ________ _

38 13 10 40 15 11

39 12 . 5 10 39 15 10.5

18 16.2 10.4 10 . 1 ____________

Totallength ___ __ ______ __ __ 99 Head and body_ .. ______ __ __ 61 TaiL. _________ _____ ______ _ 38 Ear, base to anterior point . . _ Height of tragus . . _________ _ Height of antitragus . _ _ _____ Eye to tip of snouL_ _______ Forearm .. __ _______________ [ First finger with claw _______ Second fingeL _______ __ ____

Skull: Greatest length ... ______ Condylobasal length ____ Zygomatic breadth_ _ _ __ Breadth of brain case _ __ Interorbital constriction . Upper tooth row, including canine ____ ___ Lower tooth row, entire_ Mandible. _____ ________ Upper canine, height . . _-

17 18

4 8.5

16 10.2 10 4

580 ,

536, fe-

Type United States 360, Nationfeal Mumale. seum.

male.

95 60 35 16 15 4 9 42 .5 9 39

96 62

95 63

18 15 3.8 8.5 41 8 .5 39

17 18 4.1 8.0 44 9.5 39

41 41 17 16 .5 17 1 16. 5 7 .5 6

41 18 17 6

40 17 15

18 17 7

40 14 11 41 16 1J.

40 14 10 41 15 10

38.5 13.5

39 13

16 . 8 15 10 . 1 10

18 16 10.5 10 4

18 16 . 5 10 . 2 10 4

7 7 .2

7.5

39 13 10.5 39 15 10

4.1

39 15

40 16

7.2 12 . 1

12 . 3

3 .1

94 59 35 17 15 4.3 8 .2 42 9.6 39

54 33

44.4

41 17 16 7 38.9 12 . 2 9 39 15 11 17 . 6 16 . 2 10 . 2 3.1

6.5 6J 7

1~. 8 1 __ ~~~~ __

Variation.-More than one hundred specimens of this species were collected. A large colony inhabits one of the Montalban caves near Manila where many were taken. Specimens were obtained in a small cave in the cliff wall immediately below Pagsanjan Falls, near the town of the same name, south of Manila. The amount of variation is negligible. The body length varies only a few millimeters. A few individuals are slightly more reddish brown than others. The groove on the chin is often indefinite, and the papilla on the chin is frequently inconspicuous. The numbers listed here are all from Montalban caves.

314

PHILIPPINE LAND MAMMALS

Remarks.-The species appears to be related to Ch;;erephon plicatus, from which it differs chiefly in being smaller. Total length, 94 to 100 millimeters (C. plioolus, 116 to 122) ; forearm, 42 to 44 (C. plicatus, 48 to 50). The alar expanse is less, and the tragus is shorter, the top not concave. The color apparently is distinctly different. Dr. Paul Bartsch visited the Montalban caves where most of my specimens, over one hundred, were collected, and obtained about sixty specimens of Chiroptera distributed among seven species, but not one specimen of this species was taken. On my visit nearly three hundred bats, distributed among ten species, were obtained. More specimens of Ch;;erephon were taken than of any other species. It does not seem reasonable that the species was not present at the time of Doctor Bartsch's visit, and yet it is remarkable that no specimens were obtained by him. Genus PHILIPPINOPTERUS novum

Type.-Philippinopterus lanei sp. nov. Similar to Nyctinomus and Ch;;erephon in general external characters. It agrees with Ch;;erephon in the complete condition of the premaxillary bones, and in the presence of two palatal foramina. The teeth are similar in general to those of Ch;;erephon but the first upper premolar is absent in the maxilla . . 1-1 1-1 1-1 3-3 Dental formula: 1. 2-2' c. 1-1' pm. 2-2' m. 3-3=28. The first lower premolar is similar to the same tooth in Ch;;erephon but is reduced to about one-half the size. Mormopterus Peters has the same dental formula as Philippinopterus, but the character and size of the ears of the former genus as well as many other generic differences seem to show that no close relationship exists between the two genera. PHILIPPINOPT'E RUS LANEI sp. nov.

Type.-No. 253, E. H. Taylor collection; collected April 28, 1923, at Saub, Cotabato, Mindanao, by E. H. Taylor. Alcoholic .specimen; skull separate. Diagnosis,. description of the type.-Adult female. In general conformation resembling markedly Ch;;erephon luzonus Hollister; body somewhat heavier and head broader; ears large, similar to those of C. luzonus, but somewhat thicker. The snout resembles the snout of a swine; a broad V-shaped notch above; nostrils directed forward and ,o utward; end of snout very

315

PHILIPPINOPTERUS

obliqnely truncate; upper lips thickened, vertically plicate or wrinkled, overlapping lower lips; underside of snout with numerous short modified hairs with spoon-shaped tips; posterior to these hairs a roughly circular area of hair; modified hairs with spoon-shaped tips scattered on upper lips and upper sur120

118

PHILIPPINE ISll\NDS

126

124

122 Batan Is

® Chc:er>ephon luzonus @ Philippinoptepus lanei

@ Aonyx ciner>ea

Lubsng'<::;,:

~ ""'o

MINDORO

@ Busuanga ~'®.

DCatanduanes S1uque

"}>

8u.;a~ .,•• f7bllomblon~~, ;

"--V :;;S>yan

,cao

•

l~~:~·t:l~MA.~~:;'u

10)r'

@ Cas"yan Is J'NEGROS

PAL AWAN

MINDANA

8- -. alabac 6

SOHOL .0

<;)

QCagayan Sulu

120

FIG. 16.

122

126

Distribution of Molossidre, Felidre. and Mustelidre in the Philippines.

face of snout; a few scattered elongate vibrissre on snout; eye small, nearly hidden by ears; latter extend out from sides of head, usually held on a level with top of head; outer lobe with several wrinkles; a band of reddish fur around anterior edge of conch; balance of ear almost entirely naked, except upper

316

PHILIPPINE LAND MAMMALS

inner part; tragus small, lobelike, rounded slightly on top, sides nearly equally concave, separated by a slight notch from a small conical elevation behind it; antitragus large, rounded, separated from posterior part of ear by a deep notch, and attached anteriorly at a point only a short distance posterior to angle of mouth; ears joined together anteriorly by a moderately high, skin band; neck strongly constricted; parietal region broad; posterior to ear band a longitudinally plicate, totally naked, glandular area, surrounded by long fur; lower lip only slightly wrinkled with a rounded glandular papilla on under side of jaw bearing two or three elongate hairs. Wing membranes attach on sides of body; wings slender, elongate, folding at tip, similar to Chmrephon; first digit with a rounded callous spot at base; posterior lateral wing membrane attaches to tibia about one-third of total length from knee; interfemoral membrane attaches about midway on tibia; calcar moderately well . developed; tail long, more thickened than in Chmrephon; it emerges from posterior edge of membrane and is free for about two-thirds its length; first and fifth toes with a glandular area on their outer sraes, the areas bearing numerous fine hairs and a few elongate hairs; tiI?s of other toes with elongate hairs; glandular areas on skin above femora, each area bearing a few elongate white or reddish hairs; body above and below covered with dense fur; that on ventral part shorter than dorsal; wing membranes naked except for a slight growth on membrane anterior to arm. Color in life.-Fur above dull reddish brown to orange brown, basal portion of fur buff. Below much more buff-brown, buff predominating in region of neck; ear cartilages, snout, and wings, blackish. Skull and teeth.-Dental formula given under the genus. General shape of skull similar to that of Chmrephon ~uzonus. Sagittal crest extending to nasals; brain case not especially inflated; palate complete, with two diagonal, oval, palatal foramina not completely separated. Canines with a broad anterior groove, inner edges much compressed forming a sharp edge; main axis of tooth curves out somewhat; cingulum not pronounced, posterior edge does not form a cusp; single premolar in contact with canine; two upper incisors conical, placed slightly apart, their tips not curving; lower incisors crowded in a curve between canines, all four strongly bilobed, failing by an appreciable distance to reach top of cingUlum on inner side of canines; lower canine triangular with a broad cingulum, anterior facet

PHILIPPINOPTERUS

317

with a slight ridge, inner facet smooth, posterior with a deep groove; first lower premolar decidedly smaller than second, overlapping cingulum of canine. Variation.-A young specimen (No. 299, head and body, 50 millimeters), taken at the same time as the adults, has the skin almost naked, and heavily wrinkled above. On the throat and neck there is a noticeable growth of fur, becoming scant on the breast. The abdomen is practically naked. The glandular areas on the femora are distinctly marked, each bearing several long vibrissalike hairs, some black, some white. The glandular area between the ears is marked by an elongate area of lightcolored skin; the lower jaw is covered largely with whitish skin, which extends behind the angle of the mouth. The median gular papilla is well differentiated. In a somewhat older specimen (No. 287, head and body 58 millimeters) the femoral glands are more strongly Plionounced, and the naked area between the ears is more conspicuous. The entire neck, breast, and abdomen are covered with short grayish fur, much darker, almost black on the breast and sides of head and neck. I have examined the milk dentition of the two specimens and find that in the upper jaw of each there are six slender, straight, terete teeth tapering to a sharp point and recurved at the tips ; the two corresponding to the upper incisors curve outward; the second pair also curve outward and forward somewhat and are distinctly smaller than the first pair; the next tooth on either side (the canines?) are curved backward and inward. I am unable to discern other milk teeth in the upper jaw. In the larger specimen the same teeth are persistent even though the permanent teeth are coming through the gums. In the lower jaw' of the younger specimen there are likewise six teeth, two incisors and a canine on each side. The inner incisor is bilobed as is the adult tooth and is widely separated from its fellow; between them is a pair of teeth which I presume to be the permanent incisors; the second milk incisor is a very small straight tooth; the third tooth is large and strongly recurved at the tip. The sex differences are strongly marked in the character of the teeth. The canines of males are decidedly larger and longer, their tips more widely separated; the upper incisors are larger and stand somewhat closer together; the molars likewise are larger. In the mandibles the canines stand closer together and the cingula are broader, while the lower incisors fail to reach so high a position with regard to the cingula of the canines.

37 14 10

15.2 6

18.~

232 --105 73 32 21 1 4 10 39 9 38

Fifth finger: Metacarpal _______________________________ 22 First phalanx _____________________________ -------Second phalanx ___________________________ -------Hind foot ... _. ______ .• ________________________ 12.8

Fourth finger: Metacarpal _______________________________ First phalanx _______________________ - _____ Second phalanx ___________________________

Third finger: MetacarpaL ___________________ - __ . _______ First phalanx _____________________________ Second phalanx _________________________ . _ Third phalanx .•. __________________________

Ear, from base forward .. _______________________ Tragus. ______________________________________ Antitragus .. __________________________________ Eye to end of snout .. __________________________ Forearm .. ____________________________________ First finger and claw.. _________________________ Secondfinger _________________________________

Totallength__________________________________ Head and body.:. ___________ ~ _________________ Tail ... _______________________________________

250

28 11

6 12.2

6 12.8

38 14 10

40 18 15 8

100 68 32 21 1 3 10 41 9 38

39 15 10

43 18 16 6

41 10 38

109 83 36 22 1 4

253

760

22.6 10 5

38 14 10

38 18 16 7.6

99 68 31 20 1 3.2 10 39 8.8 37

- - - - - - - --

Males.

245

271

281

290

Females.

11.8

22 10

38 13 9

40 17.5 14 7

98 66 32 21 1 3.6 10 4.6 8.8 38

6 11

28 10.3

38 13 9

40 18 14.8 6

96 65 31 20 1 4 10. 42 8 38

11 6

11 11

37 13 10

40 16.8 13 7

97 66 32 20 1 3.6 9 41 8 38

28 10.6 6

37 13

40 16.5 14 6.5

98 65 33 19 1 3.6 10 41 8 38

22 10 6 11

37 13 8.2

40 17.6 14 7

96 64 31 20 1 3.5 9.3 40 8 38

- - - - - - - - - - - - - - - --

273

E. H. T.,I" ooUoo"•• N..

[Measurements in millimeters.]

Measurements of Philippinopterus lanei sp. nov.

297

307

761

11.2

22 9 6

37 13 8

40 17.6 14 7

93 63 30 18 1 3.5 9.3 40 8 38

21 9 6 12

36 18 7

39 16.5

90 60 30 19 1 3.4 9 39.6 8 37.5

22 9.8 5 12

37 13 7.8

40 17 14 7.1

92 60 32 19 1 3.2 9.2 42 9 40

--- --- ---

r:n

> t"'

~ ~

t;j

t"'

Z t.:I:j

'"d '"d .....

~ ..... t"' .....

'"d

CJ.:) ......

Condylobasallength ___ __ ____ ______ __ ____ __ Zygomatic width ____ _____ ___ ____ ____ ___ __ _ Orbital constriction _________ _______ ____ ___ Upper molars ___ ___ ___ _____ __ __ ___________ Width between canines, basaL ______ ________ Width between canines, distal. . ___ _______ ___ Height of upper canines __________ _____ __ ___ Width between third molars ________________ Mandible length . . _____________ _______ _____ Length of lower molars. __________ __________ Height of lower canines ___________ ___ __ ____

Skull: Total length ____ ____ ___ __________ ___ ___ ___

20 17 12.6 5 5.2 2.3 5. 2 3 51 13.8 6.8 2.5

3 5.3 14 6.7 2.2

20 17 12.8 5 5.2 3 20 17 12 5 5.5 2 4.4 3 5 14 6.8

2. 8

20.2 17.2 12.3 5.1 5.7 2 4.9 3 5 14 . 8 6.8

20 17 12 5 5.5 2.5 5.3 3.3 5 14 7 2.2 18 16 11.5 4.2 5.3 2 4 2.8 4.8 18 6.7 1.8 4.8 5.5 2 4 2.8 4.8 18 6.8 1.9

19 16

19 19 15.8 15 . 8 12 - ----- -4.8 4.8 5.3 5.3 2 2.2 4.2 4.3 2.6 2 .8 4.9 5 13 13 6 .7 6. 5 1.9 2 4.9 5.3 2.1 4 2.6 5 13 6.6 2

----_._- -

19 15.7

19 15.5 11.5 5 5.5 2.2 4.2 2.6 5 13.5 6.7 2

18 16 11.5 4.2 5.3 2 4 2.8 4.8 18 6.7 1.8

CI:l

rJl

'"t1

~ ..... Z o

t"' .....

'"t1

III .....

320

PHILIPPINE LAND MAMMALS

The heads of the males are more heavily muscled and are much broader than those of the females. The growth of the fur on the posterior side of the earband is denser in the females than in the males. The males have larger average measurements than the females. RemarkS.-About forty specimens were captured at Saub, Cotabato, in a recently felled tree; which represented only a part of the colony. The species is named for Dr. H. H. Lane, professor of zoology, University of Kansas. Genus CHEIROMELES Horsfield

Cheiromeles HORSFIELD, Zool. Res. in Java (1825); DOBSON, Cat. Chirop. Brit. Mus. (1878) 405; MILLER, Bull. U. S. Nat. Mus. 4 (1901) 249.

Chiroptes GLOGER, Gernein. Hand-u. Hilfsbuch der Naturgesch. (1841) 49.

Chiromeles AGASSIZ, Nornenclator Zo'ol., Marnm., Addenda

(1846) 3.

Body robust and heavy; ears narrow, extending at right angles to body, widely separate, rather angulate; tragus low; lips not folded and wrinkled; limbs short and heavy, toes independent of each other; membranes attached near middle of back; space between membranes and body forming a spacious pouch by a second membrane from humerus; mammre in the pouches; skin greatly thickened and wrinkled; a large gland on neck covered by a large neck fold; numerous small openings of the gland in males; in females a single large orifice; body with fur scant and so short as to make body appear almost naked. Teeth and skull.-Upper incisors short, robust, their form much as in Molossus; height of crown barely equal to width through posterior expansion, shafts oblique and closely in contact with each other; a distinct space between incisor and canine; lower incisor well developed, touching canines; canines heavy, but not peculiar in form, the upper with a deep longitudinal furrow on anterior face, and a minute posterobasal cusp; upper premolar large; first lower premolar scarcely functional; skull robust and heavily built, rather smooth, sagit~al crest barely indicated except in posterior part of skull. The three species of the genus are Cheiromeles torquatus Horsfield, a widespread species occurring from India to the Philippines; C. parvidens Miller, of Celebes; and C. jacobsoni Thomas, of Simular. 'T he latter two species are somewhat smaller than the first. This is probably the first authentic record for the occurrence of a species of the genus in the Philippines.

CHEIROMELES

321

CHE.lROMELES TORQUATUS H(>rsfield

Chei1'omeles torquatus HORSFIELD, Zool. Res. in Java (1824); DoB.SON, Monog. Asiatic Chiropt. (1876) 177; Proc. Zool. Soc. London (1876) 704; Cat. Chirop. Brit. Mus. (1878) 405, pI. 21, figs. 1 and 1a; MEYER, Abhandl. Mus. Dresden 7 (1899) 48 pIs. 10, 11; TROUESSART, Cat. Mamm. 1 (1898-1899) 142; Sup. 1 (1904-1905) 101; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 249. Chiropeles torquatus GLOGElR, Gemeinnutziges Hand-u. Hilfsbuch der 'Naturgesch. 1 (1841) 49. Chiromeles tOTquatus AGASSIZ, Nomenc. ZooI., Mamm., Addenda (1846) 3. Dysopes chei'ropus TEMMINCK, Monog. Mamm. 1 (1827) 218, pI. 17. Dysopes cau,datus TEMMINCK, Monog. Mamm. 2 (1827) 348, pI. 71.

Description of the species.-(From No. 390, E. H. Taylor collection; collected at Saub, Cotabato, Mindanao, April 25, 1923, by E. H. Taylor.) Head broad and flattened; snout truncate, extending'considerably beyond lower lip; nostrils directed forward and outward; upper lip thickened, not or but slightly wrinkled; underside of snout with a triangular area covered with extremely short, spiny, modified hairs with spoon-shaped tips; a few similar spines on side and top of head; eye moderately large, for the most part posterior to anterior point of attachment of ears; ears widely separated, of simple shape; outer side of ear with seven well-defined ridges; antitragus rounded, much thickened; tragus low, rounded; lower lip with a few spiny hairs with spoon-shaped tips; head distinctly delimited from body by a heavy fold of skin which encircles neck, suggesting that head is somewhat retractile (turtle fashion); skin fold covers or, at least, partially covers a large neck gland (in females gland with a single large opening; in males gland opens through two series of small rounded pores); muzzle without hairs or spines; body heavy and robust; legs short and thick; calcar apparently wanting; toes independent, inner toe much thickened, glandular, the whole inner side covered with long reddish hair 6 to 8 millimeters in length; other toes with a tuft of hair about base of claws; tail short, thick, extending beyond interfemoral membrane for more than two-thirds of its length; interfemoral membrane thickened, attaching some distance above foot; wing membranes thick and leathery next to body; membranes between fingers thin, typically batlike; wing expanse small in comparison with size and weight of body; wing membranes fastened to near middle of back, the space between wing and body forming a pouch by means of a second membrane fastened to side of body and attached to humerus, the pouch 259453-21

322

PHILIPPINE LAND MAMMALS

opening posteriorly; first finger with a well-developed claw and with a smooth rounded pad near base; second finger with long metacarpal and a rudimentary phalanx; third and fourth fingers with three phalanges; fifth finger scarcely longer than metacarpal of second; body above naked except for a thin covering of hair 0.2 to '0.4 millimeter in length; a few scattered longer hairs on interfemoral membrane and tail; wing membranes entirely naked; a few elongate hairs on snout mostly directed forward; head and upper anterior part of ear with a thicker and somewhat longer growth of hair than body; a ruff of hair behind ears and on neck 5 to 10 millimeters long; breast covered with a denser growth of hair nearly 1 millimeter long. Color.-General color, due to color of skin, blackish; hair on body distinctly brownish to brownish red. Skull and teeth.-See under genus. M easurements of Cheiromeles torqu atus H01路sfield. [Measurements in millimeters.] -

___ _t ___

Total length .. _______ ___________ L ~ L Head and body _____ ____ ____ ___ ___ ___ ___ __ __ __ Tail ___ ___ ____ ____ ___ __ ____ ___ _________ ___ ___ Length of ear .. __ __ _________ ____ ___ ___ ____ ____ Height of antitragus. ____ ____ __ ________________ Eye to end of snouL ____ ____ ________ ___ ___ __ __ Ear. basal measurement ... _____ __ ________ __ ___ _ Length of forearm . . ___ __________ ______________ First finger without claw. ___ ___ ______ _____ ____ _ Secondfinger . . ________ ______ _____ ____ _____ ___ Third finger; Metacarpal. . ____ ____ ____ ______ __ _________ First phalanx . _____ __ _______ ____ _____ _____ Secondphalanx. ____ ____ _______ " __ ____ ____ Fourth finger, metacarpaL _______ ____ ____ ___ ___ Fifth finger, metacarpaL ___________ __ ________ _ Tibia. ____ ________ ___ __________ ____ __ ________ Skull: Condylobasal length . . ______ ___ __ ___ ___ __ __ Zygomatic width .. _____ __ _____ __ ______ ____ Narrowest constriction . . __ _______ ___ __ _____ Greatest height of skull at crest. _______ ____ Upper tooth row, including canine ... ________ Distance between canines __________________ DiBtan~ between third molars .. ____________ Lower tooth row, including canine .. _________ Length of lower jaw. ______________________ Height at coronoid process _________________

E. H. Taylor collection No. and

s=-l

387,

388,

389,

390,

female.

male.

female.

180 126 54 19.5 5.3 17.9 17.5 7.5 15.5 70

170 116 54 18.8 4.5 16.5 16.8 7.3 15 66

185 127 58 20 5.8 18.5 18 7.8 17 72

80 47 38.5 75 37 31

75 43 34 72 36 30

81 48 39 78 38 32

83 47 40 76 38 32

30 21.8 8.9 14.8 11.5 4.8 8.1 12.5 25.5 12

29 20 . 5 9 14 11.3 4.8 8.5 12 23 10

31 22 9 15 11.1 5 8.5 12 . 5 23 11

30 22 8.8 15.5 12 5 8 12.8 22.2

176 120 56 19.8 5.1 18.5 18 7.5 16.5 70

PRIMATES

323

Remarks.-The four specimens listed above were obtained at Saub, Cotabato, Mindanao, April 25, 1923, from the trunk of a tree recently felled. They were taken alive, and kept alive two days. When set free in a closed room they refused to take flight, although there were no apparent injuries. The surface of the outspread wings seems almost insufficient to support the weight of the animal. The odor produced by the gular gland is especially offensive. The curious spinelike hairs on the upper lip seem to function like a currycomb. The tips are flattened and curved like scrapers and the interstices were filled with flecks of epidermis which apparently had not grown in situ. The specimens were infested with two species of forficularid parasites of, or closely related to, the genus A1"ix enia Jordan. The parasites seemed to obtain their food from the surface of the interfemoral membrane, but whether it was blood or epidermis I could not certainly determine. As many as six or eight parasites, each 15 millimeters in length, were found in this region, holding on tightly and refusing to escape or loosen their hold when they were seized. Specimens are in the hands of Dr. K. Jordan, Tring Museum, England. Associated with this bat was another species of the family Molossidre, Philippinopterus latnei, and several dozen specimens of this species were obtained. These likewise were infested with the parasites which caused great unrest among the immature forms; at least, they were continually trying to rid themselves of the pests, which seemed to be ever able to escape their jaws. Cheitromeles torquatus was originally described from Java and has since been taken in Borneo, Sumatra, and the Malay Peninsula. I believe this is the first undoubted record of this remarkable species in the Philippine Islands. Order PRIMATES

This order includes the lemurs, monkeys, apes, and men. Dentition diphyodont and heterodont; orbit invariably surrounded by a ring of bone; clavicles well developed; radius and ulna never united; usually five digits furnished with nails in both hand and foot; pollex may be rudimentary or wanting; hallux, except in Homo, opposable to other digits; pollex, when present, opposable; crecum usually large; placenta either diffuse and nondeciduous or di.scoidal and deciduous; two mamrn.re in pectoral region (Chiromys excepted) ; testes descend into a scrotum. The suborders Lemuroidea and Anthropoidea are recognized.

324

PHILIPPINE LAND MAMMALS

The order Primates (other than Homo) is poorly represented in the fauna of the Philippines. It is only in the southern and southeastern islands of Mindanao, Sulu group, Bohol, and Samar that r epresentatives of the Lemuroidea are known. Two genera of the Anthr opoidea, Pithecus and Homo, occur throughout the ent ire group, the former represented by several island species. No consideration is given in this work to the human anthropoids. The presence of undiscovered primate forms in the Philippines is not improbable. A gibbon, Hylobates !unereus, has been r eported in Sulu (Jolo?) and is included in Hollister's first list, but omitted in the second. I have seen specimens of this form in captivity at Zamboanga and in Jolo. Young orangs are also brought to these towns from Sandakan and other Bornean ports. Travellers in the interior of Mindanao on two or three occasions, have reported seeing species of "monkeys" of large size, but the information is so vague that one must conclude that a specimen of Pithecus has been the basis of the report.

Suborder LEMUROIDEA Placenta bell-shaped, diffused and nondeciduate; allantois very large; cerebellum in an uncovered condition; pollex developed with discoidal tips on fingers; usually nocturnal. Two families of this suborder are represented in the Philippines, the Tarsiidre and the Nycticebidre. The first has three named forms, the second two. * K ey to the Philippin e families of Lemuroidea. a

FO'ur upper incisors, the twO' near the middle separated by a cO'nsiderable interval; fO'ur IO'wer incisO'rs; O'rbit nO't separated frO'm the tempO'ral fO'ssa except by a narrO'w ring O'f bO'ne bO'unding O'rbit. NYCTICEBIDJE (p. 332). Median upper incisO'rs elO'ngate, in cO'nrtact; secO'nd pair in contact with first, smaller; twO' IO'wer incisO'rs; O'rbit separalted frO'm tempO'ral fO'ssa to a large extent by bony partitiO'n; tail elO'ngate and nearly naked .................. .............. _._ ...... _._...... _.......... _...... _.... ..... T ARSIIDJE (p_ 325).

* Very probably there are two species. Hollister, PrO'c. U. S. Nat. Mus. 46 (1913) 328, repO'rt.s the species frO'm MindanaO'. The single specimen is very yO'ung bUll; HO'llister believes it probable that it is distinct from the sO'uthern S'ulu fO'rm. He places it under the name Nycticebus philippinus Cabrera.

325

TARSIUS

Family TARSIIDlE Small animals with a head and body length rarely if ever exceeding 200 millimeters; eyes very large; tail long, slender; head somewhat rounded; legs longer than arms; toes and fingers with flattened pads at tip; nails on toes flat except those on second and third digits, which are compressed and rise like a thorn on upper surface; pollex and hallux opposed to other digits; pupil of eye contracted to a slit during the day. but at night expanded to cover nearly the entire iris. Skull with enormous orbits, their diameter only slightly less than half total length of skull, and separated from each other by an interorbital space usually less than 2 millimeters; first upper incisors larger than upper canines, slightly smaller than lower, strongly in contact; second upper incisors greatly reduced, separated from canines; upper molar teeth strongly trituberculate; lower molar with four or five cusps; first premolar, both above and below, smaller than succeeding tooth; fibula slender, with: its lower half confluent with tibia; calcaneum and navicular bones in feet elongate. Only one genus of this family is known. It comprises several small species distributed from Sumatra through Java, Celebes, Borneo, and the southern islands of the eastern part of the Philippines. Genus T ARSIUS Storr

Ta,rsius

STORR,

Prodr. Meth. Mamm. (1780) 33, Tab. A.

The characters given for the family serve to distinguish the genus. The species are all nocturnal in habit. They are dull, stupid, and slow moving in daytime but show activity at night. There are three Philippine species. Key to the Philippine species of Tarrsius.

a'. Largest form, skull, 41 millimeters; teeth lal"ge and lower molars ................... T. philipp~nsis OJ 2. Similal" but smaller; skull, 37.9 millimeters. T. fraterculus a 3. Skull nanower and proportionally longer. T. carbonarius 0 ••••••••••••••••••••

heavy, especially Meyel" (p. 326). Millel" (p. 327). Heude (p. 331).

These small animals, wherever they occur, are regarded with superstition by the people. The native name of the Bohol tar-

326

PHILIPPINE LAND MAMMALS

sier IS malmag; in Samar the tarsier is known as magau or mago; among the coast Manobos of Cotabato as magatilakok. The Luzon locality usually given, due to the pre-Linnrean record of Camel, * who describes Cercopithecus luzonis minimis with a native name magu or booot, is very probably incorrect. The reason for the Luzon locality may be due to incorrect labelling, since of the native names given, the first is from the Samar Visayan for Tarsius philippensis Meyer; the second, spelled bUrut by Whitehead, is the Igorot (north-central Luzon) name for the rodent Crateromys schadenbergii Meyers. T ARSIUS

PHILIPPENSIS

Meyer

Tarsius philippensis MEYER, Abh. Mus. Dresden No.1 (1894-95) 1; No. 1 (1896-97) 9 (Samar); THOMAS, Trans. Zool. Soc. London 14 (1896) 381; MAJOR, Proc. Zool. Soc. London (1901) 137; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912') 36; Proc. U. S. Nat. Mus. 46 (1913) 338; TROUESSART, Cat. Mamm. 1 (1898-99) 67. Tarsius philippinensis TROUESSART, Cat. Mamm. S'uppl. (1904-05) 39; ELLIOT, Rev. of Primates 1 (1912) 10. Cercopithecus luzonis minimus CAMEL, Phil. Trans. 25 (1705) 2197; PETlVER, Cat. Class. Gazo. Nat. (1764) pI. 13, fig. 11. Tarsius spectrum PETERS, Monaisb. Akad. Berlin (1861) 707 (Samar); STEERE, List of Birds and Mammals Philippines (1890) 28 (Mindanao and Samar); GORGOZA, Anal. Soc. Espanola Hist. Nat. 17 (1888) 9 (Samar and "Visayas").

Description of the species.-Tarsi bare; tail almost naked, the tip covered with sparse hair; color (presumably) red brown; tail longer than in Tarsius spectrum and T. fuscomanus. Remarks.-This brief description contains all the distinctive characters published in the type description. Meyer ha:d at the time of his description an alcoholic specimen of an adult and young from Mindanao (probably Agusan or Surigao Province, collected by Semper) and two stuffed specimens from Samar. No particular one of the lot is designated as type, but I presume that Samar is the type locality since Elliot, who examined the type, records it from Samaroo Elliot gives more data on the type as follows: "Face and top of head reddish brown; upper parts reddish brown, paler than face; outer side of limbs reddish brown, lightest on legs; throat and chest reddish; under parts yellowish gray; tail dark brown." * Phil.

Trans. 25 (1705) 2197.

327

TARSIUS

Measurements of Tarsius philippensis Meyer amd T. ca.rbonarius Heude. Tarsiua Tm'si""l

philippensis.

type.

Skull, total length Occipitonasal length Zygomatic width Intertemporal width Palatal length Breadth of orbits Width of brain case Median length of nasals Length of upper molar series Length of mandible Length of lower molar series Hind foot

carbonariu8,

E. H. Taylor collection. No. 592.

mm.

32 28 22 14 18 23 7 13 26 13 '

37 34 27 22 14 17 22 6.5 13 27 14 65

Variation.-Elliot notes that some of the specimens examined are not as red as the type; but some are dark grayish on the back of the head, with the shoulders and upper back washed with reddish; hands dark brown; feet pale rufous; an example from Mindanao is pale, wood-brown with a reddish tinge. The latter may belong to a different species. A specimen in my collection from Consolacion, Leyte, No . .593, agrees fairly well with the above description of the type; the hind foot measures 66.5 millimeters. The known localities are Samar and Leyte. The Bohol and Mindanao records probably belong to other species. The record for Luzon appears to be incorrect. The types (presumably) were collected by Schadenberg in Samar and are in the Dresden Museum. TARS IUS FRATERCULUS Miller

Tarsius fraterculus MILLER, Proc. U. S. Nat. Mus. 38 (1911) 404; ELLIOT, Rev. Primates 1 (1913) 12; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 36; Proc. U. S. Nat. Mus. 46 (1913) 328-338. Ta1'sius spect?"'Um WATERHOUSE, Proc. Zool. Soc. London (1838) 67.

Description of the species.-(From No. 106, Bureau of Science collection; collected by Andres Celestino, in Sevilla, Bohol, April 9, 1906.) * Small monkeylike animal with body smaller than a * This specimen was collected by Celestino at the same time and in the same place as the types. The specimen was not examined by Miller, however.

328

PHILIPPINE LAND MAMMALS

large rat; eyes very large, set closely together; nose moderately prominent, nostrils separated by a groove; a few scattered vibrissre on snout and on lip below eye, as well as above eye; ear proportionally large, rather rounded at top; a small pocket formed in outer posterior edge below which is a small projecting lobe; a well-developed anti tragal lobe, separated by a notch from rest of ear; a traguslike flap, 6 millimeters wide, rises from lower inner part of ear; ear largely naked; a growth of hair at anterior attachment and along anterior edge of ear; growth of hair in pocket and at base of pocket in outer edge of ear; outer surface of ear furred scarcely halfway to summit; face below eye, and snout covered with short hair not exceeding 3 or 4 millimeters; on crown, about 7 or 8 millimeters; on upper part of neck, from 15 to 18; on middle of back, 12 to 14; on hind leg above, 5 to 7; belly and underside of chin sparsely furred; hand and wrist furred to base of phalanges; underside of phalanges covered with very short almost invisible scattered hairs, the fur extending to palm; digits of foreleg long, somewhat flattened, with flattened tip, each of the four fingers with somewhat compressed, partly elevated nails, pollex only with a 'flat nail; opposition of pollex to other digits not strongly marked; hand with six well-defined rounded palmar tubercles, largest at base of wrist; two others at base of pollex, largest proximal, and one each at base of third, fourth, and fifth digits; hind leg furred to heel; tarsi naked except for a short sparse growth of hair about color of skin of tarsi; toes likewise covered with extremely short, sparse, ' nearly invisible hairs; digits on foot strongly differentiated with flattened pads on tips; hallux much wider than other digits; hallux with perfectly flat nail; second digit shortest, with a compressed erect nail; third digit longer, with a similar nail; fourth digit longest; fourth and fifth with flat nails; pads on feet, four; largest at base of hallux, rectangular, partially divided by a groove; a second at base of fifth toe, elongate, running length of sole; a third at base of second and third toes running more than half length of sole; fourth also elongate, almost contiguous with the largest pad but extending farther back, joining the one on fifth to form an indistinct V; tail with twenty-seven vertebrre, first three heavily furred; following joints naked except for an extremely sparse growth of short hairs, which toward tip form a noticeable growth, gradually longer and thicker toward tip; sparse hairs at tip from 2 to 3 millimeters long.

329

TARSIUS

Color.-Upper part of head and body dull ochraceous-buff, somewhat lighter posteriorly and less noticeable on arms and neck; basal part of fur light pearl to slate-gray, outer color covering only one-third to one-fifth of hairs; belly and throat gray with a suggestion of buff at tips; tail dark brown; ears light brown growing darker towards tips. Skull and teeth.-Skull agrees with description for genus; a distinctly depressed area between posterior part of orbits; postorbital process supported by a thin bone separating orbit to a great extent from temporal fos's a; palate short, not extending behind last molar; palatine processes reach back and attach to large audital bullre which are almost contiguous medially but separated by a deep groove; condyles directly below median portion of brain case; no sagittal crest; two indistinct crests rise posterior to orbits and curve backward over brain case, widely separated. Teeth agree with description given for genus. Measurem ents of Twrsius fratercuLus Mifller. Bureau of Science.

Typ e.

No. 160. min.

Head and body 140 Tail vertebrre 245 Hind foot 64 Pollex 12 Hallux 16 First toe 15 Fourth toe 25 Navicular 25 Skull: Condylobasal length Gl"eatest orbital breadth 33 Zygomatic breadth 28 Least interorbital breadth 1.3 Breadth of brain case over root of zygomata 22 Depth of brain case middle 16 Breadth of rostrum ove;r canines 6.8 Mandible 18 Maxillary tooth row entire 16.4 Mandibular tooth row entire 15

mm .

120 210 60

30.5

31.2 26.5 1.5 21.9 16

6.6 25 16.6 13.6

Remarks.-The type and one paratype were collected by Andres Celestino, of the Bureau of Science, at Sevilla, Bohol, April 9, 1906. The specimen here described was taken at the same time. The two former specimens are deposited in the United

330

PHILIPPINE LAND MAMMALS

States National Museum.

The species is known in Bohol as

malmag.

Hugh Cuming, * who collected this form in Bohol in 1837-38, writes: The Malmag is a small animal living under the roots of trees, particularly the large bamboo of these islands. Its principal food is lizards, which it prefers to all others. When extremely hungry, I have known it to eat shrimps and cock-roaches, and give a great preference to those which are alive. It is very cleanly in its habits, never touches any kind of food that has been parrt;ly consumed, and never drinks a second time from the same water. It seldom makes any kind of noise, and when it does emit sound it is a sharp, shrill call and only once. On approaching it in its cage, it fixes its large full eyes upon the party for a ~ength of time, never moving a muscle: on drawing nearer, or putting anything near it, it draws up the muscles of its face simHar to a monkey, and shows its beautiful sharp regular set teeth. It laps water like a cat, but very slowly, and eats much for so small an animal. It springs nearly two feet at a time. It sleeps much by day, is easily tamed, and becomes quite familiar, licking the hands and face, and creeping about your person, and is fond of being caressed. It has an aversion to the light, always retiring to the darkest place. It silts upon its posteriors when it feeds, holding its food by its fore paws; when not hungry, it will ogle its food for a considerable time. A male and a female are generally seen toget.her: the natives of these islands make sure of taking the second, having secured the first. They are extremely scarce in the island of Bohol, and only found in the woods of Jagna, and the island of Mindanado. [sic]

* * *. It produces one at a time. I had the good fortune to procure a female without knowing her to be with young: one morning I was agreeably surprised to find she had brought forth. The young appeared to be rather weak, buJt a perfect resemblance to its parent: the eyes were open and [it was] covered with hair; it soon gathered strength, and was constantly sucking betwixt its parent's legs, and so well covered by its mother, that I seldom could see anything of itt but its tail: on the second day it began to creep about the cage with apparent strength, and even climb up to the top by the rods: of which the cage was composed. Upon persons wishing to see Ithe young one when covered by the mother, we had to disturb her, upon which the dam! would take the young one in its mouth, in the same manner as a cat,. and carry ilt about for some time; several times I saw her when not disturbed trying to get out of the cage, -with the young one in her mouth as before. It [the young] continued to live and increase in size for three weeks, when unfortunately some person trod upon the tail of the old one, which was protruded through the cage, a circumstance which caused its death in a few days: the young one died a few hours after, which I put into spirits.

* Proc. Zool. Soc. London (1838) 67. Waterhouse.

From field notes published by

TARS IUS

331

TARSIUS CARBONARIUS Reude

(a~'siue)

carbona?'ius HElUDE, Mem. Hist. Nat. Emp. Chinois 4 (1898) 164, pl. 32, figs. 4--7. Tarsius carbonarius HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 36; Proc. U. S. Nat. Mus. 46 (1913) 328.

Description of the sp ecies.-This species is named by Heude in a footnote. The description is as follows: "T. carbonarius, de Mindanao, golfe de Davao et vallee du rio Poulangui. Cette espece se distingue d l'espece type, par la forme de ses premolaires superieures, Ie developpment des molaires. l'etroitesse et la longeur plus grande du crane. On m'a dit qu'il mangeait du charbon !" I have a specimen of a small tarsier in my collection from Saub, Cotabato (No. 529), which may belong to Heude's species. However, his meager description, lacking data on color, markings, measurements, etc., makes it almost impossible to be certain. The figure of the skull, Plate 33, figs. 6 and 7, which Heude gives is clear, but the accuracy of the dr awing may be questioned; he points out that the anterior nasal openings are not correctly drawn. I note, furthermore, in my specimen that the posterior part of the bullre are narrower and the anterior part more inflated than the drawing of the type shows; the brain case of the type is slightly more elongate and narrower than in my specimen. The color of the face, above and surrounding the eyes, as well as the mouth region, shows a decided reddish brown; between the eyes and the snout is a small somewhat darker area continuing backward below eye. The pelage of the back and about base of the legs does not seem to have a uniform color but appears somewhat mottled with small lighter areas of buffy fur; the fur is slate gray except at its distal portion; the tarsi are covered with fine short hair leaving this part of the leg nearly naked; the fur of the belly is grayish buff, the underparts slate, of lighter shade than that of the back; the fur in the region of the neck is tinged with russet. My specimen was captured in the forest at Saub by a Manobo workman. It apparently had been injured as it did not live long after it came into my possession. M easurements.-See under Tarsius philippensis. It may be that specimens from Mindanao hitherto referred by other authors to Tarsius philippensis may belong to this form. The type locality is Davao Gulf and Pulangui River, Davao, Mindanao.

332

PHILIPPINE LAND MAMMALS

The whereabouts of the type is unknown. in the Ze Kai Wei Museum in China.

One presumes it is

Family NYCTIOEiBIDlE Small primitive primates, nocturnal in habit, slow in movement. The body is covered with soft woolly fur; eyes large; ears erect; legs subequal in length; tarsi short; tail short, vestigial, or wanting altogether; third upper premolar smaller than first and possessing one large external cusp, last molar with two, three, or four cusps, in different genera. The recognized genera are Nycticebus, Arctocebus, Perodicticus, and Loris; only the first i,s known in the Philippines. Genus NYCTICEBUS Geoffroy

Nycticebus E.

GEOFFROY,

Ann. 1\I[us. Hist. Nat. Paris 19 (1812) 162.

Small lorises, nocturnal in habit; body rather heavy with rounded head, short muzzle, eyes very large, placed close together directly in front of face; neck short; tail short; limbs well developed but short.; skull shortened, with large flaring eye sockets separated by a narrow flat space; crests prominent; premaxillre not extending very far forward; p.aJate slightly emarginate behind, emargination not extending forward farther than second molar; canines strongly developed, the upper greater than lower, incisors above very unequal, strongly separated medially; first upper premolar elongate. One young born . 2-2 1-1 3-3 3-3 at a time. Dental formula: 1. 2-2' c. 1-1' pm. 3-3' m. 3-3 = 36. The twelve species of this genus are confined to the Oriental Region, and inhabit India, Siam, Malay Peninsula, Sumatra, Java, Borneo, and the extreme southern part of the Philippines. Two of the species have been described as having only two incisors; the others as having four. Only one authentic species is known to occur in the Philippines. This is Nycticebus menagensis Lydekker.* Cabrera's t name Nycticebus philippinus was published to replace N. menagensis, supposed to be without stan'ding in nomenclature, and ill consequence is presumably a pure synonym. The element of doubt enters, however, since Cabrera's specimen presumably came from Mindanao, and if so may

* Zool. Rec.

t Bol.

(1892) 29, 25. Real Soc. Espanola Rist. Nat. 8 (1908) 137.

โ ข"-

333

NYCTICEBUS

represent a distinct species. Hollister * reports a very young specimen from Catagan, Mindanao. He remarks: The discovery of a specimen of the slow lemur in the collection from Mindanao is of very great interest. The specimen is preserved in alcohol and is too young to show if the characters described of the type by Cabrera are constant. It seems best, however, to recognize Ithe Mindanao form, as further collecting will almost surely prove the species distinct from the Tawi Tawi form, N. m,.enagensis. Cabrera's name was proposed chiefly to replace menagensis, supposed at the time to be without Sltanding in nomenclature, but now becomes available for the Mindanao slow lemur.

The presence of this form in Mindanao is a surprise, and it must be very rare. I believe Elera's t records of Bohol, Samar, and Leyte should be disregarded. The question of a species in Mindanao, differing from menagensis is still completely unsettled. I shall disregard the name Nycticebus phitippinus Cabrera except as a synonym of N. menagensis Lydekker. NYCTICEBUS MENAGENSIS

(Lydekker)

Lemur menagensis LYDEKKER, Zoo!. Record; Mamm. 29 (1893) 25. - - menagensis Zool. Anzeig. 15 (1892) 147. (Original name and description disclaimed by N achtdeb.) Nycticebus menagensis TROUESSART, Cat. Mamm. 1 (1898-99) 63; STONE and REHN, Proc. Acad. Nat. Sci. Philad. (1902) 132; THOMAS, Ann. & Mag. Nat. HiiSt. VIII 1 (1908) 469; LYON, Proc. BioI. Soc. Washington 22 (1909) 89; ELLIOTT, Rev. of Primates 1 (1912) 32; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 35; Proc. U. S. Nat. Mus. 46 (1913) 338. Nycticebus philippinus CABRERA, Bo!. Real Soc. Espanola Hist. Nat.

a (1908) 137. Description of the species.-Head round; snout short and flat;

eyes brown, large and round, and slightly prominent; ears hardly projecting beyond hair of head. Neck so short as to give head appearance of being set squarely on shoulders. Nails of hands flat. Hind legs (bowed) with feet turned sharply inward. Color.-General color light rufous, hairs being dark at base, then gray changing to light rufous, with very short gray tips. M easurements.-Total length, 266.7 millimeters; tail, 15.8. Remarks.-The species is presumably confined to the southern part of the Philippines, in fact the southern part of the Sulu group. Capt. Francis Link, many years a resident of Jolo, with

* Proc.

U. S. Nat. Mus. 46 (1913) 328. Fauna Filipina (1915) 49 N.

t Contrib. (Linn.)

(Nycticebus)

tardigradus

334

PHILIPPINE LAND MAMMALS

extensive 路 information on the fauna and flora, writes me that Nycticebu8 mena.gensis does not occur on Jolo, but is common in Tawitawi and on most of the larger islands south of J 010. He states that the name throughout Sulu is kuka.m or kuka.ng. The species is frequently kept as a pet by the natives and carried from one island to another. It is surprising that the form is not more widespread. The following data are taken from Elliot, apparently quoted:* White line between eyes extending backward 1 in. from base of nose. Face around eyes dark rufous, the markings extending upwards on forehead. The effect produced is a heart-shaped mark of dal'k rufous on face, the point of heart being 'on forehead, the eyes occupying two lobes and separated by the white mark which does nolt run to tip of heart. Sides of head at back, and of neck, have hairs broadly tipped with gray. Broad stripe of dark brown extends backwards for 5~ in. along spine tapering to a point. Hairs of arms lighter rufous than that of back. Back of head gray, nearly white. Back of feet grayish. Hair on back of body, arms and legs thick and so:fit, making a fine ful' like that of Ga,leopithecus. On under surface of body hair is thinner and somewhat lighter in color than on back. About the genitals is buffy white. This curious little ani~al is known to the natives of the region it inhabits as eocane. An adult specimen from which the description was taken, was kept alive by Us for seven days. Its movements were sluggish except in biting when its sudden and unexpected activity proved a painful surprise. It moved with equal ease along the upper and lower sides of a small branch or rope, and progressed quite as rapidly backwards as forward. On the floor it was nolt at home and presented a most ludicrous appearance, tumbling along on all fours with feet nearly as far apart as a turtle and its body barely raised from the boards. It spent most of rthe days asleep, rolled up into a furry ball with its head buried between its thighs. If disturbed when actively climbing about, it had a curious way of folding its hands over its eyes and from hence earned the name of "shame-face" which it shares with Tarsius sp.ec't rwm (T. philippinensis). Irt had two notes, a low complaining grunt, and a sharp squeal. During its confinement it took little food, turning up its nose at lemons, but occasionally eating a little banana or egg. We had no inseCJts or small mammals to offer it. In drinking irt lapped up water like a dog. After five days of semi-starvation its strength seemed almost unimpaired, and it showed remarkable tenacity of life.

Elera gives a spirited account of the habits of this animal. Vive este animal formando reducidos grupos que pasan ell dia durmiendo en los huecos 6 ramas de los arboles, enrollado como un erizo, oculta la cabeza entre las piernas, doblando para ello e1 cuerpo, sostenit3ndose con

* Zool.

Anzeig. 15 (1892) 147.

ANTHROPOIDEA

335

los miembros de una rama vertical u horizontal; respira tranquila y profundamente veintidos veces por minuto. Apenas se oculta el sol se despierta, se frota los ojos como hace el hombre cuando esta aun medio dormido, se relame, y limpia como un gato y, dirigiendo sus miradas alrededor, empieza a moverse en busca de alimento, consistente este en frutos suculentos y azucarados, pero prefiere insectos, huevos y pequefios pajarillos, aunque en cautividad es omnivero; no Ie gusta la carne si esta de algun modo cocida, y el pan que come tambien Ie desecha si esta mojado; no bebe nunca agua. EI mejor alimento que se Ie puede ofrecer es un pajarillo que caza admirablemente cogiendole con una rapidez asombrosa; se acerca a el sigilosamente, y cuando est.a a un pie de distancia, se para y poniendose recto sobre las extremidades posteriores, avanza y extendiendo los bl'azos de repente, 10 coge con las' dos manos; Ie deshace y estrangula con singular presteza y 10 devora: 10 primero que se come son los sesos que chupa del craneo, despues de haberlo roto con los dientes, sigue a esto el resto del cuerpo sin desplumarlo antes y se va comiendo la carne y los huesos, despidiendo las plumas con los labios. * * * Como animal nocturno es muy sensible a la liuz del dia, pero de noche ve perfectamente y sus ojos brillan mucho en la oscuridad. Su oido es tan fino que Ie des~ierta el leve rumor que producen a su alrededor los insectos. Parece que es susceptible de alguna educaci6n; ligeros. castigos bastan para que no muerdan y se muestra bastante carifioso con el que Ie cuida, sin embargo no es amigo de muchas bromas. Donwsticado, si se Ie llama, atiende y se acerca pero siempre con mucha precauci6n y timidez. Excepto cuando se irrita, raras veces produce sonidos: su voz consiste ordinariamente en una especie de gemido que varia segun expresa dolor, impaciencia 6 placer: en la cOlera hace oir sonidos agudos y penetrantes, bastante parecidos a los de la ardilla. Guando se intenta quitarle la presa, su mirada se altera, y produce una especie de inspiraci6n, de voz temblona y cuyo sonido es muy agudo; si se lie irrita, grita y repite jay, ay, ay! fuertemente y con un tono plafiidero. Su orina, que es muy abundante y frecuente, despide un olor muy fuerte y desagradable, y sus excrementos tienen la forma de los de las ovej as; y todo el animal despide un 0101' repugnante. En algunos puntos donde habita creese pOI' los indigenas 10 que de la especie siguiente, esto es, que su presencia acarrea peligros, enfermedades, muertes u otras desgracias parecidas, por 10 que huyen y evitan la vista y encuentro con este animal. Los indigenas de Filipinas, ni de est a ni de la especie siguiente (Tarsius spectrum) tienen semejante ridicula prevenci6n.

The common name for the species is slow lemur. the name tonger vergonzoso.

Elera gives

Suborder ANTHROPOIDEA

Unguiculate, claviculate placental mammals with orbits surrounded by bone; three kinds of teeth; brain always with a

336

PHILIPPINE LAND MAMMALS

posterior lobe; innermost digits of at least one pair of extremities opposable; hallux with a flat nail or none; a well-developed crecum; penis pendulous; testes scrotal; always two pectoral mammre. (After E'lliot.) * This group contains the larger forms of the order which include the anthropoid apes and man ; only a single family t and a single genus Pithecus, in the Philippines other than Homo. Family CERCOPITHECIDJE Genus PITHECUS E. Geoffroy

Pithecus GEOFFROY, Mag. Encyc1. 3 (1795) 462; ELLIOT? Review of Primates 2 (1912) 176.

M acaca LACEPEDE, Tabl. Mamm. (1799) 4. Silenus GOLDFUSS, Handb. Z061. 2 (1820) 479. Mairnon WAGNER, Schreb. Saugeth. Suppl. 1 (1840) 141 and 148. Rhesus LESSON, Rev. Zoo1. (1840) 49, 95. Pithex HODGSON, Journ. Asiat. Soc. Bengal 9 (1840) 1212, pI. 2. Cynamolgos Rm0HENBACH, V ollstand. N aturg. Affen. (186Z) 130, pI. 23, figs. 237, 344.

Body heavy, compact; limbs short, stout; thumb pointing backwards; nose long, not extending beyond upper lip; nostrils opening downwards, not placed at extreme end of nose; eyes approximate, surmounted by heavy bony ridges, ears naked, pointed; callosities large, buttocks nude; tail of various lengths, and tufted; hair of head sometimes long or manelike about face and extending downward on neck to shoulders; cheek pouches present. Skull with small brain case, and prominent orbital ridges; canines long and formidable; first and second lower molars with four cusps; last molar, largest, with five cusps and posterior talon. (After Elliot.) This genus is represented by five t island species in the Philippines. The differences are not striking.

* Review

of Primates 1 (1912) XI.

t Elera, Contr. a la Faun. Filipina Manila 1915 lists Semnopithecus albipes Geoff. from J 010, Cynopithecus othiops Tim. J 010, Negros, Guimaras. These records may be disregarded. Two other species, Macacus speciosus and Macacus nernestrinus var. philippina (subsp. nov.?), are listed. l In the Mehan Gardens, Manila, I observed (1923) a specimen of Pithecus said to be from Panay. The specimen was light reddish brown in marked contrast with other specimens on exhibition in the gardens.

PITHECUS

337

Key to the Philippine species of Pithecus. a '. Outer side of arms and legs mixed gray and ochraceous tawny becoming duller, almost old gold near wrists and ankles; hands and feet grayish buff. (Mindoro.) ............ P. mindorus Hollister .(p. 337). a". Outer side of arms and limbs mummy brown mixed with smoke; gray on wrists, ankles, feet, and hands. (Luzon.) P. philippinensis (Geoffroy) (p. 338). a I. Outer surface of limbs tawny-olive, becoming drab-gray on fingers and toes. (Mindanao and Basilan. L .. P. mindanensis (Mearns) (p. 340). a '. Skull larger than rnindanel1sis with wider frontal and zygomatic arches; rostrum shorter and broader; a udi tal bullre more rounded; color unknown. (Jolo.) ...... .......................... P. suluensis (Mearns) (p. 342). a". Outer surface of limbs olive-brown like back above but fading at knee :md elbow joints to drab-gray on forearms , hands and feet. (Cagayan Sulu.) ... ................................. P. eagayanus (Mearns) (p. 342). PITHECUS MINDORUS Hollister

Pithee-ILS mindorus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 328, pIs. 27 and 28 (skull and teeth).

Description of the species.-Large; coloration dark; externally resembling Pithec'us philippinensis, of Luzon, but darker and richer colored; skull much like that of P. mindanensis, of Mindanao and Basilan, but larger. Color.-General color of upper parts dark, rich, russet-brown, most intense on head and shoulders; hairs long and coarse, deep mouse gray at bases and marked on terminal half with alternate rings of black and rich ochraceous-tawny; usual black markings on head; sides and hips lighter than back, with less black and more yellow; outer sides of arms and legs mixed gray and ochraceous-tawny, becoming duller, almost old gold, near wrists and ankles; hands and feet grayish buff; underparts, including inner sides of limbs, dirty buffy gray; tail blackish above near base, becoming dark gray at end; below dark buffy gray; iris yellowish brown; bare space above eye grayish white; bare skin of muzzle light brown; feet olive-plumbeous; claws plumbeous-black; scrotum brownish flesh color. Skull and teeth.-The skull differs greatly in general shape from that of Pithecus philippinensis which is long and comparatively narrow with a deep narrow palate; it is shaped almost exactly like that of the Mindanao species, broad and rounded, with wide and heavy rostrum and wide shallow palate. 269463-22

338

PHILIPPINE LAND MAMMALS

but it is decidedly larger, with a heavier mandible; teeth show small supplementary middle inner cusps on molars as in P. mindanensis. These cusps are apparently absent in P. philippinens~s. (After the type description.) Measurements of the type Qf Pithecus mindorus HoUister. mm.

Total length Tail Hind foot Girth of chest Skull: Greatest length Condylobasal length Breadth of rostrum above canine Breadth of brain case over roots of zygoma Pala,t al length Postpalatal length Upper molar-premolar series Mandible Mandibular molar-premolar series

1,100 580 140 390 136 112 369 62.9 57.8 45 32.9 102 42.2

Remarks.-The type and two other specimens, in the United States National Museum, were collected in Mindoro by Edgar A. Mearns in November, 1906. Three localities are represented; namely, Alag River, Bulalacao, and Mount Halcon. Hollister states: "While differing specifically from the Luzon and Mindanao species, the Mindoro macaque exhibits a curious combination of the distinguishing characteristics of both." PITHECUS PHILIPPINENSIS

(Geoffroy)

Macacus philippinensis GEOFFROY, Archive du Mus. d'Hist. Nat. 2 (1843) 568 pI. 33 (1841) (Luzon); STEERE, List of Birds and Mammals Collected by Steere Exp. (July 14, 1890) 28 (part.). Macacus palpebrosus GEOFFROY, Cat. des Primates 93 (Manila). Macacus fur SLACK, Proc. Acad. Nat. Sci. Philadelphia 19 (1867) 36, pI. 1. Macacus cristatus GRAY, Cat. Monk., Lemurs, and Fruit-eat. Bats in Brit. Mus. (1870) 30. Cynamolgos philippinensis RIillCHElNBACH, Vollstand. Naturg. Affen (1862) 134:, pI. 23, fig. 340 (albino) . Macacus cynomolgus THOMAS, Trans. Zool. Soc. London 14 (1898) Luzon, Palawan, Mindoro, N egros. Cynamolgos palpebrosus REICHENBACH, Vollstand. Naturg. Affen (1862) 137. Pithecus (Macacus) philippinensis DAHLBERG, Stud. Zool. Fam. Reg. Anim. Nat. fasc. 1 (1856) 118, 120. Cynomolgus phil'ippinensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 426.

339

PITHECUS

Macacus cynomolgus c~~1ningii GRAY, Cat. Monkeys, Lemurs, and Fruit-eating Bats (1870) 30. Pithecus philippinensis ELLIOT, Rev. of Primates 2 (1912) 248. Pithecu8 syrichta HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 37; Proc. U. S. Nat. Mus. 46 (1913) 328.

Description of the species.-This form is slenderer than Pithecus mindanensis Mearns, but of the same general characters. Color.-Like all species of this group, the pelage of upper parts and outer surface of limbs is annulated with light and dark; light rings and glossy texture of hair cause colors and shades to change in different lights'; upper parts generally mummy brown, the hairs everywhere annulated with blackish; top of head strongly tinged with chestnut; sides of head grizzled yellowish olive; forehead ornamented with a row of stiff black hairs scarcely separated from naked orbits by a narrow rim of short, brown hair, which is cut off by intervening black in center; underparts, including inner surface of limbs, scantily clothed with pale drab-gray hairs of a silky texture; genitalia broccoli brown; outer surface of limbs mummy brown mixed with smoke gray on wrists, ankles, feet, and hands; tail grayish black above, hair brown below, with a gradual blending of these colors on sides of tail. (After Mearns.) Measu1'ements of Pithecus philippinensis (Geoffroy)." mm.

Total length Tail Foot Skull: Total length Occipitonasal length Interfemoral width Zygomatic width Breadth of brain case Length of nasals Palatal length Length of upper molar series Length of lower molar series Length of mandible

1,665 550 120 125 104.4 41.9 77.5 60.4 28.5 48 29.8 38.3 90.9

n From Elliot, Review of Primates 2 (1912) 249.

Remarks.-Elliot, while accepting Mearns's species from J ala and Cagayan, considers the Mindanao species as identical with the species occurring on Luzon. The measurements here given are from a southern Luzon specimen.

340

PHILIPPINE LAND MAMMALS

That all of the specimens reported by Steere under the name Macacus phUippinensis, from Paragua, Mindanao, Basilan, Negros, Samar, Leyte and Luzon, actually belong to this species is to be doubted. More probably the southernmost specimens represent Pithecus mindanensi,c;. Mearns. The specimens from Palawan ("Paragua") may represent an undescribed species of equal value with the other island forms. The specimen reported by Thomas from Mindoro may very probably be Pithecus mindorus Hollister. The common custom of transporting pet monkeys from island to island and their frequent escape have doubtless resulted in mixing of the island forms so that the original distinctive characteristics may be somewhat obliterated. This species appears in the recent lists * as Pithecus syrichta Linnreus. Cabrera t has recently shown that this name is based on a species of Tarsius and cannot be used for the Philippine macaque. PITH~CUS

MINDANENSIS

(Mearns)

Cynomolgus mindanensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 428.

Cynomolgus mindanensis a-poensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 429.

Pithecus minda-nensis mindanensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 37. Pithecus mindanensis apoensis HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 37.

Pithecus mtindanensis HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 329.

Description of the species.-Stouter than Pithecus philipp~"nensis and different in color. Color.-Upper parts raw umber, .hairs everywhere annulated with blackish; top of' head strongly washed with burnt umber; sides of head pale grayish olive; forehead with grizzled brown front, ornamented with long stiff black hairs, forming a bushy brow; underparts pale drab-gray; genitalia broccoli-brown; outer surface of limbs tawny-olive becoming drab-gray on fingers and toes; tail slaty black at base above, becoming paler toward extremity, and olive-drab below, with a few tawny annuli to hairs of upper side near base. The young are grayer and less brownish.

* Hollister, t

Philip. Journ. Sci. ยง D 7 (1912) 37; Proc. U. S. Nat. Mus. 46 (1913) 328. Cabrera, Journ. Mamm. 4 (1923) 89-90.

341

PITHECUS

Skull and teeth.-Foramen magnum larger and lambdoidal crests more arched and wider, showing less posterior expansion of brain case than in P. philippinensis; teeth very similar. (After type description.) Measu?'ements of the type of Pithecus mindanensis (Mearns). mm.

Total length Tail vertebrre Tail with hairs Hind foot Head Ear above crown Ear above notch at base Skull: Greatest length without incisors Basal length Basilar length Palatal breadth between first molars Zygomatic breadth Mastoid breadth GreaJtest breadth of brain case Orbital breadth Mandible Maxillary tooth row exclusive of incisors Mandibular tooth row exclusive of incisors

1,100 600 633 137 143 32 39 125 97 94 24 83 64 57 62 91 38 46

Remarks.-The type was collected by Edgar A. Mearns at Pantar, Davao, Mindanao, August 26, 1903. Nine other specimens were collected in Davao and one in Basilan. All are in the United States National Museum. There are two specimens in my collection from near Kraan Point, on the Cotabato coast of Mindanao. The variety apoensis Mearns differs slightly in color as follows: Characters similar to Pithecus mindanensis. but paler and different in color. I believe this to be individual variation. Color.-Upper parts yellowish olive, hairs everywhere annulated with blackish; top of head slightly washed with burnt umber; sides of head pale olivaceous gray, annulated with darker; forehead pale olivaceous gray, separated from crown by a strong fringe of stiff black hairs; face gray; underparts whitish smoke gray; genitalia bister; outer surface of limbs pale yellowish olive, changing to olive-gray on lower portion and on hands and feet; upper side of tail slate-black at base, fading to mouse gray on terminal half and smoke-gray on underside. Young and nearly mature specimens are colored almost exactly like the two adult females.

342

PHILIPPINE LAND MAMMALS

Skull and teeth.-Compared with P. mindanensis mindanensis,' Rostrum shorter, brain case wider; zygomatic arches more expanded; orbital breadth slightly greater; molariform teeth slightly narrower. (After the type description of Pithecus mindanensis apoensis. Mearns.) PITHECUS SULUENSIS (Mearns)

Cynomolgus suluensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 430. Pithecus suluensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 37; Proc. U. S. Nat. Mus. 46 (1913) 328.

Description of the species.-Larger than Pithecus mindanensis and P~ philippinensis, with different coloration and cranial characters. Skull and teeth.-Compared with C. mindanensis the skull is larger with relatively wider frontal and zygomatic arches; rostrum shorter and broader; audital bul1re more rounded; molariform teeth smaller without marginal tubercles on inner border. Measurements of the type of Pithecus suluensis (Mearns). mm.

Skull: Greatest length, exclusive of ineisors Basal length Basilar length Least palatal length Palatal breadth between the front molars Zygomatic breadth Mastoid breadth Greatest breadth of brain case Orbital breadth Greatest depth of brain case, excluding sagittal crest Mandible Maxillary tooth row, excluding incisors Mandibular tooth row, excluding incisors

126 96 92 51 22 91 64 58 73

49 93 40 43

Remarks.-The type, in the United States National Museum, was collected at the foot of Crater Lake Mountain, J 010 Island, November 1.6, 1903, by Edgar A. Mearns. The skin was not retained. PITHECUS CAGA Y ANUS (Mearns)

Cynomolgus cagayamts MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 431. Pithecus cagayanus HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 36; Proc. U. S. Nat. Mus. 46 (1913) 330.

De8cription of the species.-This is the smallest Philippine species, and is about three-fifths the bulk of Pithecus mindanensis or P. philippinensis.

343

CARNIVORA

Color.-(From the type skin in alcohol.) Upper parts olivebrown with scarcely a trace of chestnut or burnt umber on head; sides of head brownish gray, face smoke-gray; forehead grayish in front, behind which a crest of stiff black hair arises; underparts pale drab-gray; outer surface of limbs like back above, but fading at knee and elbow joints to drab-gray upon forearms, hands, legs, and feet; tail above slate-black at base, fading to mouse-gray towards end, and drab-gray on underside. Skull and teeth.-The skull of the type is similar to that of the male of Pithecus mindanensis but is only as large as the female of that species; the proportion of weight in the sexes being as 9 to 15. The size, therefore, would be only threefifths of that of P. mindanensis. In form the skull closely resembles that of P. suluensis, but the dentition is relatively much heavier; the audital bull~ are small but inflated and prominent apically. Measurements of the type of Pithecus cagayanus (Mearna). mm.

Skull: Greatest length, exclusive of ~ncisors Basal length Basilar length Least palatal length Pa~atal breadth, between first molars Zygomatic breaclth Mastoid breadth Least breadth (behind orbits) of brain case Greatest breadth above roots of zygomata Orbital breadth Greatest depth of brain case, exclusive of sagittal crest Mandible Greatest depth of ramus Maxillary tooth row, exclusive of incisors ' Mandibular tooth row, exclusive of incisors

110 81 77 46 20.5 77 60 38 55 59 44 81 116 39 44.2

Remarks.-The type, in the United States National Museum, was collected in Cagayan Sulu, an island in' the Sulu Sea, north of Borneo, February 25, 1904, by Edgar A. Mearns. Only the type is known. It represents a dwarfed, small island form. Order CARNIVORA

This order includes a very diversified group of animals. The common characters of the order are numerous, but no especially important character separates the members from all other orders. The following characters are found in all of the families: Un-

344

PHILIPPINE LAND MAMMALS

guiculate, having at least four digits on limbs, usually with pointed nails; pollex and hallux never opposed; stomach simple; seminal vesicles wanting; -uterus bicornuate ; mammre abdominal; placenta deciduate; fibula always distinct; dentition diphyodont and heterodont. The following characters are more or less variable: Three incisors on each side of jaw, first smallest, third largest; strong, recurved, pointed canines; teeth not lobulated by infolding of enamel, usually with conical cusps; clavicle frequently absent and never complete when present; condyle of lower jaw a transverse half-cylinder, working in a deep glenoid fossa. Most of the species are carnivorous, although many use plant food and many are omnivorous. They usually prefer freshly killed warm-blooded animals. The following are the general divisions of the order: Section Arctoidea. Family Canidre; dogs, wolves, and foxes. Family Procyonidre; raccoons. Family Ursidre; bears. Family Mustelidre ; martens, weasels, and otters. Section Aeluroidea. Family Viverridre; palm civets. Family Protelidre; aard wolves. Family Hyaenidre; hyaenas. Family Felidre; cats.

Only the families Canidre (domestic), Mustelidre, Viverridre. and Felidre are represented in the Philippines; most of the species are small, and the Philippine species of several of the genera are found only in Palawan. Key to the PhiliP'JYine families of Carnivora . a 1. Typical cats; dental formula

. 3

3' c. l' pm. 2' m'I = 30 .

FELIDlE (p. 345) . Typical dogs; dental formula for family variable (usually in Canis) . 3 1 4 2 1. 3' c. I' pm. 4' m. 3 = 42 --- - - . ------ --- - - -- ----- CANIDlE (p. 348).

Martens, otters, and skunks; dentition variable; anal gland in Mydaus producing an offensive odor _._. ___...... _... _...... _... _._. MUSTELIDlE (p. 348). a'. Palm civets; catlike animals with elongate bodies and tails; a glandular follicle on perineum secreting a musklike odor. VlVERRIDlE (p. 353).

345

FELIS

Distribution of the genera of Carnivora in the Philippines.

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Family FELIDJE The cat family is 路r epresented in the Philippines by a large domestic cat (sometimes feral) and a small wild cat. The domestic cat is a variety of Felis c'a tus Linnreus; the wild cat is longer limbed, slenderer, and has a longer neck and a longer tail. Genus FELIS Linnmus FELIS CA,TUS Linnlllu8

Felis catus LINNlEUS, Syst; Nat. 10 led. 1 (1758), 42; POCOCK, Proc. Zool. Sec. London (1907) 149; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 24; Proc. U. S. Nat. Mus. 46 (1913) 336. Felis domestica THOMAS, Trans. Zool. Soc. London 14 (1898) 387; Ann. & Mag. Nat. Hist. VI 18 (1896) 245.

The Philippine domestic cat is too well known to warrant description. The variation in the "tabby" coloration is not especially great. A large proportion of the specimens are of the Manx short-tailed or (occasionally) tailless variety. It is highly probable that this form originated in the Oriental region as a result of artificial selection. Pocock * says: The conclusion that the shortness or complete absence of the tail in 80called "Manx cats" has been brought about by selective breeding from favored sports must be regarded as beyond dispute. There is no such thing as a tailless species of Felis among fossil or recent forms. The tail

* Proc.

Zool. Soc. London (1907) 144.

346

PHILIPPINE LAND MAMMALS

is short in Lynxes, but it cannot be claimed that an intermixture with any known specieS! of Lynx has contributed to the abbreviation of the tail in the "Manx." When and where this breed arose has been much debated. Some have suggested Spain as its original country, others China. But in al] probability the "sport" has appeared independently and been preserved by selective crossing on many occasions and in many places.

Feral forms of the domestic cats are usually larger than the "domestic" sorts. Thomas, (loc. cit.) quotes J. Whitehead as follows: In North Luzon we obtained a very large specimen of a Wildcat, on the mountains at an altitude of 7,000 feet. This animal, I am told, is a feral race of the Domestic Cat, Felis domestica, but it is unlike any cat that exists in the native villages of today, being nearly double the size of any Igorrote Cat, and tabby marked, on a rather sandy ground. My friend, Mr. A. H. Everett, however, informs me that he obtained a Wild Cat very like it in Celebes, which turned out to be an offspring of some escaped Domestic Cat. FELIS MINUTA Temminck

Felis minuta TEMMINCK, Monog. des Mamm. 1 (1827) 130; GUNTHER, Proc. Zoo1. Soc. London (1879) 75; THOMAS, Trans. Zoo1. Soc. London 14 (1898) 387; SANCHEZ, Anales de la Sociedad Espanola de Hist. Nat. II 27 (1898) 93 (Palawan); ALLEN, Bull. Am. Mus. Nat. Hist. 28 (1910) 13; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 24; Proc. U. S. Nat. Mus. 46 (1913) 336. Felis javanica ELElRA, Contr. Fauna Filipina (1915) 167, pI. (unnumbered) .

Description of the species.-(From a specimen in possession of Mr. Macasaet, Iwahig Penal Colony, Iwahig, Palawan, collected at this same locality.) Typical cat form, body slender, somewhat elongate; tail not abbreviated but at least half the length of body; ears shorter than in the domestic cat and somewhat farther from eye. Color and markings.-Ground color of fur above mouse-gray at base, distal part of hair tipped with dull brownish white, growing grayish brown at extreme tip of about half the hairs, in others silvery gray; a brown stripe on nose passes onto forehead, bordered on either side by a short whitish stripe, which merges into the ground color of fur; beginning above level of eyes four dark lines continue more or less irregularly behind shoulders where they become broken into spots; two or thrl~e short dark and white lines on sides of face; chin pure white; a dark collar on sides of neck passes under throat where it is dim; line on neck preceded by a diagonal spot; back, sides, and limbs spotted blackish brown, forming indistinct lil1es on sides;

FELIS

347

spots on legs same shade of brown as those low on sides; breast white, with a few dark brown lateral spots; belly gray-white with dark brown spots; tail spotted only at base; greater part of tail smoky brown, growing silvery white toward tip; lower part of hind leg unspotted, smoky gray brown; ears above dark brown with a lighter area in the middle. Measurements.-The mounted specimen stands about 240 millimeters high; head and body, 440; tail, 190. Remarks.-The specirpen here described was a mounted one in the private collection of Mr. Macasaet, an official of the Iwahig Penal Colony, Iwahig, Palawan. The skull was mounted in the skin and consequently I could not examine it. The flpecimen was caught in a rice paddy near I wahig when the stacked rice was being moved. Mr. Macasaet assures me that the species is relatively rare, only a few specimens having been seen in the colony. . I examined a privately owned specimen when it was in the Bureau of Science for mounting. The notes and measurements made have been mislaid and are not available. This specimen was from N egros. Besides these two lo~alities the species is known from Panay, Calamianes, and Cebu (possibly also Samar) in the Philippines. It occurs in Java, which is the type locality, Borneo, and Sumatra, as well as on the Asiatic mainland. It is known that cats of this species are beneficial, as they destroy rats in rice and sugar-cane plantations. Whitehead * states: This handsome little Cat is apparently found only in the islands of Panay, N egros and Cebu; but as it also occurs on the great continental island of Borneo, doubtless it will some day be found in Mindanao. One of my hunters declared that he shot a Wild Cat in Samar among some rough broken up limestone, into which the wOllntled animal unfortunately disappeared. I think we may say for certain that this Cat does not occur in Luzon, which is so well cultivated that it could scarcely have escaped detection. In Mindoro, it might be possible for this animal to have escaped detection, as the island is perhaps, after Minda.nao, the wildest and most densely covered with forest of the whole gyoup. In Negros where we obtained a specimen of Felis minuta the animal frequented the sugar-plantations, where it finds an abundance of rats. During the harvesting operations this Cat is often captured lJy the natives, who form a ring around the last patch of standing cane. One of -my collectors said that he saw this animal as high as 6,000 feet, on CanIaon volcano.

* Trans.

Zool. Soc. London 14 (1898) 387.

348

PHILIPPINE LAND MAMMALS

The Visayan name for the species, according to GUnther, is tamaral. Elera gives the names singalong and milo, but does not state where they are used. Family CANIDJE The dogs of the Philippines are too well known to warrant description. They appear to belong to a stock developed in southern Asia and scattered widely in various parts of the Malay Archipelago. Except for few recent importations, the domestic dog is rather uniform in character. Genus CANIS Linnreus

Only the genus Canis, with a single domestic species, a variety of Canis familiaris LinTIlreus, occurs in the Philippines. Studer has described a Canis familiaris sumatranus.* Whether or not this represents a form similar to the Philippine dog I am unable to determine. CANIS F AMILIARIS LinnreuB

Canis familiaris

HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912); Proc. U. S. Nat. Mus. 46 (1913) 313. ? Canis pollokensis ELERA, Contr. Fauna Filipina (1915) 193.

No description of this common dog is appended. Elera describes a dog under the name Canis pollokensis as follows: Canis de Mindanao que Haman Ira. Es pequeno y mal tallado, de orejas cortas y enhiestas, valiente y cazador; en vez de ladrar, aulla; de dos palmos de alto; los hay de varios colores y algunos atigrados.

I am of the opinion that this does not constitute a form worthy of even subspecific designation. Family MUSTELIDJE Molars 1f2 (rarely Yt) ; no alisphenoid canal; in upper molar the inner tubercular portion always longer in the anteroposterior direction than the secant external portion; audital bullre very slightly inflated; palate much produced behind molars; postglenoid process of cranium generally considerably curved over glenoid fossa, so as to hold condyle of mandible firmly. Animals of this family are usually known as otters, weasels, badgers, skunks, polecats, martens, etc. The family is divided by Osborne t into three subfamilies and sixteen living genera.

* Abhandl.

t Age

Schweiz. paleont. Gesell. 28 (1901) 1-37. of Mammals (1910) 530.

MELINJE

349

Two of the subfamilies occur in the Philippines and a third is of doubtful occurrence. [Subfamily Mustelinre (doubtful).] * [Genus Mustela Linnreus.] [Mu stela nudi pes Desmarest.] Subfamily Melinre. Genus Mydau s Cuvier. Mydaus mar chei Huet. Mydaus schadenbergii J entink. Subfamily Lutrinre. Genus Aonyx Lesson. Aonyx ciner ea (Illiger).

Key to the Phi lippine su bfa'l1~ilies of the Mu s teli dre. a ' . Feet not webbed __ __ _____ ___ ___________________ _____ ____________ ____________ MELINlE (p. 349). (t ". Feet webbed _________ __ ____ _______ ___________ ___ ____ _________________________ ___ LUTRINlE (p. 352). Subfamily MELINยฃ

Form variable; body usually somewhat elongated, feet elongated; with straight toes ; claws non retractile, slightly curved, subcompressed, blunt ; those of forefoot especially enlarged; upper molar variable. Dental formula varies, the teeth total either 32, 34, or 38. In the genus Mydaus, which occurs in the Philippines, there are 38 teeth. Mart es and Mus tela, belonging to this subfamily, have been reported in the Palawan group and certain Sulu Islands. Sanchez t excludes the forms from the Philippines, and while Hollister:j: noted them in his first list as doubtful, he omits them entirely in the second list. The species given are Mustela fia,vigula henricii Westerman = Martes henricii (Westerman) (Sulu) and Putorius nudip es (Desmarest) = Mustela nudip es Desmarest (from Palawan, Calamianes, Sulu, etc.). While it is highly probable that they or other representatives of the same genera occur in the Islands, J am aware of no * The genus Ma'rtes, of the subfamily Mustelinre, has been reported by Trouessart, Cat. Mamm. Suppl. (1904) 203, under' the name Mustela fia'1:igula henricii, in Sulu. The locality is so indefinite that it seems best to omit this record until more evidence is forthcoming. Hollister admits the species to his first list, Philip. Journ. Sci. ยง D 7 (,1 912) 20, with doubt. but omits it from his second Philippine list, Proc. U. S. Nat. Mus. 46 (1913). l' An. de la Soc. Espanola de Hist. Nat. II 29 (1900) 177-290. :j: Philip. Journ. Sci. ยง D 7 (1912) 20-21.

350

PHILIPPINE LAND MAMMALS

authentic records or actual specimens. Consequently, I shall follow Hollister in omitting them from the Philippine list. Genus MYDAUS Cuvier

Mydans F.

CUVIER,

Hist. Nat. Mamm. 3 (1821) 27.

Small badger like animals; head pointed in front; snout produced, mobile, obliquely truncated, nostrils inferior; legs short and stout, tail extremely short, clothed with rather long bushy hair; anal glands well developed, emitting an odor somewhat like American skunks. Teeth with cusps rather acutely pointed; cranium elongated, face narrow and produced; suborbital foramen small, palate produced backward about midway between last molar and glenoid fossa. Vertebrre: Cervical, 7; dorsal, 14 or 15; lumbar, 6 or 5; sacral, 3; caudal, 12. Dental formula:

i.~, c.

pm.

i, m. l = 38.

The species Mydaus schadenbergii J,e ntink and M. marchei Huet have been described from the Philippines. MYDAUS SCHADENBERGII Jentink

Mydaus schadenbergii JENTINK, Notes Leyden Mus. 17 (1895) 46; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 21; Proc. U. S. Nat. Mus. 46 (1913) 336.

Description of the species.-Characters of the genus; fur soft to touch; about same size as Mydaus meliceps of Borneo; claws much less developed than in other forms. Color.-Albove blackJ with white hairs scattered here and there; no white crown spot; no trace of white line along middle of back; tail without white tip, all these parts being uniform black like the rest of back. Skull and teeth.-Skull smaller than in other species; lower jaw more curved and its depth and length less than in other forms; hindmost upper molar very large, and penultimate lower molar a much better developed tooth than in other species; claws much less developed than in other forms. M easurements.-The measurements of the type have not been published except that of. the lower molar tooth, which is 10 millimeters. Remarks.-Only the type is known. This was collected by Schadenberg in the Calamian Islands. The exact locality is unknown. I presume that it is actually from Busuanga Island,

351

MYDAUS

since it is known that Schadenberg collected at Malbato, a hacienda near Coron, Busuanga. MYDAUS MARCHEI

Duet

Mydaus ma1'chei HUET, Le Naturaliste II 9 (1887) 149 (Palawan); SANCHEZ, Anal. de la Soc. Espanola de Hist. Nat. II 29 (1900) 177290; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 21; Proc. U. S. Nat. Mus. 46 (1913) 312; MEYER, Abh. und Ber. des Konig. Zool. Mus. Dresden (1894-95) No. 13, 1-4. Mydaus meliceps ELERA, Contr. Fauna Filipina (1915) 7, 194, pI. unnumbered.

Description of the species.- (From an unnumbered specimen in the collection of Mr. Macasaet, Iwahig Penal Colony, Iwahig, Palawan. The specimen was collected at Iwahig.) Head rather elongate, with a pointed truncate snout like a pig; eyes small; sides of snout with a few vibrissre about 10 millimeters long; ear very small, scarcely raised above surrounding skin, distance from posterior tip of ear to eye 30 millimeters; eye to end of snout, 30 millimeters; nostrils opening on anterior end of snout; snout unfurred; legs short, body covered with coarse fur; tail short, covered sparsely with hair, which extends much beyond tip. Skull and teeth.-The dentition of this specimen is imperfect as some of the teeth are wanting. Color.-Above uniformly dark brown, the hair growing gray toward base; fur on posterior part of hind leg and anal region light brown; fur on belly and neck of about same shade of brown as back; a triangular white spot on head and neck. Measurements of Mydaus marchei Huet. mm.

Head and body, estimated Skull: Total length Width of brain case Zygomatic breadth Length of palate Height of brain case Distance between canines Mandible Length of large lower molar Width of large lower molar Claws on forefeet Claws on hind feet

450 91 43 50 44 27 9

54 9 5 15 10

Remarks.-A second specimen in Mr. Macasaet's collection differs from the described specimen in having a smaller silvery

352

PHILIPPINE LAND MAMMALS

white spot on head and neck. Both specimens were captured on the farm of the Iwahig Penal Colony and are now mounted. The stench resulting from the secretions of the anal glands is said to be extremely offensive. The animal is known as pantot in the dialect spoken at Iwahig. Elera reproduces a photograph of a mounted specimen of "Mydaus meliceps" which shows the face nearly white with a dark streak between the eyes. He lists Paragua, Calamianes, and Balabac as localities. Meyer is of the opinion that Mydaus schadenbergii Jentink is a synonym of M. marchm Huet. Subfamily LUTRINJE

Body generally elongate; feet short, rounded; toes webbed; claws small; head much depressed. Teeth diverse in form, with a single upper molar, which is enlarged and squarish, inner tubercular portion somewhat expanded anteroposteriorly. Aquatic. Of the two or three living genera, only Aonyx is represented in the Philippines. Genus AONYX Lesson

Aonyx Lutra

LESSON,

Man. Mammalogie (1827) 157. Syst. Regn. Anim. (1777) 448.

ERXLEBEN,

Characters of the subfamily. Skull broad, depressed, with a short anterior portion; brain case large; first upper premolar small, sometimes wanting; upper carnassial (third premolar) with a trenchant tricuspid blade, inner lobe large, hollowed on the free surface, with a raised sharp cutting edge, and extending along two-thirds or more of length of blade. Body long; ears short and rounded; legs short, feet more or less completely webbed; claws usually developed but may be vestigial or absent; tail long, thick at base and tapering, somewhat depressed; fur short and close; five toes. Cervical vertebrre, 7; dorsal, 14 or 15; lumbar, 6 or 5; sacral, 3; caudal, 20 to 26 . . 3 1 4 1 Dental formula: 1."3' c. l' pm. '3' m. "2 = 36.

Aonyx cinerea (Illiger) has been found in the Philippines only in Palawan and probably does not occur outside the Palawan group. It is also known from Asia, Sumatra, Java, and Borneo.

353

VlVERRIDlE AONYX CINEREA (Illiger)

liutra cinerea ILLIGER, Abh. Ak. Berlin 1811 (1815) 99; BONHOTE, Fasc. Malay (1903) 11. ? L(utra) sumatrana ELERA, Contr. Fauna Filipina (1915) 197, pI. 1 (Paragua and Balabac). Aonyx cinerea LYON, Proc. U. S. Nat. Mus. 36 (1909) 485, pI. 39; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 21; Proc. U. S. Nat. Mus. 46 (1913) 312 (Palawan); ALLEN, Bull. Am. Mus. Nat. Hist. 28 (1910) 17 (Palawan).

Description of the species.-Characters of the genus. Color.-Uniform chocolate brown above, somewhat lighter brown below. Skull and teeth.-Skull short, facial portion especially abbreviated; zygomatic arches expanded; palatal foramina short, not extending behind posterior edge of canines; palate extending a considerable distance beyond molars posteriorly; palate with a V-shaped notch; palate extending less than half the entire length of skull; postorbital constriction marked. Incisors forming a. continuous, slightly curved line, separated by a considerable space f.rom canine; first premolar small, crowded out of line of teeth and frequently absent in adult specimens; second premolar usually somewha~ out of line of tooth row, less than half the diameter of third premolar; fourth premolar molari~orm; molar quadrate. Mea8urements of Aonyx cinerea (Illiger). mm.

Head and body Tail Hind foot Basal length of skull Zygomatic width Maxillary tooth row, alveoli

495 290 95 82 57.5 29

Remarks.-The measurements here given are those of Lyon for a specimen from Rhio Linga Archipelago. Lyon calls attention to considerable variation in the skulls of this species from various localities in the Malayan region. Family VIVERRIDlE Body usually elongate; tail variable in length; legs usually rather short; digits usually five but pollex or hallux or both 259453-23

354

PHILIPPINE LAND MAMMALS

may be wanting; dorsal vertebrre 13 or 14; auditory bullre externally constricted and divided by a septum; upper carnassial usually with an anterior lobe, lower one with a well-developed talon; second lower incisor raised above level of first and third. The family includes the palm civets and mongooses and has four recognized subfamilies; namely', Euplerinre, Viverrinre, Herpestinre, and Cryptoproctinre. Only the Viverrinre and the Herpestinre are represented in the Philippines. The family is confined to the Old World. Key to the Philippine subfamilie8 of Viverridm.

a'. A perineal scent gland; claws strongly curved and retractile. VlVERRINlE

(p. 354).

a '. No perineal scent gland; claws elongated and nonretractile. HERPESTINlE

(p. 364).

Subfamily VIVERRINLE

The genera Viverra, Paradoxurus, and Arctictis of this subfamily are represented in the Philippines. The first two are spread throughout the Islands, but A rctictis is, so far as is known, confined to the Palawan group in the Philippines. Key to the Philippine genera of Viverrinm. a', Ears with hair tufts, tail prehensile. b 1. Large, total length, 1,400 millimeters; tail as l(mg as head and body. Arctictis Temminck (p. 354). a'. Ears withou.t hair tufts i Itail" not prehensile. b 1. Medium, total length, 900 millimeters; tail half the length of head and body .......................__ ................... ........ V 'i verra Linnreus (p. 357). b '. Smaller, total length, 800 millimeters; tail about two-thirds the length of body.................................... Parado xwrus Cuvier (p. 360).

Genus ARCTICTIS Temrninck

Arctictis TEMMINCK, Prosp. de Monog. des Mamm. Palmer, Index Gen. Mamm. (1904) 117; HOLLISTER, Philip. Journ. Sci. ยง D 7 (191~) 23. Arctitis (err. typ.) ALLEN, Bull. Am. Mus. Nat. Hist. 28 (1910) 15.

Body elongate; head broad posteriorly, with a small pointed face; vibrissre long and numerous; ears small, rounded, bearing a tuft of long hairs; eyes small; fur on body coarse and harsh, elongate; tail elongate, prehensile; legs short, soles and palms broad and naked. Cervical vertebrre, 7; dorsal, 14; lumbar, 5; sacral, 3; caudal, 34.

355

ARCTIC TIS

Dental formula:

. 3

2 3' c. 1l' pm. 44' m. 2 = 40.

A single species of the "bear cat," Arctictis whitei Allen, is known from Palawan. The species reported from Palawan by Everett and Oustalet as A rctictis binturong. may very probably be referable to Allen's species. ARCTICTIS WHITEI

Allen

Arctictis whitei ALLEN, Bull. Am. Mus. Nat. Rist. 28 (1910) 15 (err.) . ? Arctictis binturong (part.) OUSTALET, Bull. du Mus. d'Rist. Nat. 7 (1901) 318-320; EVERETT, Proc. Zool. Soc. London (1889) 223 (Palawan); ELERA, Contr. Fauna Filipina (1915) 188, pl. unnumbered. Arctictis whitei HOLLIl:?TER, Philip. Journ. Sci. ยง D 7 (1912) 23; Proc. U. S. Nat. Mus. 46 (1913) 336. Viveria binturong RAFFLES, Trans. Linn. Soc. 12 :352.

Description of the species.-(After the type description.) Generally similar in character of pelage and general features of coloration to other forms of the genus, but differing in character of tooth structure. Pelage long and coarse, except on head, where it is short, especially on nose and facial region. Color.-General color black, washed with fulvous ; basal part of pelage deep black, hairs broadly tipped with fulvous, the fulvous tint forming the surface color on forelegs, and prevailing tint over greater part of body; face generally dark, the hairs narrowly tipped with whitish; all around neck and terminal fifth of tail black; outer surface of ears heavily tufted with black, with a narrow white or yellowish white border; ventral surface rather thinly haired, hairs crinkled and more or less woolly, blackish at base, broadly tipped with dull fulvous on chest and inside of forelegs; black on sides encroaches on thoracic region; abdominal region pale, silvery fulvous, continuous over anal region and along underside of tail as a broad mesial band for two-thirds length of tail; vibrissre on snout elongate, reaching ears, white or yellowish white; mammre four, abdominal. Skull and teet h.-Skull distinctly fiat on top, decidedly different from Arctictis binturong in which the skull rises rapidly from proximal third of nasals to its highest point behind orbital region.

356

PHILIPPINE LAND MAMMALS

The character of the fourth upper molar differs in A. binturong from this species; the tooth is triangular with a well-developed anterointernal cusp, and the first upper molar has the inner 118

120'

PHILIPPINE ISLANDS

.122 Batan Is

11'4

126

® Ar>ctidis whitei @ Viver>r>a tangalunga © Par>adoxur>us minax @ Par>adoxurus philippinensis ® Par>adoxurus tor>vus ® Mungos palawanus @ Mungos paflvus

'\,

LEYTE ~Oinagat

~ ~Siapgaol".l ~ I7Suca " t»

10'

;t.

"'Cagayan Sulu

122

FIG. 16.

124'

126

Distrihut,ion of Viverridre in the Pb.ilippines.

moiety much narrower than the outer; in A. whitei the teeth are obtusely oval instead of triangular in outline, and the inner half of the tooth is less reduced and less oblique to the axis of the tooth. The formula for teeth is same as that of the genus.

357

VIVERRA

Measurements of the types of A?'ctictis whitei Allen. Type. mm.

1,310 Total length 700 Head and body 610 Tail vertebra! 100 Hind foot, without claws 107 Hind foot, with claws Skull: 75 Zygomatic breadth 27.5 Interorbital breadth 32 Postorbital breadth 40 Across postorbital processes 27.5 Length of nasals 14 Breadth of nasals at front border 67.5 Palatal length Palatal breadth, outside first upper molar 36.5 16 Length of canine above alveolus 40 Upper tooth row, canine to second molar Breadth of incisors, alveolar border 15

Paratype. mm.

28 34.3 41 27 14 65 36 19

Remarks.-I have examined the types and paratypes, in the American Museum of Natural History, which consist of the type, a female specimen, from Iwahig, Palawan, collected by John R. White; a paratype, also a female (both with occipital skull region nrissing) ; a male skin without skull, and a flat imperfect skin. The species was named for John R. White, formerly colonel in the Philippine Constabulary and in charge of Iwahig Penal Colony. Genus VIVERRA Linnmus

Viverra LINNlEUS, Syst. Nat. 10 ed. 1 (1758) 4'3.

Largest species of the subfamily Viverrime. Fur long and loose, usually elongated in median line of body, forming a low crest or mane; neck with one or two black rings; pupil circular when contracted; perineal glands greatly developed; teeth stout, less compressed than in other genera, second llPper molar larger; audital bullre relatively small and pointed in front; tail much less than half length of body; claws long, not completely retractile. Civet, a secretion fr<1In the perineal glands in the species of this genus, is used in making perfumes. One species, Viverra tangalunga Gray, occurs in the Philippines. It occurs also in Malacca, Sumatra, Borneo, Celebes, and Amboyna.

358

PHILIPPINE LAND MAMMALS

Elera * lists Viverricula malacensis from Cagayan, Luzon, and V (iverra?) zibetha from Paragua, Negros, and Panay. These species probably do not occur in the Philippines. VIVERRA TANGALUNGA Gray Viverra tangalunga GRAY, Proc. Zool. SOC. London (1832) 63; (1864) 512; Cat. Mamm. Brit. Mus. 48; THOMAS, Trans. Zool. Soc. London 14 (1898) 388 (Luzon and Palawan); MEYER, Abhandl. und Berichte Konigl. Zool. Mus. Dresden 7 (1899) 20; BOURNS and WORCESTER, Occ. Pap. Minnesota Acad. Nat. Sci. 1 (1894) 1-64 (Calamianes, Mindanao, Mindoro, Panay, and Siquijor); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 22; Proc. U. S. Nat. Mus. 46 (1913) 312 (Luzon, Lubang, Palawan, Mindanao); ALLEN, Bull. Am. Mus. Nat. Hist. 28 (1910) 15 (Palawan).

Description of the species.-(From No.4, Bureau of Science collection; collected February 13, 1902, by R. C. McGregor and A. Celestino, near Mariveles, Bataan Province, Luzon.) Head pointed anteriorly, snout naked, truncate; vibrissre on side of head not reaching tip of ears; a second group of vibrissre arising behind angle of mouth below ear; a few elongate vibrissre arising from above eye; no crest or mane present; ear small, rounded above, but distinctly emarginate on external edge; fur on neck, sides, and back 20 to 24 millimeters long, on belly about 18 millimeters; on median line of back, hairs about 30 millimeters long; legs rather short. Color.-General color above a mixture of grayish white and blackish brown; face grizzled, darker on snout and on forehead; lighter above eyes and along mouth; eye encircled with a black rim; ear grayish brown behind near tip, below this a darker spot; side of neck with three black lines, the upper continued toward foreleg; the two lower, on each side, cross throat and meet their fellows from the opposite side; black lines separated by grayish to white interspaces; white and dark bands in strong contrast on undersurface of neck; chin dark brown, becoming gradually grayer posteriorly until first black band is reached; neck above reticulated with black-brown, a black stripe 1.5 centimeters wide from shoulders to end of tail; two or three rather indistinct dark stripes separated by lighter interspaces on posterior part of body; sides mottled and reticulated with irregular spots; all legs brown, with dull blackish markings; most of belly

* Contr.

Fauna Filipina (1915) 173-179.

359

VIVERRA

hair brownish except for whitish tips; belly also mottled with darker; about anal opening hair brilliant reddish brown in strong contrast; tail blackish above, with lighter bands below. Skull and teeth.-Skull generally slender, arches greatly expanded; a slight depression on nasals; postorbital and jugal processes not or scarcely visible; interorbital constriction less than constriction in postorbital region, a distinct sagittal crest on posterior skull and strongly developed lambdoid crests; audital bullre large, diagonally placed, somewhat compressed; palate extends some distance behind last molar; palatal foramina extend slightly beyond posterior edge of canines; antorbital foramen small, somewhat rounded. Incisors forming a slightly curved row in contact or nearly so, separated from canines by a well-defined space much greater than diameter of outer incisor; canines long, curved; first premolar halfway between canine and second premolar; second and third teeth separated by an appreciable distance; three posterior teeth in contact with each other; first lower premolar slightly nearer second than cl'tnine. MeaSU1'ements of Viverra tangalunga Gray. mm.

TO!tal length of skin (stretched?) Tail Skull: Greatest length Zygomatic width, greatest Greatest depth Width of rostrum Interorbital constriction Postorbital constriction Width of brain case Length of palate from foramina Length of palatal foramina Length of audital bullre Upper tooth row including canine Mandible, incisors to condyle Lower tooth row including canine

690 310

112 62' 38 20 20

16 35 41 路

8.5 19 46.5

84 51

Remarks.-A specimen in my collection from Cotabato, Mindanao, has the darker spots and lines composed of light buff brown and blackish; facial markings similar but not so well defined; the bands on underside of tail almost encircle it. The

PHILI PPINE LAND MAMMALS

360

specim en does not have its compl ete adult dentit ion so it is impossib le to make a comp arison of the denta l chara cters. The specie s is wides pread . The type locali ty is "India n Archi pelago ." It is know n from all the larger island s of the Philip pines. A skin in my collection is from Poli1lo Island . Genus P ARAD OXUR US Cuvier HOLLISParado xurus F. CUYlER, Hist. Nat. Mamm . 3 (1824) livr. 24; 22. TER, Philip. Journ. Sci. ยง D 7 (1912)

Body rathe r elong ate; head pointe d in front; ears small, t round ed; legs not especi ally shorte ned, palms and soles almos a of ention entire ly naked , and joinin g footpa ds witho ut interv hairy space ; claws compl etely retrac tile; eye mode rate, pupil , vertic al; tail long, non prehe nsile. Cervic al verteb rre, 7; dorsal 13; lumba r, 7; sacral , 3; cauda l, 29 to 36 . 2 4 1 . . 3 Denta l formu la. 1. 3' c. l' pm. 4' m. 2 = 40. The denta l formu la is identi cal with that of Ardic tis. The genus is confined to south ern and south easter n Asia and the Malay Archi pelago . It occurs throu ghout the Philip pine Island s. Three specie s are curren tly recogn ized. Key to the Philipp ine species of Parado xury,s Cuvier.* a '. Smalle r. small spots. b '. Withou t bands on back and sides, marked with many P. philipp inensis Jourda n (p. 360). withou t lighter b '. Genera l color dark copper y brown ; face blackis h 94 millim eters. skull, lines; black dim three with back markin gs; P. torvus Thoma s (p. 361).

a 2 โ ข Larger . broken b'. Unifor m brown less mottle d than in P. philipp inensis ; three

uous; lines of dark spots on back; light frontal band not conspic skull, 105 millim eters. P. minax Thoma s (p. 362).

PARADOXURUS PHILIPPINENSIS Jourdan Paris Parado xurus philipp inensis JOURDAN, Compt . rend. Acad. Sci. 339; 41) (1835. 5 (1837) 523; TEMMINCK, Monog . de Mamm London Soc. Zoo!. Proc. OGLIBY, Zoo!. Journ. No. 15, 300; GRAY, (prob(1864) 537; GUNTHER, Proc. Zoo!. Soc. London (1879) 75 15 (1910) 28 Hist. Nat. Mus. Am. ably P. minax ); ALLEN~ Bull. Acad. ota Minnes Papers Oce. STER, WORCE (Palaw an); BOURNS and

) Contr. a la Fauna Filipin a (1915) 180-18 8, lists P(arad oxurus crOBxuTUS Parado India, and nigrifr ons Gray (var. from Luzon, Cagay an 8ii Gray var. Luzon, India, Borneo , probab ly erroneo usly.

* Elera,

PARADOXURUS

361

Nat. Sci. 1 (1894) 1-64 (Panay, Mindoro); THOMAS, Trans. Zool. Soc. London 14 (1898) 388 (North Luzon); STEERE, List of Birds and Mammals Steere Expd. (1890) 29 (Mindanao? Negros, Marinduque); TROUESSART, Cat. Mamm. Sup pI. (1904) 254'; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 22; Proc. U. S. Nat. Mus. 46 (1913) 313 (Luzon, Ticao, Mindoro, Palawan) ; ELERA, Contr. Fauna Filipina (1915) 182, pI.

Description of the species.-Body elongate; tail long, less than length of head and body; head triangular, nose pointed; vibrissre on snout reaches to ear; fur soft, consisting of a soft, frequently wavy underfur with longer, much coarser overfur; ears l'ather small, rounded, not rising much above horizontal line of top of head; legs rather short; claws retractile; scent glands well developed . . Color.-Above, somewhat yellowish brown, sometimes with a golden luster; back somewhat darker; head with a grayish band above eyes across forehead; breast grayish yellow; top of head and ' ears coffee-brown; tail uniformly brownish; feet and legs both inside and outside coffee-brown; a brownish to brownish black band present, dim in adults but fairly distinct in young. Skull, teeth, and measurements.-I have been unable to obtain a specimen of this species for examination. Remarks.-Temminck says, "Espece indiquee primitivement par M. Jourdan de Lyon, qu'il decrit sous Ie nom d' Ombliodon dore." However, the first reference in literature I have found is that of Jourdan, where he refers to the form as Paradoxure des Philippines, Paradoxurus philippinensis (nobis). The description is extremely short and the type is not mentioned. The type locality is "Rabite les iles Philippines Luzon et Mindanao." The species is not especially rare but is nocturnal in habit and rarely encountered. Elera figures two mounted albino specimens. I saw a very young uniformly blackish specimen of Paradoxurus in Tablas, in captivity. PARADOXURUS TORVUS Thomas

Po,1"o,doxurus torvus THOMAS, Ann. & Mag. Nat. Hist. 38 (1909) 376; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912') 23; Proc. U. S. Nat. Mus. 46 (1913) 336.

Description of the species.-Size about as in Paradoxurus philippinensis, with similar body characteristics. Color.-General color dark coppery brown, (between mummy brown and Vandyke brown), approaching that of the South Indian species P. jerdoni; face wholly blackish without lighter

362

PHILIPPINE LAND MAMMALS

markings; back of ears, nape of neck to groin, and rump also blackish; back with three, indistinct, inconspicuous black lines; legs and tail blackish brown; undersurface dark chocolate brown. Skull a,nd teeth.-Skull similar to that of P. philiJppinensis; teeth with a similar rounded character, larger individually than those of P. sa,ba,mt-S, the species most closely related geographically; posterior edge of palate of reversed V-shape found in the genus Pa,g~~ma" although it is about the same distance behind molars as is usual in Pa,ra,doxurus. (From the type description.) Mea,surements.-See under Para,doxurm minax Thomas. Rema,rks.-The type locality is Bungau Island, Tawitawi Group, Sulu, P. 1. The type was collected by A. Everett, July, 1893. Thomas remarks: Even if the peculiar coppery brown color of this Paradoxure should prove to be due in any degree to a melanoid suffusion (and there is no evidence of this suggestionJ the continuation of the dark color of the head down to the withers, the large size of the teeth compared with P. sabanus and the Paguma-like snape of the posterior nares will distinguish the species from any of its a~lies,

The species probably also occurs on Tawitawi, since Bungau is separated from the southern point of this island by only a few hundred meters. I failed to 0btain any mammals in Bungau. PARADOXURUS MINAX Thomas

Paradoxurus min ax THOMAS, Ann. & Mag. Nat. Rist. III 8 (1909) 375; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 22; Proc. U. S. Nat. Mus. 46 (1913) 313 (Mindanao, Baganga, Catagan, Mount Malindang, Mount Apo, and Zamboanga). Paradox7,wus zelanicus GRAY, List of Mamm. Brit. Mus. (1843) 55 (nomen nudum) (Camiguin Island, Mindanao) .

Description of the species.-(After the type description.) Markedly larger than P. philippinensis; about equalling P. niger. Color.-General color un~form dark brown, less mottled than in P. philippinensis; three broken lines formed of black spots fairly well defined on the back; light frontal band inconspicuous; crown and back of ears blackish brown; legs, feet, and tail uniformly dark. Skull a,nd teeth.-Skull similar to that of P. philippinensis but larger; teeth of the same general rounded shape, the carnassial with a heavy posterointernal ledge.

363

PARADOXURUS

Measu?'ements of the types of Parado'Xurus minax Thomas anw p, torV'l.48 Thomas. PtDradozurutl minaz.

Paradozurus torvus .

mm.

580 Head and body Tail (in a specimen other than type) • 440 75 Hind foot Skull: 98 Basal length 105 Condylobasal length 60 Greatest breadth 19.5 Length of nasals in middle line 19 Interorbital breadth 34 Breadth of brain case 5.5 Palatal foramina 47 Palatal length 8.4 Length (outer edge) of fourth premolar 9.6 Greatest diagonal diameter (same) • Not the type.

Type .

490 410 64 ·39 94 57.5 14' 15.5 33.5 5 42 8 8.7

Sic.

Remarks.-The type, an adult male, was collected by M. P. Anderson, in Davao, Mindanao. Thomas refers Gray's Paradoxurus zeylarllicus from Camiguin Island, Mindanao, to this form. Hollister says: * The material representing this genus from the Philippines is far too little to work out the forms satisfactorily. The characters given by Thomas for his P. rwinax are mostly untrustworthy, as shown by our material, limited as it is. The Mindanao animal does not seem to differ in size, or size of skull from the Luzon form. It does average darker in color, so far as shown by the few available skins, and doubtless will prove to be a recognizable race.

A small specimen in my collection, brought to me alive at my hotel in Zamboanga, has the following coloration: Body above generally buckthorn brown (Ridgway) with irregular darker markings (almost a reticulation) on back and rump; sides lighter; the color results from the fact that tips of overfur for about one-half of its length are buff, extreme tip black, basal part of overfur and underfur gray; head blackish brown with an indefinite frontal band passing in front of ears, coming to a point between eyes; numerous white hairs forming a spot below eye; vibrissre blackish with white tips; hands and feet dark blackish brown; underside of head, neck, and breast much lighter than sides, buff color absent, longer hairs silvery white at ends;

* Proc.

U. S. Nat. Mus. 46 (1913).

364

PHILIPPINE LAND MAMMALS

extreme tips scarcely darker than outer half; basal half of tail with grizzled, wavy underfur with longer dark blackish overfur; djstal half much darker. The skull, which measures 61 millimeters in total length, shows the permanent teeth emerging from the gums. It is probable that Gunther's * reference to Paradoxurus philippinensis, North Mindanao, should be P. minax Thomas. Subfamily HERPESTIN.JE

Auditory bullre very prominent, pear-shaped, posterior part large, rounded, and generally with its greatest prominence on outer side; toes straight, claws lengthened, exserted, nonretractile; no perineal glands; orbit may be surrounded by bone; anus usually opens in a sa-dike depression. Mungos is the only genus of the subfamily occurring in the Philippines. Genus MUNGOS Geoffroy and Cuvier

Mungos GEOFFROY and CUVIER, Mag. Encycl. 2 (1795) 184. Herpestes ILLIGER, Prodr. Syst. Mamm. (1811) 135.

Head pointed in front; ears short, rounded; body long and slender; legs short; five toes on each foot, the first, especially that of hind foot, very short; toes free or but slightly palmated; palms generally naked; distal portion of soles naked, undersurface of tarsus and metatarsus usually clothed with hair, but considerable variation obtains; tail long or moderate, generally thick at base and sometimes covered with more or less elongated hair; longer hairs on body and tail almost always annulated; teeth generally well developed, with sharply pointed cusps; skull elongated, constricted behind orbits; face short, compressed; frontal region broad and arched; postorbital processes of frontal and jugal bones well developed, generally meeting so as to complete circle of orbit. Cervical vertebrre, 7; dorsal, 13; lumbar, 7; sacral, 3; caudal, 21 to 26 . . 3 1 4 3 2 Dental formula: 1. 3' c. l' pm. 4 (or 3)' m. 2 = 40 or 36. The mongoose is widely distributed and species occur from Africa to Asia and the Malay Archipelago. In the Philippines the genus is confined to Palawan, the Calamianes, and adjacent islands. Two species, Mungos pa[,a"wanus Allen and M. parvus (Jentink), have been described. Whether the two forms are specifically distinct I cannot say. Allen described M. palawanus

* Proc.

Zoo!. Soc. London (1879) 75.

365

MUNGOS

from a skin only and had not seen the skull. It appears that th~ description of M. parvus was unknown to him since he states: "The mongoose so long known from the Philippines appears to have always been referred to Herpestes brachyurus Gray, the type locality of which is Malacca." Apparently only the type specimen of each is known. MUNGOS P ALAW ANUS Allen

Mung08 pala,wa,nus ALLEN, Bull. Am. Mus. Nat. Hist. 28 (1910) 17 (Palawan); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 23; Proc. U. S. Nat. Mus. 46 (1913) 336.

Description of the species.-Related to Herpestes brachyurus Gray.

Color.-Color above generally yellowish red punctuated with black; head much paler and grayer than body; ventral surface reddish brown, hairs slightly tipped with yellow; forelegs dusky; hind legs like sides and back; individual hairs of dorsal surface reddish, annulated subapically with black and tipped with yellow, the black annulations more or less visible at surface. Skull and teeth.-Characters unknown. Measurements of Mungos pa,la,wa,nus Allen. mm.

Length of head and body Tail vertebrre Hind foot without claws Hind foot with claws

395 190 50 56

Remarks.-Allen states: A comparison of the present specimen with Gray's description and figure leaves little room for doubt that the Palawan form is a quite different animal from the true M. bra,chyu?'Us of Malacca. It is also so different from the short-tailed mongoose from Sumatra, commonly referred to M. b?'achyurus, as to need no minute comparison with it, the latter beIng a much larger and a much darker animal.

The type, in the American Museum of Natural History, is an a.dult male, collected at Iwahig, Palawan, by John R. White. Only the type is known. * MUNGOS PARVUS (Jentink)

Herpestes parv'Us JENTINK, Notes Leyden Mus. 17 (1895) 4'8, Mungos parvus HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 23; Proc. U. S. Nat. Mus. 46 (1913) 336.

* Everett, Proc. Zool. Soc. London (1889) 223, lists H. b?'achyurus from the Palawan group of island~.

366

PHILIPPINE LAND MAMMALS

. Description of the species.-About the size of an adult Mungos brachyurus but differs in having a short tail. Color.-Lighter colored than Mungos brachyurus; extremities not black; light rings, producing grizzled appearance, reddish brown; tail less hairy, hairs brownish black. Skull and teeth.-The type is very old and many of the teeth are wanting. Measurement of the skull of Mungos parvus (Jentink). mm.

Greatest length Breadth of brain case Length of upper molar series Length of lower molar series

16 31

25 28

Remarks.-The type was collected by Schadenberg in the Calamian Islands (probably in Busuanga). See preceding remarks under the generic description. The type is in the Leyden Museum. Order RODENTIA

The order Rodentia is distinguished by the presence of large curved incisors, above and below, and the total absence of canines. Most of the living forms are small; dentition diphyodont, usually with only one premolar or none; number of molars variable; feet plantigrade, or semiplantigrade, generally with five digits, usually unguicula;te; clavicles present as a rule, but may be imperfect or rudimentary. Orbits communicating freely with temporal fossre; intestine with crecum; testes inguinal or abdominal; cerebral hemispheres not covering cerebellum. The order is represented in the Philippines by the Sciuridre, Petauristidre, Muridre, and Hystricidre. Key to the Philippine families of Rodentia. a'. Arboreal or terrestrial squirrels; premolars 7i; cylindrical hairy tails, or flattened tails. b 1. No membrane between fore and hind legs for flight. SCIURIDlE (p. 367).

b', A well-developed membrane between fore and hind legs adapted for sailing through the air (flying squirrels). PETAURISTIDlE (p. 383). a 2. Arboreal, terrestrial, fossorial, and natatorial rats; no premolars present; tail round, usually nearly naked and scaly ____ MURIDlE (p. 390). a路. Body covered with very strong elongate spines, with occasional long stiffened hairs; premolars present; (porcupines). H YSTRICIDlE (p. 489).

SCIURIDJE

367

Family SCIURIDJE The family Sciuridre comprises the true squirrels, the ground squirrels, the pigmy squirrels, and' the marmots. The flying squirrels belong to the family Petauristidre. The group consists of arboreal, terrestrial, or burrowing forms with: cylindrical tails without scales; body without lateral membranes; skull with distinct postorbital processes and small infraorbital foramen; palate only moderately broad; ears moderate or small, sometimes tufted; two upper premolar~ usually present (absent in young and in Eosciurus) , sometimes deciduous; molars rooted, tubercular. . 1-1 2-2 (1-1) 3-3 Dental formula: 1. 1-1' pm. 1-1 or 1-1 ,m. 3-3 = 22 or 20. The subfamilies Nannosciurinre and Sciurinre are known in the Philippines. The N annosciurinre, represented by a sIngle genus, are confined to Samar, Leyte, Mindanao, and Basilan. The Sciurinre are found in the same islands but also occur in the islands of the Palawan group. The question as to the derivation of the species of squirrels is an unsettled one, particularly as regards those species in the Eastern part of the Philippine Archipelago. Species in Palawan have undoubtedly been received from the Bornean fauna. Celebes has at least seven wen-defined species; namely, Sciurus 1nurinus Muller and Schlegel, S. rubriventer .Muller and Schlegel, S. lenconus Muller and Schlegel, and S. prevosti Desmarest, in North Celebes; S. rosenbergi J entink, in Sangi; S. weberi J entink, in Central Celebes; S. notatus Boddaert, in South Celebes. There are seven species recognized in the Philippines. Meyer * states that S. philippinensis is regarded by Jentink as being closely allied to the Malayan and Bornean species S. tem('is Horsfield; and that the Philippine species S. steerii is similar to the widely spread S. exilis Muller and Schlegel. Meyer remarks on his S. cagsi= (S. mindanensis) : ' The general coloration of this Squirrel is similar to that of the widespread Sc. tenuis Horsf. but that species is only half its size and has no white on the tail. Sr.:. ca,gsi also reminds one of Sc. leucomus M. & Sohl., from Celebes, in its general coloration; but the last named species has white behind the ears, which are long-haired, and reddish underparts, and offers, besides, other differences.

Borneo has a number of species including S. giganteus McClelland. Four of the seven species of Sciurinre known from the

* Meyer,

Proc. Zool. Soc. London (1890) 600.

368

PHILIPPINE LAND MAMMALS

Philippines are confined to the Palawan group, and three to the islands of the southern part of the eastern group. Key to the Philippine subfamtilies of SciuridaJ. a t. Small squirrels about size of small rats; head and body under 125 millimeters ................................ NANNOSCIURIN.iE Trouessart (p. 368).

a'. Larger squirrels; head and body over 200 millimeters. SCIURIN.iE Linnreus (p. 375). Subfamily NANNOSCrURIN..tE

N annosciur,inlÂŁ FORSYTH-MAJOR, Pro c .. Zool. Soc. London (1899) 189.

Forsyth-Major has separated the genus Nannosciurus from the Sciurinre and placed it in a subfamily on the basis of the characters of the skull and teeth, as listed in the generic description. Genus NANNOSCIURUS Trouessart

Nannosciurus TROUESSART, Le Naturaliste 2 (1880) 292 (Oct. 1. No. 37) (subgenus); Calt. Mamm. 1 (1898-99) 445 (genus); Cat. Mamm., Suppl. (1904-05) 344 ; FORSYTH-MAJOR, Proc. Zool. Soc. London (1893) 189 (genus); MOLLISTER, Philip. Journ. Sci. Â§ D 7 (1912) 24; Proc. U. S. Nat. Mus. 46 (1913) 313.

Small squirrellike for ms with moderately long flattened tails; fur soft, short, without spines; forefoot with four well-developed digits, vestigial pollex scarcely distinguishable; hind foot with well-developed toes. Skull with a small postorbital process, and a small process arising from zygomatic arch; anterior zygomatic root a perfectly vertical narrow septum and placed so far backwards that the facial part of cranium appears very elongate while the frontal region is broader than in other squirrel genera; infraorbital foramen small, vertical; palate moderately broad; orbits formed by a circular bony ring. A small anterior premolar appears to be present in adults of all species of the genus, although absent in the juvenile type of Nannosciur us concinnus; second premolar much smaller than first molars; pattern of crown of molars usually shows three complete transverse crests in both upper and lower molars; transverse valleys in teeth comparatively very deep . . 1-1 2-2 3- 3 Dental formula: 1. 1-1' p. 1-1' m. 3- 3 = 22. Key to the Philippine species of Nannosciurus. a

t. Molar teeth with three continuous ridges across surface. b 1. Color rather reddish brown to ochraceous tawny, variegated. N. concinnu.s Thomas (p. 369).

NANNOSCIURUS

369

b %. Less reddish, with a somewhat longer skull than N. concinnus. N. surrutilus Hollister (p. 370). b B. More grayish to gray-olive, grizzled .... N. samaricus Thomas (p. 372). a'. Middle ridge on upper molar teeth broken near middle forming two unequal cusps ........................................ N. luncefordi sp. nov. (p. 373). NANNOSCIURUS CONCINNUS (Thomas)

Sciurus concinnus THOMAS, Ann. & Mag. Nat. Hist. VI 2 (1888) 407 (Basilan) . Nannosciurus concinnus FORSYTH-MAJOR, Proc. Zool. Soc. London (1893) 181-189, pI. 10, fig. 10; pI. Ii, fig. 7; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 24; Proc. U. S. Nat. Mus. 46 (1913) 313.

Description of the species.-(From No. 45, E. H. Taylor collection; collected at Abung-abung, Basilan, October 20, 1920, by E. H. Taylor.) Adult male. Small; forefoot elongate, fur on heel extending to within a short distance of base of 'posterior tubercle; ears small, rounded in general outline; whiskers extending beyond ears; posterior pads on forefeet large, close together, the inner involving the vestigial pollex; posterior sole pad oval and situated farther back on foot than in other species, distance between posterior pad and middle digital pad being 6 to 7 millimeters; fur short, dense, bicolored, that on back and sides 4 to 5 millimeters long, a few long hairs extending as much as 7 to 8.5 millimeters; fur on belly about 6 to 7 millimeters, on tail, 13 to 15; hair on foot somewhat ochraceoustawny and hairs about base of toes buff; ears without tufts, haired both inside and outside. Color.-General color of body and head rusty rufous; basal part of fur bluish gray to slate; rufous color on outer third of hair, and outer end tipped with black; many hairs with alternating bands of black and dull rufous; belly only slightly lighter; tail dull rufous with indefinite darker areas; hairs banded on outer end but rufous toward base. Skull and teeth.-Skull broadened, with short muzzle; greatest breadth of the two nasals more than half their , length; postorbital processes distinct; angle formed by orbital ring with skull nearly a right angle. Anterior premolar especially small, its grinding surface approximately one-fifth that of second premolar; width of upper molars and premolars distinctly greater than length; first two molars with prominent transverse median ridges; not so well defined in other cheek teeth; mandibular teeth not or scarcely wider than long; upper tooth rows distinctly diverging anteriorly; distance between posterior molars, 2.6 millimeters; between anterior premolars, 3.8. 269463-24

370

PHILIPPINE LAND MAMMALS

Measurements of Nannosciurus concinnus Thomas. Type

mm.

Head and body Tail, without hairs Hind foot Heel to front of last foot pad Heel to front of anterior digital pad Ear, contracted Skull: Basal length Greatest length Greatest breadth of brain case Greatest zygomatic width Nasals, length Nasals, width of both Diastema between incisors and molars Length of tooth series Least interorbital width Greatest length of mandible, including incisor Mandibular tooth row

E. H. Taylor collection No. 45.

mm.

85 59 25.5 11.4 18.4 4.8

86 59 25.1 11.5 17.8 5.0

19.6

20 27 15.4 17.5 7.0 3.8 6.0 4.1 10

15 7.1 3.9 6.3 3.8

18.6 4.2

Remarks.-The speeimen here described (No. 45) differs from the type chiefly in the presence of the small upper premolar; the tooth is very minute having a diameter of only about 0.38 millimeter and a surface area of only about one-fourth to one-fifth of the second premolar; there is a single premolar in the mandibular series. The slight difference in the length of the upper tooth row from that of the type is due to the presence of the small premolar. ,My specimen Was shot running along the trunk of a fallen tree in a dense forest in southern Basilan, at a point approximately 25 kilometers directly south of Isabela, the type locality. The specimen consists of a dry skin (hair on tip of tail missing) and skull. This was the only specimen observed in one and one-half months collecting on the island. NANNOSCIURUS SURRUTILUS Hollister

Nannosciurus 8UrrutJiJ,US HOLLISTER, Proc. U. S. Nait. Mus. 46 (1913) 313; (Mt. Bliss, Mindanao).

Description of the species.-(From the type description.) Like Nannosciurus concinnus of Basilan, but darker, less reddish; with a longer and slenderer skull. Color.-General color of upper parts dark reddish brown, finely marked with black; much darker and less reddish than in N. concinnus; the underfur slate-black; terminal half of hairs

371

NANNOSCIURUS

ringed with black and ochraceous-tawny, and tipped with black; cheeks and sides more buffy, less reddish; underparts with underfur deep neutral gray, the hairs tipped with buckthorn brown; hands and feet like back but brighter, more reddish 118

122 Batan I.

120

PHILIPPINE ISLANDS

126

@ NannosciupU5 concinnus

® Nannosciurus surruti Ius © Nannosciurus samaricus @

Nannosciurus funcefordi

® Sciuropterus nigripes nigripes ® Sciuropterus n. efassodontus I

Sciuroptel'U5 crinitu5

Lubaog'~"

\:i.

MINQORO Busuanga ~

'«'

guque ~~

;

J!tI"

6up; •• ,

f7bRomblon~f; ..

lab1f S;~yan

"; J~::i~'f:l?}:;'t,;,t I<aO

"'J' ,.g.". ,. 'NE"OS~

~PALAWAN

MINOAN"

S' BalaOac

®@ B "

IZ'

tr~

-®

DCatanduanes "-

.(J

"Cagayan Sulu 6'

0' 120

FIG. 17.

122

124"

126"

Distribution of N atn/nosciUTU8 and S ciUToPte'1'U8 in the Philippines.

orange; tail above and below mixed black and Sudan brown, with more black at tip. Skull and teeth.-Skull longer and slenderer than that of N. concinnus, nasals and rostrum narrower and brain case elongated; audital bullre slightly larger; teeth as in N. concinnus.

372

PHILIPPINE LAND MAMMALS

MeasU1'ements of the type of Nannosciurus swrrutilus Hollister. mm.

Total length Tail vertebrre Tail to end of hairs Hind foot Skull: Greatest length Condylobasal length Zygomatic breadth Interorbital breadth Maxillary tooth row Mandible Mandibular tooth row

164 77 113 25

27.7 24.9 16.8 10.2 4.2 16.4 4

Remarks.-The type, in the United States National Museum, together with three paratypes, was collected on Mount Bliss and Mount Malindang, northern Mindanao, May, 1906, by Edgar A. Mearns. This species differs widely from N. samaricus Thomas in color markings. NANNOSCIURUS SAMARICUS Thomas

Nannosciurus samaricus THOMAS, Minutes Proc. Zool. Soc. (June 15, 1897); Trans. Zool. Soc. London 14 (1898) 389, pJ. 30, fig. 2; HOLPhilip. JOUTn. Sci. ยง D 7 (1912) 24; Proc. U. S. Nat. Mus. 46 (1913) 313.

LISTER,

Description of the species.-(From the type description.) Size and characters much as in Nannosciurus concinnus Thomas; fur much shorter and more velvetlike, hairs about 5 millimeters long on back; undersurface rather thinly haired; character of sole pads apparently much as in N. concinnus,' tail very similar, but rufous rings on hairs less developed and black rings more pronounced. . Color.-Grayer and less rufous than N. concinnus: general color of head and body finely grizzled olive-gray, with only a faint tinge of rufous on back; undersurface dirty grayish not defined on sides; legs dusky, upper sides of hands and feet dusky grizzled gray; a few orange-tipped hairs on digits. Skull and teeth.-Skull very similar to that of Nannoscinrus concinnus, but nasals somewhat narrower; two premolars normally present, the anterior minute, styliform, circular, the posterior considerably larger but still smaller than first upper molar; molars all much more rounded than in N. concinnus, their transverse diameter scarcely exceeding their longitudinal diameter.

373

NANNOSCIURUS

Measurements of the type of Nannosciurus scvmaricus Thomas. mm.

Head and bo.dy Tail, witho.ut hair Tail, with hair Hind fo.o.t, mo.istened Skull: Greatest breadth Nasal length Nasal breadth Intero.rbital breadth Tip to. tip o.f po.sto.rbital pro.ceslSes Diastema Length o.f cheek teeth series, with last premo.lar Length o.f three mo.lars Lo.wer jaw, co.ndyle to. inciso.r tip Bo.ne o.nly

88 69 94 25.2 16.2 7.7 3.1 10 12.6 6 4.1 3 18.2 15.5

Remarks.-The type of the species, an adult female, was collected by J. Whitehead in Samar, June 30, 1896. It is now in the British Museum. Thomas gives a very good color plate of the species, and remarks that this form is probably only the representative of Nannosciurus concinnus in Samar, since it seems probable that there are two premolars in N. concinnus in the adult. He described the original specimen as having only one premolar, but finds that this single one is a milk premolar. The specimen of that species which I describe in this paper proves that Thomas's conclusion is correct. NANNOSCIURUS LUNCEFORD I sp. nGv.

Type.-No. 594, E. H. Taylor collection; collected at Saub, Cotabato, Mindanao, at sea level, April 29, 1923, by E. H. Taylor. Adult male; dry skin with skull. Diagnosis; description oj the type.-Related to Nannosciurus concinnus and surrutilus but apparently differing from both in numerous characters. Hair on forehead, about 4 millimeters long; on neck above, 5; longest hairs, about 7; on middle of back, 6; longest hairs, about 8 to 9; Iowan sides, 8 to 9; on sides of base of tail, 121 to 14; near tip, 14 to 15. (Tip of tail missing.) Ear apparently larger and more flattened than in N. concinnus; tail distinctly less bushy, fur softer than in concinnus; a few white or white-tipped hairs about mouth; shorter vibrissre also tipped with white. Color.-Reddish brown on crown, back, and sides, the bulk of the hairs bicolored, deep slaty black on proximal part, distal

374

PHILIPPINE LAND MAMMALS

part with a ring of ochraceous-tawny to rufous, varying in width, and extreme tip black; a few of the longer hairs with a narrow tawny band about middle, outer half of hair black; a few of the longer hairs blackish for their entire length; low on sides and on belly the outer part of fur dull wood brown to tawny, underfur dull dark gray-slate; hair on tail a~ extreme base mixed, the darker predominating, followed by a ring of ochraceous-tawny; then a broad black band, tip usually reddish tawny; underside of tail appears striped (the upper part somewhat less so), with three stripes on each side, the median stripe widest, black. Skull and teeth.-Skull rather stout; nasals distinctly broader than in other species; region above and anterior to orbits distinctly inflated; postorbital process very small, with a small spur rising from zygomatic arch; skull somewhat less deflected, and bullre distinctly larger than in other Philippine forms; teeth with surface nearly horizontal, somewhat concave, directed outward and downward; transverse ridges not continuous at equal height and thickness across tooth in two first molars, but median ridge higher and thicker internally and separated from outer portion of ridge by a ' deep depression; a second ridge rising from outer posterior part of tooth goes nearly halfway across tooth making a V -shaped figure with outer portion of median ridge. Thus, in outer profi]e, the tooth row shows the following cusp series: Second premolar, a large anterior, and a small posterior cusp; first molar, a very small cusp followed by a medium-sized cusp, separated by a deep groove From the other larger cusp; second molar with small anterior cusp still more reduced, the two following, equal, separated by a deep groove. In vertical profile the molars show notches on outer side; first premolar very small, styliform, not unlike that in N. con(Jinnus; all teeth slightly larger than in N. concinnus; lower teeth rather similar to N. concinnus. Measurements of Nannosciurus luncefQ1路di sp. nov. E- H. Taylor collection No. 594.

mm.

Head and body Tail (tip missing) Hind foot Heel to front of last footpad Heel 1\;0 front of anterior digital pad Ear

82 50 25.2 11

18 6.3

375

SCIURUS

Measurements of Nannosciurus lunceford!i sp. nov.-Continued. E. H. Taylor colleqtion No. 594.

mm.

Skull: Basal length Greatest length Greatest width of brain case Zygomatic width Nasals, length Nasals, width of both Diastema Upper tooth series Orbit Interorbital widJth Mandible with incisor Mandibular bone Mandibular tooth row

22 26

14 16.6 7 4 6.2 3.9

5.6 10.2 18.1 16.3 4

Remarks.-Only a single specimen was captured, although others were glimpsed in the forest. The characters of the teeth seem to separate this from the other Philippine species. The species is named ,for my frieNd Albert Lunceford, jr., of Ottawa, Kap.sas, who has accompanied me on many collecting trips and collected many valuable specimens for me. Subfamily SCIURIN.IE

This group includes the true squirrels, the ground squirrels, and the marmots. They are characterized by the absence of wing membranes and are of moderate size. Forsyth-Major, * who reviewed the group in 1893, reduced the genera to six, but listed some eight subgenera. Only the genus Sciurus is recorded from the Philippines, with seven named forms. Genus SCIURUS Linnreus

Key to the Philippine species of Sciu'f'Us. a 1. Body generally reddish chestnut, grizzled, witl,l a few longer black bristles; belly uniformly reddish chestnut or reddish buff. S. steerii Gunther (p. 376). a 2. Similar to S. steerii but sides and shoulders grayer rih an admixture of whitish hair; belly either chestnut-red or white. , S. juvencus Thomas (p. 379). a 3. Similar, but with white on tail tip ........ S. albicauda Matschie (p. 379). a'. Varying from preceding form only slightly, with a white line under base of tail ........................................ S. mollendorffi Matschie (p. 380).

* Proc.

Zoo!. Soc. London (1893) 179.

376

PHILIPPINE LAND MAMMALS

a'. Upper parts blackish, variegated or grizzled with yellowish brown. S. mindanensis Steere (p. 381). a'. General upper color deep rieh brown grizzled; under parts grayish white; head and forelegs grayish.

S. philippinensis Waterhouse (p. 382). a '. Head colored like back; larger; feet grayish. S. samarensis Steere (p. 382). SCIURUS STEERII Giinther

steer;ii GUNTHER, Proc. Zool. Soc. London (1876) 735, pI. 69 (part.) (Balabac); THOMAS, Ann. & Mag. Nat. Hist. 2 -(1908) 498 (Balabac); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 25; , Proc. U. S. Nat. Mus. 46 (1913); FORSYTH-MAJOR, Proc. Zool. Soc. London (1893) 189 (part.).

Sciu1"'uS

-Description of the species.-(Fro.m No.. 21, E. H. Taylor collection; collected at Sir J. Brooke Point, Palawan, September, 1923, by E. H. Taylor.) Adult male. Hair on median anterior part of muzzle forming a low distinct roach; ear small, rather flattened against head, covered with long haIr behind and short hair over anterior surface; fur composed of well-defined, woolly underfur and longer straight overfur; tail rather small, more flattened than bushy, hair at tip elongate (longest hairs, 53 millimeters) ; fur on upper neck, 10 to 15 millimeters long; on middle of back, 15 to 18; longest hair on sides of tail, 28; hair on belly uneven, about 7 to 9; o.n sides of tail, 23. Color.-Upper parts of body, sides and outer side of legs rather grizzled, reddish, chestnut brown, lower part of hairs grayish on anterio.r part of body and mo.re slaty black on back and rump; each hair with a broad band of reddish chestnut and tipped with gray to slate black, the black terminal band wider along median part of back and rump; entire undersurface of body and limbs bright orange-buff to. brownish buff, base of hairs very light gray; vibrissre entirely black; tail more reddish than back, some of the hairs with black bands arranged so. as to. form indistinct blackish bands around tail; proximal part of hair reddish chestnut; a few hairs at tip of tail with bro.ad blackish band near reddish base, but entire outer half reddish chestnut. Skull and teeth.-Facial portion of skull moderately short; a strong postorbital process and a slight lachrymal process; infraorbital foramen large, vertical, with a bony proj ection external to it; palate with a small median projection; bullre large, flattened. . The skull is apparently that of a young animal since the upper premolar present appears to be a milk tooth; small premolar vlhich scarcely rises above the surface of the palate pro.bably

377

SCIURUS

represents the first premolar; the second permanent premolar can be seen below the present tooth; the first premolar probably is absent in the adult; the teeth with exception of the last molar are built on the same plan. The folds rise on the outer 118

120

I 4

122

1<16

Batan I.

PHILIPPINE ISLANDS

® Celcenomys silaceus <Ii)

Rhynchomys sOl'icoides

Chpo!omys whiteheadi

@ Crunomys fallax

® Cpunomys melanius ® Sciupus albicauda «V Sciupus mindanensis ® Sciupus mollendopffi CD Sciupus Q) Sciupus ® Sciupus <D Sciupus

juvencus steepii samapensis philippinensis

< "

"Cagey.. n Sulu

122

FIG. 18.

124'

126

Distribution of Celamomys, Rhynchomy8, Chrotomys, Cru/T/,OmYB, and Sciurva in the Philippines.

edge of teeth to form three cusps, the anterior in each case greatly reduced; surface of tooth traversed by three valleys; inner valleys deep and wide, outer very narrow, valleys ending at the single large longitudinal ridge on inner edge of teeth; incisors compressed, rather short, In lower premolar-molar se-

378

PHILIPPINE LAND MAMMALS

ries each tooth with four well-defined cusps, the inner anterior much the highest; on outer edge a distinct vertical groove, and a lesser inner; on two posterior lower molars a groove enters anterior edge of tooth, then turns outward isolating a low cusp; upper and lower tooth rows practically parallel; outer surface of incisors deep orange. Measurements of Sciurus steerii Gunther. mm.

Head and body, from a dry skin Tail from a dry skin Height of ear Width of ear Hind foot Naked sole of hind foot to base of claw Skull: Condyle to incisor tip Total length Width of brain case Zygomatic width Diameter of orbit Diastema Width between tooth rows Orbit to nasal tips Upper tooth row Lower tooth row Mandible to incisor tip Mandible, bone only

215 178 16.1 10

47 36.1 41 49 20.2 28

9.8 12.5 6.5 20.1 9.4 10 34.5 28.8

Remarks.-Thomas has separated the original species into two forms and given each specific rank. The name steerii is applied to the form from Balabac and the name iw;vencus is applied to the specimens (presumably) largely from Puerto Princesa, Palawan. He states: In 1876 Dr. Gunther described and figured two squirrels collected by Dr. Steere on the islands of Balabac and Palawan as Sciurus steerii, he being under the impression that they represented but a single species inhabiting the two islands. Since then, however, a number of these squirrels have come to Europe, and all those from Balabac prove to be precisely identical with Dr. Gunther's No. 1 from that island, while all thQSe from Palawan, although considerably more variable, inter se, than the Balabac ones, agree in differing from the latter by their browner, less rufous colour, their grayish sides, and black-tipped tail, as described by Dt. Gunther under 2. In belly-color, however, they may be white or either partially or wholly rufous. This being the case, it is evident that the squirrels from the two islands should be considered as distinct species, and I would propose to restrict the name S. steerii to Dr. Gunther's No.1, the foremost figure on his plate, with its type B. M. No. 76. 10. 4. 4, and to assign the name S.

379

SCIURUS

juvencus to the Palawan species, described by Dr. Gunther under No.2 and drawn in the background of his plate, its type being B. M. No. 76. 10. 4. 3. With regard to the variation in the belly-colour of S. juvencus it is to be noticed that the greatest extremes, all white and all red, are shown by specimens from the same place, Puerto Princesa.

My specimen from Sir J. Brooke Point, Palawan, agrees much more nearly with the Balabac form than with the specimens described as S. juvencus from Palawan. The chief differences noted is a tendency of the black on tail to appear in dull rings when held in certain lights. The shade of red in the type figure is somewhat more reddish than my specimen. The specimen was caught at the base of a tree in a trap where an hour before I had removed a specimen of Tupaia. The two forms bear a very marked resemblance to each other. SCIURUS JUVENCUS Thomas

Sciur;us steerii (part.) GUNTHER, Proc. Zool. Soc. London (1876) 736, pI. 69 (part.) Sciurus juvencus THOMAS, Ann. & Mag. Nat. Hist. 2 (1908) 498 (Palawan); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 25; Proc. U. S. Nat. Mus. 46 (1913) 314 (Puerto Princesa, Palawan). ?Sciurus stee'rei STEERE, List Birds and Mammals Steere Expedition (1890) 29.

Description of the species.-Allied to S. steerii Gunther but differing in color. Color.-Upper part of head, middle of back, and outer side of legs grizzled with grayish brown and black, each hair having a black ring and black tip; sides of body similarly grizzled, but mixed with numerous white hairs; lower parts pure white, with the exception of root of tail; anal region and borders of white abdomen, brownish red; tail bright brownish red, tip black; hairs moderately long with a broad black ring. Measu?'ements of Sciurus juvencus Thomas. mm.

End of Length Length Length

snout to root of tail of tail of naked sole of hind foot of skull

228.6 177.8 38.1 50.8

Remarks.-See remarks under S. steerii. SCIURUS ALBICAUDA Matschie

Sciurus albicauda MATSCHIE, Sitz.-Ber. Ges. Nait. Freunde, Berlin No. 5 (May, 1898) 42; TROUESSART, Cat. Mamm., Suppl. (1904-05) 316; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 24; Proc. U. S. Nat. Mus. 46 (1912) 337.

380

PHILIPPINE LAND MAMMALS

Description of the type.-A squirrel allied to Sciurus mollendorfji, but differing in having ears black-brown, head grayish white, and terminal hair of tail white. Measurements .-Total length, 450 millimeters; tail, 210; caudal vertebrre, 160; foot, 50. Remarks.-Matschie states that he doubts that this species comes from Culion which was the locality placed on the tags, but that they probably originated on some nearby island. He gives the type locality as Calamianes Islands. The type and four paratypes were collected by von Mollendorff. Regarding the markings Matschie states: Die vorliegenden Exemplare zeichnen sich durch den hellen, weisslichen Kopf aus, dessen Haare weiss sind mit ganz kurzen, dunklen Spitzen. Der Riicken ist bei einigen Stiicken hell braun mit hellen S-triche1chen besprengt, bei anderen fast lehmfallbig ohne deutliche Strichelchen. Die Korperseiten sind sehr hell sandfarbig. Der Schwanz is an der Basis ebenso vie der Riicken gefarbt weiterhin sind die Haare dunklebraun oder fahlbraun mit langen weissen Spitzen. Die ohren sint dunklebraun behaart, die Vorderbeine wie die Korperseiten, die Hinterbeine etwas dunkler gefarbt. Die Unterseite und die Innenseite der Gliedmaassen sint weiss.

In this species the nasal bones are said to be less than the distance between the posterior nasal spine and the incisive foramina. SCIURUS MBLLENDORFFI Matschie

Sciurus mo.llendor/fi MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde, Berlin No.5 (May, 1898) 41; TROUESSART, Cat. Mamm. (1904-05) 316; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 25; Proc. U. S. Nat. Mus. 46 (1913) 337.

Description of the specie.s.-Sides of body strongly mixed with white hair; white area on belly extending for some distance to underside of base of tail as a well-defined line for from 2 to 5 centimeters; white hairs in shoulder regions, in old males a whitish shoulder stripe; distance from posterior nasal spine to posterior edge of incisive foramen equal to or greater than length of nasals. Measurements.-Total length, 400 to 470 millimeters; tail, 200 to 230; caudal vertebrre, 185; foot, 50. Remarks.-This species is described from several specimens collected by von Mollendorff in Culion, Calamian Islands. One specimen is mentioned as having white hair mixed in the pelage on the back and also on the top of the tail. The type is in the Berlin Museum.

SCIURUS

381

SCIURUS MINDANENSIS Steere

Sciurus mindanensis STEERE, List Birds and Mammals Steere ' Exp. Philippines (1890) 29; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 25; Proc. U. S. Nat. Mus. 46 (1913) 314 (Mindanao; Catagan, Lower Rio Grande Valley, Mount Malindang). Sciurus cagsi MEYER, Proc. Zool. Soc. London (1890) 599.

Description of the species.-(After Meyer's description of S. cagsi.) Fur short and rather stiff; upper parts entirely blackish, variegated or grizzled with light yellowish brown, each hair black, ringed with yellow near ~ip; bristles entirely black as well as whiskers; ears covered with short hair of same color as body; cheeks and side parts of head and neck rather more yellowish; underparts grayish or brownish white, underbase of tail more or less ferruginous; tail bushy, variegated with reddish brown, black and white above; beneath, brown edged laterally on each side by a longitudinal black stripe bordered externally with white; end of tail white. A skeleton examined by Meyer presumably of this species has 12 costal vertebrre, 7 or 8 lumbar, 3 sacral, and 26 caudal. Skull an.d teeth.-Skull elongated; two upper premolars, anteri-or minute; molars broad; second upper molar 3 millimeters; series of teeth parallel. Remarks.-Steere's description is very brief, as follows: Sciurus mindanensis sp. nov. Mindanao. Size [length] 14.50, body, 7.50, tail to tip of hairs 7.00. Above dark gray. Hairs black at their bases then buffy tipped with black. Darker along the middle of the back. Thighs and upper surface of fore and hind feet rufous gray. Nose and space about the eye rufous. Under surface of legs and body to the tail ashy. Tail nearly uniform with the back.

My specimen was collected at Saub, Cotabato, Mindanao, April, 1923. It was obtained from natives who caught it in a fallen tree. The coloring about the eyes and nose is not rufous as mentioned by Steere but the buffy color ("light yellow brown" of Meyer) is somewhat more predominant. The "ashy" color of the throat is mingled with buff, and the posterior part of the abdomen is distinctly reddish buff; the tnil markings are identical with those recorded for S. cagsi. The permanent dentition is not in place. The first (permanent?) premolar, at least, is scarcely to the alveolar surface; the second premolar is a milk tooth, the permanent tooth can be seen below it. Compared with the teeth of S. steerii (a specimen with the milk premolar in place) I note that the transverse ridges are less heavy, the valleys shallower, and the cusps

382

PHILIPPINE LAND MAMMALS

more pointed and less bulky; the vertical groove on outer side of molars is very much less distinct than in steerii; the audital bullre are larger and distinctly more inflated; the postorbital processes are more slender and the lachrymal process is more prominent. The skull on the whole is less heavy than S. steerii and the teeth are smaller, more noticeable in the lower teeth than the upper. SCIURUS PHILIPPINENSIS Waterhouse

Sciurus phiUppinensis WATERHOUSE, Proc. Zool.

SOC. London (1839) 117 (Mindanao); STEERE, List Birds and Mammals Steere Exp. Philippines (1890) 29 (Basilan); MEYER, Proc. Zool. Soc. London (1890) 600; THOMAS, Trans. Zool. Soc. London 14 (1898) 389, (Basilan); HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 25; Proc. U. S. Nat. Mus. 46 (1913) 314 (Basilan).

Description of the sp'ecies.-(After the type description.) Size moderate, tail not very bushy. This species is rather larger than Sc. palmarum and less than Sc. bivittatus. The general hue of the upper parts, sides of the body, and outer side of the hinder legs, is deep brown, a much richer and deeper color than the same parts in Sc. bivittatus; this tint is produced by the admixture of rust color and black, the hairs being of the latter color and rather broadly annulated with rusty-red near the apex. The tail is not very bushy; the hairs are black, with two bright rusty bars. The under parts of the body are grayish white, with a faint yellow tint; the head and forelegs are grayish, and the feet are black slightly grizzled with rust color. Measurements of the type of Sciurus philippinensis Waterhouse. mm.

Length Length Length Length Length

snout to base of tail of tail of snout to base of ears of tarsus and digits of ear

165 160 38 44.3 7.3

Rem{Jff'ks.-The type was collected in Mindanao by H. Cuming in 1839. It is now in the British Museum. Everett also collected the species at Zamboanga, Mindanao, and in Basilan. Casto de 'E lera records, probably incorrectly, a specimen from Balabac. SCIURUS SAMARENSIS Steere

Sciurus samarensis STEERE, List Birds and Mammals Steere Exp. Philippines (1890) 30 (Samar and Leyte) ; MEYER, Abh. Mus. Dresden (1896-97) 29, pI. 11, fig. 2; THOMAS, Trans. Zool. Soc. London (1898) 388; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 25; Proc. U. S. Nat. Mus. 46 (1913) 314. (Samar and Leyte).

SCIUROPTERUS

383

Description of the species.-(After the type description.) General color above and below as in S. rruindanensis. It differs from that species in being of larger size and in having the thighs and nose and space about eye uniform in (;olor with the back, and the upper surface of feet dark gray. Measur'ements of the type of Sciurus samarensis Steere.Length, 406.4 millimeters; tail, 203.2; body, 203.2. Remarks.-No definite type is designated and the locality given is Samar and Leyte. The short descriptions are very inadequate. Hollister states that there are two skins in the United States National Museum collected by Steere in "Samar or Leyte." The specimen figured by Meyer is thought by Thomas to be overcolored, since it does not agree with specimens in the British Museum as regards the black color of the feet. Family PETAURISTIDJE Body with lateral membranous expansion of body skin attaching to wrists and ankles; adapted for glissant flight. Only the genus Sciuropterus is represented in the Philippine fauna. Genus SCIUROPTERUS Cuvier

Sciuropterus F. CUVIER, Dents Mammif. (1825) 255; and Ann. der Mus. 1 (1825) 126; FLOWER and LYDEKKER, Mamm. Liv. and Extinct (1891) 453; 179.

FORSYTH-MAJOR,

Proc. Zoo!. Soc. London (1893)

Squirrellike animals with limbs connected by a membrane which attaches along the sides of the body and to the fore and hind feet; a second membrane from neck to forearm; fur rather elongate, without spines; ears large, thin, not pressed close to head; tail elongate, nearly equal to head and body, somewhat flattened, hair longer on the sides, so as to exaggerate the flatness. ., Skull strongly deflected, alveolar line passing through audital bulla above meatus; interorbital region concave"; postorbital processes present; anterior attachment of zygoma greatly widened; its lower edge pierced by the foramen, immediately below which is a small projection of the bone; in the anterior edge of orbit a slight projection suggests a preorbital process; zygomatic arch broad, heavy; palate extending a short distance behind line of posterior molars; bullre large, somewhat flattened. 1-1 0-0 2-2 3-3 Dental formula: 1.. , C. - , pm. - , m. = 22. 1-1 0-0 1-1 3-3

384

PHILIPPINE LAND MAMMALS

Incisors wedge-shaped, upper shorter than lower; anterior upper premolar small, less than one-third of following tooth; second premolar, and first and second molars with an obscure W- or M-shaped enamel fold forming three outer cusps; inner surface of teeth concave to edge; lower teeth with the folding less pronounced. This genus has been divided into four subgenera, two of which are known to occur in the Philippines. Hylopetes is represented by Sciuropterus nigripes nigripes Thomas and S. n. ew,ssodontus Osgood in the Palawan group. The other subgenus Petinomys is represented by Sciuropterus crinitus Hollister in Basilan. Only these three forms are known from the Islands. This genus has a wide distribution, occurring in North Amer- . ica, Malay Archipelago, and Asia, one species extending through Siberia to Europe. Key to the Philippine species of SC1tu?'opterus Cuvier. a 1. No ear tufts; ears very large, thinly haired terminally but with longer

yellowish hair on basal surface externally. Parachute thinly haired above ............_..._. __._._... __ .......... ......... __....... _.. S. nigripes Thomas (p. 384). a'. Three long tufts of hair, one below and one, on each side; smaller than nigripes; ears very small; fur on parachute long and heavy. S. crinitus Hollister (p. 389). SCIUROP11ERUS NIGRIPES Thomas

Sciuropterus nigripes THOMAS, Ann. & Mag. Nat. Hist. VI 12 (1893) 30.

Two subspecies are recognized. The typical form, presumably confined to Palawan (possibly also to the Calamianes), and the second form known only from Bankalan Island, south of Palawan. The Bankalan form differs in the following points: Distinctly smaller and skull smaller; nasals contracted postel'iorly; front edge of coronoid process unbeaded; incisor teeth decidedly slenderer; cheek teeth average 10 per cent smaller. SCIUROPTERUS NIGRIPES NIGRIPES (Thomas)

Sciuropte1'US nigripes THOMAS, Ann. & Mag. Nat. Hist. VI 12 (1893) 30; ELLIOT, Cat. ColI. Mamm. Field. Col. Misc. Pub. 8 (Palawan); ALLEN, Bull. Am. Mus. Nat. Hist. 28 HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 26; Proc. Mus. 46 (1913) 315; ELERA, Contrib. Fauna Filipina pf. facing p. 140.

(1907) 186 (1910) 14; U. S. Nat. (1915) 139,

SCIUROPTERUS

385

Description of the species.-(From No. 647, E. H. Taylor collection; collected at Sir J. Brooke Point, southern Palawan, October 10, 1925, by E. H. Taylor.) Adult female. Ears large, inner surface naked, outer surface well furred on basal third, hair reaching a higher point posteriorly than anteriorly, upper portion of ear with very thin, extremely short hair; vibrissre heavy, elongate, reaching nearly double the distance from snout to ear; longest hair, 65 millimeters long; fur on forehead short, 18 millimeters long; on neck, 15 to 17, the longest hairs 20; on back, 28 to 32; on wing membranes, of similar length but much thinner; edge of membrane with a fringe of dense hair 15 millimeters long just behind attachment to foot, but only about 8 millimeters on sides medially; on edge of membrane near hind leg tufted edge much less dense; fur moderately dense behind hind leg (similar to back) but long, 35 millimeters (longest hairs, 40) ; tail somewhat flattened, fur about 43 to 45 millimeters long near base, 35 near distal end, and 30 at tip; fur on belly uneven, longest hair rarely exceeding 22 millimeters, outer longer hair straight, shorter underfur woolly on back. _Color.-Ears blackish at tips; upper and lower lips blackish, cheek and throat pale. Above, indefinite grizzled gray-brown with many hairs tipped with blackish along middle of back, fewer laterally, while other hairs are tipped with brownish drab; underfur slate to mouse gray; head, particularly crown and forehead, similarly colored; side of face, chin, and sides of neck indefinite whitish to grayish buff, the hairs frequently tipped with black or brownish and basal color variable from buff to gray; wing membrane above with hair tipped with brownish buff, basal color slate-black, edge of membrane with brownish buff hair, not bicolored or very slightly so; hair on abdomen and under wing membranes blotched with pure buff, brownish buff, slate, pure white, and dim brownish gray, darkest in median abdominal region; tail at base, above grayish, laterally buff; hairs become black tipped, and continuing backwards the black portion of hair increases and basal color changes from brownish gray to nearly solid black towards tip; below, tail at base, strong buff, extreme tips of hairs blackish, basal color continues buff, mixed with black hair for nearly half the length of tail where it becomes distinctly silvery on sides; near tip the entire hair is black. Black band on tip of hair continually increases to tip of tail until almost the entire hair is black. 269463-26

386

PHILIPPINE LAND MAMMALS

Skull and teet h.-Skull rather strongly deflected, alveolar line passing through upper part of bullre; bullre large, moderately inflated; nasals extending beyond incisors; a well developed postorbital process and a small projecting process in anterior part of orbit; a slight rounded notch just anterior to base of postorbital processes; zygoma attached anteriorly by broad base; base pierced by infraorbital foramen, which has a bony process bounding its lower side; a semicircular notch behind posterior base of zygoma, and circular notches in sides of palate immediately behind tooth rows; nasals narrowed posteriorly, broadened an~ teriorly with a slight median anterior groove, their length distinctly less than frontal; a distinct depression in interorbital region. Upper incisors elongate; combined width at tip, 3.9 millimeters; length, 8.3; first lower premolar very small, inserted at a strong angle leaning backward against second premolar; second premolar large, molariform, enamel folded and ridged, leaving three prominent outer cusps with a small accessory cusp between two posterior, on anterior side of third cusp; enamel covering sides of ridges somewhat beaded and pitted; on first and second molars, folds do not entirely cross teeth -(scarcely more than halfway), inner surface somewhat depressed; both molars with three prominent outer cus路p s; third molar, apparently more worn than others, with only two prominent cusps; four large cheek teeth all three-rooted (first premolar with only one root) ; posterior teeth of series tending to turn their crowns somewhat outward; lower incisors longer than upper; four lower cheek teeth, the first three with two prominent outer cusps formed by an enamel fold; inner edge with two less-strongly defined cusps, the anterior on each tooth highest. Remarks.-I have examined several specimens of Sciuropterus nigripes nigrilpes in the possession of Mr. V. M. M.acasaet, of Puerto Princesa. One specimen is nearly white. There are a few brownish marks on the head, on the anterior tip of the ears and on the anterior part of the arm. On one side there is a rather large patch of brown behind the arm, four very small spots on the wing and one on the groin; on the left side there is a spot on the side and one on the edge of the membrane; the tail has two irregular spots. The remainder of the fur above is paper white; the entire underside of the body is pure white without a mark. The body measures (mounted skin) in length 310 millimeters; tail, about 305.

387

SCIUROPTERUS Measu?"ements of S ciuropt erus nigripes nigripes (Tho mas ). [Measuremen ts in millimeters. ] N o. 9.nd sex.

--'---;----;---\ Iieadand body ... _____ ______ _____ _____ ____ ______ ___ ____ _ T ail to tip of hair . . ... ______ _____ ______ ____ _________ __ ___ _ Hind foot. m oist .. __ _______ ___ ______ ______ _____ __ ___ ____ _ Ear from notch _. _____ __ ____ __ - _____ ______ ______ _____1__ _ _

T yp e, female.

648 , male.

330 250 50 26

298 309 51 26

Skull: Upper lengtb __. ___ _____ ________ ___________ ______ ____ 51. 5 Basal length , incisors to condyle _____ ____ __ ___ ____ _____ ____ __ ___ _ Zygomatic breadth _____ __ ________ ___________________ : 32.5 Nasals, length._ ___ _ ___ ______ ___ ___ ____ _____ _____ __ __ 15.1 Nasals, greatest breadth â&#x20AC;˘. ___ ____ ____ _________ _____ ___ 8.6 Frontal length ____ ____ ________ ______ ,_ .,,____ ________ ____ ______ ~

Interorbital breadth, ignoring supra orbital notches ~. ___ __ Postorbital breadth _~______ _ _______________________ _

48 33 17 9.5 23 . 5

12 12.2

13 12 . 2

13 232 24

D iastema, u pper_.. -- ~~ - --~ -- - -- ~ - - - ~ ~ ~- ~ ~ ~ ~ 10 . 6 L ength of upper tooth series. alveolaL ~~ ~~---T-- - ---~ Length of fou r large teeth __ . ..,- _. __. . . . ~ ____ ..,________ 11.4 L ength of mandible to condyle_______________ ________ ______ ____

10 . 1 12 11 30

__ __... _.

308 310 53 26

51 44 31.2 16 8.1 20

Diameter of orbit ... ____ ___ ----- - ---- - -- - ---- - ------ _____ _____ 1 Breadth from tip of pastorbit al processes ____ __________ _ ______ ____ Palate . .. _____ -- ___ -- __-",- ---- __ - -- __-- __ - ----- -- ___ 28

------r -

I I

12.3 12.2

647 , female.

I I

13 . 3 22 . 5 26 11 . 5 12.2 11.2 33 . 4

L ength from incisor tip to condyle. - - - -- ___ --- - - - --- _ 1_- - -- - -- -35. 8 38 . 5 ~~~stema, lower . - - - - -- - - -- - - - - - - - - - - - --- - --- - - - - -- - - - - - - - - -- .. 4 .2 . . ~ ~wer tooth r ow_ -- ------ --------- --- ------------ -==- - - -~-- -~l_ ~~

The specimen was collected in 1919 when a clearing was being made at Iwahig, and Mr. Macasaet tells me that many dozens of the white form were found together with the darker formsometimes both forms issuing from the same cavity in the tree. It is certainly not a sex difference as both sexes were found in both forms. Mr. Macasaet further states that the spotting is very irregular and that sometimes specimens are found that are nearly half brown-and that the specimen described above was the one showing the least amount of spotting. The skull is mounted in the skin so it was impossible to compare the skulls and teeth. A number of skins in Mr. Macasaet's collection of the dark form . showed considerable variation. Most of the specimens had a mouse to slate-gray underfur with a banding of graybrown on the outer tips, the gray predominating on the membranes.

388

PHILIPPINE LAND MAMMALS

One specimen, a large male, had the membrane fur deep brown; the fur above on back mouse gray, below uniform grayish tipped with lighter color; and in the groin and posterior part of the belly the fur had a red-brown cast. All specimens have the basal area of the tail smoky white to buff-white, distal end for three-fourths of length black or black-brown, basal parts of fur lighter, but less so toward tip. These s~uirrels live for the most part in cavities of trees. In the neighborhood of Iwahig they are very common, at times becoming something of a pest to fields near the forest. They sometimes eat green corn, squash, and beans. These squirrels sustain themselves in the air by spreading their limbs with the membranes and volplaning from a higher to a lower point. The specimen listed as No. 648 was presented to me by Mr. V. M. Macasaet, of Puerto Princesa. The types were collected by A. Everett, 1877-7$, at Puerto Princesa, Palawan. SCIUROPTERUS NIGRIPES ELASSODONTUS Osgood

Sciuropterus nigripes ela,ssodontus ton 31 (1918) 1.

OSGOOD,

Proc. BioI. Soc. Washing-

Description of the type.-(After Osgood.) General characters those of SciJuropterus nigripes but of much smaller size; skull and teeth re}atively small. Color.-Upper parts g'rayish, hair brown overlying a deep neutral gray basal color; top of head slightly more grayish but essentially ,like back; patagium rather thinly haired and colored like back; sides of nape bordered by dull grayish, continuous with sides of neck and postauricular region; under parts chiefly with a neutral gray basal color overlaid by pale creamy; a few scattered and irregular areas wholly cream-colored or whitish; cheek, chin, sides of neck and front of breast dull mouse gray; ears dusky; feet brownish black; tail, in proximal third, wood brown, slightly, or not at all, tipped with black on under side and rather broadly tipped with it, at least medially, on upper side; distal two-thirds brownish black, the hairs more or less brownish fawn at their bases. Skull and teeth.-Skull smaller than that of Sciuropterus nigripes nigripes; nasals rather contracted posteriorly and definitely exceeding premaxillre; interparietal with transverse dimension exceeding longitudinal; frontoparietal suture acutely emarginate; front edge of coronoid process unbeaded; incisor teeth decidedly slenderer than in Sciur()pterus nigripes nigripes; cheek teeth essentially the same, but one-tenth smaller.

389

SCIUROPTERUS

Measurements of the type Scituropterrus nigripes elassodontus OsgOOlA mm.

Total length Tail Hind foot Ear Skun: Occipitonasal length Basal length Zygomatic breadth Length of nasals Anterior breadth of nasals Posterior (least) breadth of nasals Length of palate Diastema Upper cheek teeth Length of large premolar

500 250 45

25 45.7 42'.5 28 14.5 8.1 3.6

21.5 8.8 10.8

2.5

Remarks.-The type and one cotype were collected on Bancal an Island, north Balabac Strait, Philippines, July 16, 1916, by C. M. Weber. They are No. 21, 774, in the Field Museum of Natural Hiistory, Chicago. SCIUROPTERUS CRINITUS Hollister

SciuropteruB crinitus HOLLISTER, Proc. Bi()l. SOC. Washington 24 (1911) 185; Philip. Journ. Sci. ยง D 7 (1912) 25; Proc. U. S. Nat. Mus. 46 (1913) 337.

Description of the species.-A flying squirrel, externally somewhat like Sciuropterus nigripes, but smaller; ears very small with three long tufts of hair, one from below and one from each side; fur on back and upper side of parachute long and heavy; belly thinly haired. Color.-General color of upper parts pale chestnut, finely mixed with black; nose and lips gray: crown, back, and upper surface of parachute and tail pale chestnut, finely lined with black; tail blackish near tip; underfur slate color, hairs tipped with chestnut and black; underparts of body and limbs buffy, underfur gray; underside of tail pale chestnut, lighter along median line, ear tufts brown at base, black on terminal half; vibrissre elongate, black. Skull a'Yl!d teeth.-The skull resembles that of other members of Petinomys, subgenus of Sciuropterus, but with bullre slightly less flattened and cheek teeth with the small accessory cusps at outer exit of the valley between the two main transverse ridges very minute; mastoids not inflated; small premolar robust and other cheek teeth slightly incre~sing in size posteriorly; crowns

390

PHILIPPINE LAND MAMMALS

low; enamel finely sculptured; ridges low with transverse ridges somewhat broken, supplementary cusps between them small, but plainly visible on last upper premolar and less on first upper molar; obsolete on second molar. Measur'em ents of the type of SciU'J"opterus crinitus HollisteT. mm.

Total length 570 Tail 260 Hind foot 42 Ear from notch, on dry skin 13.5 Length of ear tufts 36 Skull: Greatsst length 53 Condylobasal length 48.6 Basal length 45.1 Zygomatic width 32 Least interorbital breadth 10.1 Greatest breadth of nasals 9.2 Mastoid breadth 22 Length of maxillary tooth row, alveoli 11.6 Length of mandible from anterior surface of symphysis to condyle 32.5

Remarks.-The type specimen, an adult male from Basilan Isl,and, was collected by R. C. McGregor and A. Celestino, January 12, 1907. This is the only specimen known. Family MURIDJE Skull variable, usually with a strongly constricted interorbital area; zygomatic processes present; jugal short and slender, generally reduced to a splint between zygomatic process of maxilla and squamosal; lower root of anterior zygomatic process flattened into a perpendicular plate leaving infraorbital vacuity tall and wide above, narrow below; lower incisors compressed, chisel-like; no premolars; molars rootless or with roots, tuberculate or with angular enamel folds; pollex vestigial; hallux usually with claw; tail usually nearly naked with scales arranged in annulations. Chiefly terrestrial. The family is divided into several subfamilies, two of which are represented in the Philippines. Fifty or more genera are known, of which sixteen occur in the Philippines. Key to the PhiliPPine subfamilies of MuridlÂŁ. a'. Molars divided into transverse laminre; tails well furred; larger. PHLCEOMYIN)E (p. 391). a, 2. Molars tuberculate, rooted; tails usually nearly naked; typical rats and mice; smaller ........................ :............................... MURIN)E (p. 400).

'391

PHL<EOMYS Subfamily PHLrnOMYIN....lE

This subfamily, apparently confined to the Philippinel'\, is 'c haracterized by the characters of the molars which are formed of transverse lamellre. The pelage is heavy, elongate; tail heavily furred. Two genera are known. Key to the geneTa of

~he

Phlreomyinre.

a'. Muzzle blunt; tail about 350 millimeters and thickly haired; colors

a 2.

usually .gray and blackish or reddish brown and yellow. I Phlreomys Waterhouse (p. 391). Muzzle pointed, tail about 425 millimeters and more bushy than in Phlreomys; grayish black to blackish. Crateromys Thomas (p. 397). Genus PHL(EOMYS Waterhouse

Phlreomys WATERHOUSE, Proc. Zool. Soc. London 7 (1839) 108; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 27. Phlreomys GRAY, List Mamm. Brit. Mus. (1843) 115; ELERA, Cat. Sisto Fauna .E'ilipinas 1 (1895) 21,; ContT. Fauna Filipina (1915) 144; TROUESSART, Cat. Mamm. (1898) 459.

Large rodents; tail shorter than bead and body; muzzle blunt; ears hairy externally; tail thickly haired; incisor teeth very broad. Skull. and路 teeth.-Skull rather ovate, occipital portion rather elongate and somewhat contracted; a distinct postorbital process; interparietal bone almost circular; anterior molar of upper jaw consisting of three transverse lami~re, second ~nd third consisting of two transverse lamirue each; in lower jaw anterior molar consisting of four laminre, a small rounded lobe in front followed by two transverse laminre, the anterior smaller; there follows a small transverse posterior lobe; second molar consisting of two equal transverse laminre and a small lobe bE:hind them; last molar of two simple transverse laminre. Three species of the genus have been described; namely, Phlmomys cumingi Waterhouse, the type of the genus, P. pallidus Nehring, and P. elegans (Cabrera). Elera lists a fourth form, P. albayensis. Key to the Philippine species of Phlreomys Waterhouse. a 1, Usually blackish brown washed with lighter. P. cumingi (Waterhouse) (p. 392). a 2. Usually paler, dark anteriorly; posteriorly brown, heavily washed with grayish white ....................................... . P. pallidus (Nehring) (p. 394). a'. Similar to above but general body color reddish yellow. P. elegans (Cabrera) (p. 396).

392

PHILIPPINE LAND MAMMALS PHL<EOMYS CUMINGI (Waterhouse)

Mus (Phlreomys) cumingi WATERHOUSE, Proc. Zool. SOC. London (1839) 108; EYDOUX and SOULEYET, Voyage de Ia Bonite (1841) 4'3, pIs. 7 and 8. 118

120

PHILIPPINE ISLANDS

I 4

122 Batan I.

126

® ® © ® ® ® ® ®

Phloeomys cuminSI Phl~omys pailidus Crateromys schadenbergi Apomys hylocretes " Apomys insignis Apomys insignis bardus I A pomys petra:!us Apomys major CD A pomys musculus Q) Apomys microdon 16

'"'0

<tJ C,

C"l

'">\

<"'l

..... ~ (..)

"'Cagey"n Sulu

120

FIG. 19.

122

124

126

Distribution of Phleeo1nys. Crateromys. and ApDm4/.s in the Philippines.

PhZreomys cttming"ii ELERA, Cat. Sisto Fauna Filipinas 1 (1895) 21; ContI'. Fauna Filipina (1915) 144; GRAY, List Mamm. Brit. Mus. (1848) 115; TROUESSART, Cat. Mamm. (1898) 459. PhZreomys cumingi MEYER, Abhand. und Berichte des Konigl. Zool. Mus. Dresden (1896) 29, pIs. 12 and 13; THOMAS, Trans. Zool. Soc. London 14 (1898) 400; HOLLISTER, Philip. J ourn. Sci. § D 7 (1912)

393

PHL<EOMYS

27; Proc. U. S. Nat. Mus. 46 (1913) 337; STEERE, List Birds and Mammals Steere Expedition (1890) 29 (Marinduque). ? Phlremys albayensis (nomen nudum) ELERA, Cat. Sisto Fauna Filipinas 1 (1895) 21; Contr. Fauna Filipina (1915) 144.

Description of the species.-Fur rough, suberect, intermixed with longer hairs; ear moderate, bearing elongate hairs; feet large and wide, bare below; tail moderate, covered with elongate fur. Color.-Above, blackish brown, washed with dirty yellowish; paler below; tail blackish; longer hairs, on head and back, dark. Skull and teeth.-"The skull compared with that of the common rat differs in being of a more ovate form, the occipital portion being somewhat elongated, and considerably contracted; the width between the orbits being comparatively great; and behind the 路orbits the frontal bones are expanded, and join with the temporal to form a distinct postorbital process. The interparietal bone, instead of being transverse, is almost circular. The auditory bullre are very small. The interdental portion of the palate is slightly contracted in front, so that the molars diverge posteriorly; the rami of the lower jaw are less deeply emarginated behind, the coronoid portion is smaller, and the descending ramus is broader and rounded; the symphysis menti is of considerable extent. The incisor teeth are less compressed and less deep from front to back. The molar teeth are of a more simple structure; the anterior molar of the upper jaw consists of three transverse lobes, and the second and third each of two transverse lobes. In the lower jaw the anterior molar consists of four lobes, a small rounded lobe in front followed by two transverse lobes, of which the anterior is the smaller, and finally a small transverse posterior lobe; the second molar consists of two equal transverse lobes, and a small lobe behind them; the last molar consists of two simple transverse lobes." (Waterhouse.) Measurements of PhlCBomys cumingi (Waterhouse). mm.

Length, head and body Tail Forefoot, excluding claws Tarsus Ear Skull: Length Width

482.6 320 43.4 70 25.4 59.2 43.3

Rematrks.-The type was collected in Luzon by H. Cuming; the exact locality is unknown. The species has been reported from Luzon, Marinduque (Steere), and Mindoro (Trouessart).

394

PHILIPPINE LAND MAMMALS

The measurements are from the type, changed to millimeters from inches and lines. The type locality as given by Waterhouse is "apud insulani Luzon." PHL<EOMYS PALLIDUS (Nehring)

PhZmomys cumingi var. pallidus NEHRING, Sitz.-Ber. Ges. Nat. Freunde Berlin (1890) 106 (Luzon). P hZmomys pallidus THOMAS, Trans. Zoo1. Soc. London 14 (1898) 399 (La Trinidad, Benguet; Cape Engafio; Lepanto; Mount Data) ; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 27; Proc. U. S. Nat. Mus. 46 (1913) 315 (Haight's Place and Irisan, Benguet).

Description of the species.-(From No. 650, E. H. Taylor collection; collected at Balbalan, Kalinga, April 23, 1920, by E; H. Taylor.) Larger than Phlmomys cumingi, with longer and softer fur; females with inguinal mammre

1:; 1

ears short, naked

within, with scattered hair on ears- extending beyond tip for more than one-third length of ear; longest hair of moustache reaching ear; fur on forehead and crown erect; forefoot moderate, hallux with a fiat nail; tail furred with short hairs, shorter than fur on body,' not wholly concealing annulations on tail, of which there are eleven or twelve to the centimeter; tail ,shorter than head and body; pollex with strong claw not smaller than thos~ on other digits. Color.-Fur on anterior part of body deep black on distal third and grayish to gray-brown on basal two-thirds; forelimbs above and below, back, sides, and belly with distal third of longer hair white; with basal two-thirds and all the shorter fur grayish brown or (on hind limbs and belly) reddish brown ~ crest on forehead and crown with distal half grizzled white, basal half fading into gray; ears black with black hairs; cheek fur gray, tips gray or black; black about eyes and on tip of snout; hair of tail deep black to brownish black; skin blackbrow,n ; a few gray hairs near base of tail. Skull and teeth.-Top of skull slightly concave medially, supraorbital ridges rising considerably above surface of brain case, forming small blunt postorbital processes; interparietal a distinct triangle; tooth series directed outward at a considerable angle; cutting edges of crowns continuous, transverse; incisors not especially long, their color somewhat orange externally; first upper molar with three cusps, second and last with two; anterior lower molar with four cusps, the anterior small and about half as

395

PHL<EOMYS

wide as adj oining rows; second lower molar with three cusps, the third small; third lower molar with two cusps; lower molar series directed strongly inwards. Measurements of PhZm01nys pallidus (Nehring). rom.

375 315 26 51 73

Head and body, from skin in alcohol Tail to tip of hair Ear from crown Forefoot with claws Hind foot with claw Skull: Condylobasal length Zygomatic breadth Interorbital breadth Mandible Upper tooth row Lower tooth row

85 4'5

15.8 56 18.6 19.2

The following table shows comparative measurements of the skulls of four specimens all of which, presumably, represent this form.

Measurements of fe11lale SlJecimens of Phlmmys pallid~ls (Nehring), from . M o1fntain Province, Luzon. [Measurements in millimeters.] Balbalan, Irisan, Kalinga. Haights. Benguet. Benguet. E. H. Tay- United States National Bureau of lor Museum. Science collection. collection. Condylobasallength .. ________________________ _ Zygomatic breadtb . _____.___________ __ ________ _ Length of nasals _____________________________ _ Interorbital breadth . _________________________ _ .Manclible _________________ . _________________ _

~:!:: ::::~ ::::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I Q

85 45 (a)

15.8 56 18.6 19.2

85 46.4 34.4 16 57.3路 19.8 20.3

84.5 43.4 15.8 55.9 18.9 20.4

82.3 40.5

13.8 64 . 6 19.3 20 . 3

Broken.

Remarks.-The type locality for the species is "Luzon." Thomas reports the species from several localities in northern Luzon. I obtained my specimen from some Kalingas who had just captured it with the aid of their dogs. I skinned the animal and turned over the body for food to a boy who assisted me. It is regarded by the Kalingas as a delicacy. The native name is

yut-yut.

396

PHILIPPINE LAND MAMMALS PHL<EOMYS ELEGANS (Cabrera)

Capromys elegans CABRERA, Bo!. SOC. Espanola Hist. Nat. (1901) 372 fig. (Cuba?). Phlmomys elegans CABRERA, Trab. Mus. Cien. Nat. No. 3 (1912) 30 (Philippine Islands). Phlmomys elegans HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 337.

Description of the species.-(After the type description 'Of Capromys elegans.) Body and limbs covered with thick fur, harsh and rough on head; hairs on f'Orehead erect and somewhat stiff, forming a small transverse crest which hides external base 'Of ears; tail rather large, c'Overed above moderately well by elongate stiff hairs; undersurface of tail naked. Color.-General body color brilliant reddish yellow; head, upper part of neck, limbs, and hairs on tail dull hazel brown; a small tawny-yellow spot between eyes; greater part of back covered by a blotch in the f'Orm 'Of a lance, the tip pointing toward lumbar region, the base somewhat widened on shoulders, c'Ontours of blotch P'Oorly defined; blotch intense reddish brown, less definite anteriorly; numerous white hairs surrounding the SP'Ot, forming a wide irregular border; on shoulders, anterior part of hind legs, and above base of tail many white hairs mixed with those of the general gr'Ound color, but not f'Orming spots. Claws large, much curved, yell'Owish gray; incisor teeth yellowish. M e.aswrement8 of PnlmoWJI8 elegans (Cabrera). mm.

Head and body Tail Head Hind foot, without claws

485 200

95 75

Remarks.-This species was described from an old mounted specimen from which the 'Original tag was lost. Cabrera believed that it had 'Originated in Cuba and described it as a member 'Of the West Indian genus Capromys, without examining the skull due to the fact that the skull had been m'Ounted with the specimen. Later Cabrera * transferred the species to Phlreomys. The exact type locality is unknown. Since the measurements are made fr'Om a m'Ounted specimen it is imp'Ossible t'O make definite comparison with measurements of the other tW'O forms. I suspect that the species may actually belong t'O one 'Or the 'Other 'Of the two, preceding forms.

* Trab.

Mus. Cien. Nat. No.3 (1912) 30.

CRATEROMYS

397

Genus eRA TEROMYS Thomas

Phlreomys (part.) MEYER, Abh. Mus. Dresden 6 (1894'-5) 1. Crateromys THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 163; Trans. Zoo!. Soc. London 14 (1898) 408; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 33; Proc. U. S. Nat. Mus. 46 (1913) 326.

Large, long-haired rodents; ears small, eyes small; vibrissre much elongated; pollex not prominent, with a well-developed flattened nail; other toes on forefoot relatively short; claws slender; tail elongate, covered with long bushy hair, no longer at tip than on sides; hair much elongate over body, particularly on rump; pelage generally woolly. Skull elongate, muzzle slender, zygoma greatly widened; a distinct interorbital depression; brain case not greatly expanded; edges of interorbital area raised, passing back on top of brain case as low ridges; palatal foramina much elongated; strong palatal emargination behind; ptergoids well developed; bullre small, only slightly inflated; palatal foramina long; palatal area between tooth rows narrow, with a median ridge and two deep lateral grooves; incisors moderately developed, very light, dull yellow on anterior side, flattened anteriorly and somewhat rounded laterally; upper molars large, heavy; anterior molar with three large, nearly equal, external, and three, greatly reduced, internal cusps; second with two, large, external cusps and three internal; last molar with two, large, external cusps and three internal, last internal and last external cusps not separated by a posterior groove; longitudinal groove separating inner and outer cusps distinctly nearer inner edge than outer edge; point of separation between cusps marked by a deep infolding of enamel wall of lamina, a notch so deep that it cuts lamina in two; in lower molars, inner and outer cusps nearly equal and longitudinal groove on tooth row nearly medial; anterior lower molar with three outer and three inner cusps, the anterior inner cusp of series largest and nearly medial; a small posterior supplementary median cusp; second molar with two inner and two outer cusps, and a small supplementary median posterior cusp; posterior molar with two inner and two outer cusps; lamina bent backwards so that axes of cusps meet medially at right angles. Orateromys schadenbergi (Meyer) is the only species of the genus known. The type was a skin collected by A. Schadenberg. No skull accompanied it and Meyer placed the species in Phlceomys. The specimens on which the genus was founded were collected by J. Whitehead on Mount Data, northern Luzon, February,

398

PHILIPPINE LAND MAMMALS

1895. The relationship is not clear. Thomas has suggested a relationship with Lenomys. It is certainly not closely related to Phlceomys. CRATEROMYS SCHADENBERGI (Meyer)

PhlCEO'lnys (?) schadenbergi MEYER, Abh. Mus. Dresden 6 (1894-5) 1.*

C'rateromys schadenbergi THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 163; MEYER, Abh. Mus. Dresden No.6 (1896-7) 32, pI. 13, figs. 3-6 (skull), pI. 14 (animal); THOMAS, Trans. Zoo!. Soc. London 14 (1898) 409, pI. 36, fig. 2 (teeth); HOLLISTER, Philip. Joum. Sci. ยง D 7 (1912) 33; Proc. U. S. Nat. Mus. 46 (1913) 326 (Baguio; "Northern Luzon").

Description of the 'species.-(From No. 139, Bureau of Science collection; collected at Sitio de Uyong, Mount Data, Luzon, P. I., elevation 2,168 meters, by Victor E. Lednicky.) Body elongate; with long, heavily furred tail; "ibrissre on snout elongate, the longest 90 millimeters in length; two elongate vibrissre above eye, and two posterior to eye; ear small, almost completely concealed in fur; fur on sides of head short, composed of short underwool and slightly longer straight hairs; anterior part of face and forehead wit~ a crest, short anteriorly but lengthening toward crown where longest hairs measure more than 40 millimeters; pelage of back dense, consisting of a woolly underfur with longer nearly straight or wavy overfur, both growing longer towards rump; underfur on neck, 40 millimeters long; overfur, 65; underfur on rump, 70 to 80; overfur, 100; overfur very scant on belly, 35 millimeters long; underfur, 25 to 30; underfur scant and short on tail, 20 to 25 millimeters; overfur long and wavy, 55 to 60; hair on tip of tail and below shorter; limbs moderate; pollex very short with a distinct flattened nail; other four toes with well-developed claws; on hind limb each of the five toes with well-developed claws; feet with short fur rather concealing skin; strong tufts of hair about base of each claw. Color.-General color of pelage blackish; crest on head somewhat brownish black; outer fur on neck and shoulders black from tip to skin; underfur gray externally, becoming dark gray to black near skin; color of underfur becoming darker on back posteriorly until it is nearly uniform dull black; color remains grayish on sides; entire ventral side, and generally iron gray,

* See also Meyer, ZooI. Garten 31 (1890) 195-200 and Lpz. 111. Ztg. (1891) No. 2479, S. 21, for data on the type specimens of this species, kept alive in Germany.

399

CRA'I'EROMYS

black where overfur is slightly denser; and area behind ears is without overfur and appears gray. Skull and teeth.-See description under genus Crateromys. Measwrements of Crate1'omys sohadenbe1'gi (Meyer). mm.

Total length, from skin Head and body Tail, without hair Ear height Skull: Greatest length Condylobasal length Zygomatic width Interorbital width, least Width of brain case, greatest Upper tooth row, alveolar Lower tooth row, alveolar Diastema, upper Diastema, lower Mandible to tip of incisors Length of nasals Palatal length

700 345 355 20 67 64 37 8 23 14.8 16.4 17.5 9 45.5 25.2 31

Rema路r ks.-This curious rodent is presumably confined to the high plateau region of northern Luzon. Mount Data is the type locality. , The animal is very variable in color. Sometimes the anterior 'Part of the body is whitish or brownish, and on the und~rside there is frequently a white area of greater or lesser extent. The Igorots of Mount Data are accustomed to capture the an.., imal and sell the pelts in the Baguio Market. I obtained there four badly stretched and mutilated skins (having no scientific value), which were made into a neck furpiece. The woollike fur is very striking and serviceable. Whitehead * states: Schadenberg's great Rat seems to be fairly common among the high mountains of Central N. W. Luzon. Like most Rodents, it is of nocturnal habits, and therefore the domestic economy of this Rat, or perhaps SquirrelRat, is difficult to describe. The Igorrotes, however, captured a number of specimens for me, some, they said, from holes in trees, others from holes among tree-roots; they described the animal as feeding on fruits up in the trees, and not on the fallen ones. As this Rat was nearly always br(}ught. to me alive, I often allowe.d it to climb the pine-trees, which it did with perfect ease. In the day these animals tried to hide from the sun as. much as possible, and I f(}rmed an opinion that they were dull inoffensive

* In

Thomas, Trans. Zool. Soc. London 14 (1898) 410.

400

PHILIPPINE LAND MAMMALS

creatures, until one day directly an Igorrote opened the basket in which he carried and captured the Rat, the animal sprang out, and was back in the basket again in a second, but the Igorrote's thumb had the top nearly bit ten off. The cry of the Crateromys is a curious 'Thewo thewo thewo,' uttered so shrilly that the notes might proceed from some of the peculiar forest insects. Generally speaking, Crater omys is jet-black; about 30 per cent. are of a beautiful white-grey, and some 15 per cent. piebald, black and white. This distribution has nothing to do with age, as both grey and black young ones were obtained. The Igorrote name for this curio-u s animal is

'Bii-ut'. Distribution.-High mountains of Central Northern Luzon.

The specimen which I have described bears a tag in Mr. Lednicky's handwriting, as follows: "Edge of pine and scrub oaks. Lives in holes or cavities of pines. Eats buds and bark of young sprouts of pine trees. Caught in snares." Hollister * gives a list of measurements of five specimens in the United States National Museum as follows: Measurements of skulls of Crateromys 8chadenbergi (Meyer). [Measurements in millimeters.]

Northern Luzon. Baguio. No. 44. female.

Condylobasallength ___ ___ Zygomatic breadth ________ Interorbital breadth ___ __ __ Length of nasals __ __ ______ Mastoid breadth __________ Palatallength ____________ Mandible___ _____________ Maxillary tooth row __ ____ Mandibular tooth row _____

65.7 36 . 7 8 ,1 25.7 22 . 2 34.8 43.5 15.9 16.2

No. 102546, No. 102544, No. 102545, No. 102554, female. male. male. male. 67.1 36.6 8.1 26.8 22 . 6 36 . 8 44 . 8 16 . 5 17.7

67.8 37.7 8.2 26.4 22 . 2 37.7 45 . 5 17 . 8 17.9

65.1 35.4 7.4 26.8 21.1 35 42.9 15 15 . 8

67.5

-- ---------7.9 27.8 22.8 36.4

JiJ 16.3 16.2

Subfamily MURIN.ÂŁ

This group contains the typical rats and mice, well known on account of their domestic habits. Upper molars have the tubercles usually arranged in three longitudinal series; teeth are usually 3 or 4-rooted; members usually resemble each other rather closely in general characteristics and habitats. For the 'most part the colors are dull and similar throughout the group.

* Proc.

U. S. Nat. Mus. 46 (1913) 327.

MURINJE

401

The Philippines are rich in genera; many are endemic and frequently confined to very limited areas. Thus, the high mountains of central northern Luzon have yielded the following endemic genera not known elsewhere: Cela3nomys, Chrotomys, Rhynchomys, Tryphomys, Batomys, and CarlJOmys. There are several other genera not known outside the Philippines. These are Bullimus (Mindanao and Luzon), Limnomys (Mindanao and Mindoro), Tarsomys (Mindanao), Apomys (Mindanao, Catanduanes, Luzon), and Insula3m~ts (Busuanga). Aside from the endemic genera only the wide-spread genera Rattus and Mus are known in the Philippines. Key to the Philippine genera of the subfamily MurinIB.

a'. Two upper and two lower molars. b'. Molars large; muzzle not greatly elongated. Celamomys Thomas (p. 402). b '. Molars greatly reduced; muzzle elongate, shrewlike. Rhynchomys Thomas (p. 404). a '".Three upper and three lower molars. b 1. Back prominently striped longitudinally; tail relatively short, 111 millimeters; head and body, 196; hind foot, 35. Ch1'otomys Thomas (p. 406). b'. Back uniformly colored, not striped. c 1. Pelage spiny. d '. Pelage short and very spiny; tail short; annuli on tail numerous (18 to 1 centimeter); mammre, two pairs pectoral, one pail' inguinal . ___ ._ ... _.... _..... ___ . __ .... _... __ ..... _.. Crunomys Thomas (p. 407). d'. Pelage long, spines slender; tail moderately long; hind foot, 54 millimeters; mammre, one pair pectoral, t.hree pairs inguinal. Bullimus Mearns (p. 411). d 8. Pelage variable; size from very large to small. Sections of the genus having spiny pelage; norvegicus group; R. keUe?'i of the alexandrinus group; most forms of the rattus group, and those of the everetti group._ .......................... Rattus Fischer (p. 414). 0'. Pelage without spines. d 1. Two pairs of inguinal mammre. e 1. Teeth simple; no tuft of hair on tip of tail; hallux with claw. Apomys Mearns (p. 459). e'. Teeth extremely complex; a well-developed tuft of hair on tail; hallux with flat naiL. .. _.... _. ____ .. _. Insulmmus g. nov. (p. 469). e 8. Teeth with laminre transverse; no tuft on tip of tail, but tail generally hairy; hallux with claw. Carpomys Thomas (p. 471). d '. Mammre, two pairs pectoral, one pair abdominal, two inguinal; fur harsh; ouier digits of foot reduced. Tryphomys Miller (p. 476). d 8. Mammre, three pairs pectoral-abdominal, two pairs inguinal; very small mice .................................................. Mus Linnreus (p. 477). 269453-26

402

PHILIPPINE LAND MAMMALS

d'. :M:amm.re variable or unknown. e 1. Eye surrounded by a naked or finely haired ring; large, heavily furred rat; tail hairy ____ _______ ___ Batomys Thomas (p. 480). e'. Eye not surrounded by a naked or finely haired ring. fl. Medium-sized rat; heavy fur above j belly densely furred; tubercles of feet flattened, <the posteroexternal tubercle of hind foot obsolete; no additional tubercle. Limnomys Mearns (p. 484). f 2. Medium-sized rats; pelage long and coarse; hind feet 6-tuberculate, many tubercles bilobed or with small accessory tubercles______ __________ ________ ____________ T(Lrsomys Mearns (p. 487). f 3 . Hind feet typically 6-tuberculate; a few species lacking spines of g enus Rattu s ___________ ______ ____ _________ Rattus Fischer (p. 414).

It is not improbable that CelflJnomys, Chrotomys, and Crunomys are referable to the subfamily Hydromyime. Thomas * has tentatively, at least, placed the three genera in this subfamily. Genus CELlENOMYS Thomas

X erom ys (?) THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 161: Celrenomlls THOMAs, Trans. Zoo1. Soc. London 14 (1898) 390; HOLLISTER, Philip. J ·ourn. Sci. § D 7 (1912) 26.

Related to Chrotomys, but the pelage without striking marking; fur soft and velvety. Skull and teeth.-Skull broad and strong, wedge-shaped in lateral profile owjng to height of brain case and the way the f'rontonasal and palatal profiles a-pproach each other; no ridges; nasals short, not projecting beyond incisors; brain case smooth and rounded; ante orbital foramen little expanded above front; edge of outer plate vertical; palatal foramina small. Incisors thrown forward, simple, rounded and bevelled in front; two upper and two lower molars similar to two anterior molars in Chrotomys, but ridges and crests apparently less sharp; no trace of third molar. Only the species Celmnomys silaceus (Thomas) is known. CELlENOMYS SILACEUS (Thomas)

Xeromys (?) si laceus THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 161. Celrenomys siZ(Lceu s THOMAS, Trans. Zoo1. Soc. London 14 (1898) 391, pI. 31, fig. 16 (animal); pI. 35, figs. 11 and 12 (skull and teeth); HOLLISTER, Philip. Journ. Sci. § :0 7 (1912) 26; Proc. U. S. Nat. Mus. 46 (1913) 315.

* Trans.

Zoo1. Soc. London 14 (1898) 394.

403

CELlENOMYS

Description of the specie's .-Size of a common rat, with soft, close, velvety fur; hairs on posterior back 10 to 12 millimeters long; eye small, not surrounded by a black ring; ears short; tail thinly haired, a little shorter than body without head. Color.-General color uniform slaty gray, very finely grizzled with whitish; sides of muzzle nearly black; undersurface paler than back, not sharply defined, the hairs slaty gray basally, washed with buffy white terminally; ears uniform grayish; feet dark gray as far as metapodials; digits whitish or flesh colored; tail brown above basally, whitish below and at tip. Skull and teeth.-Skull broad and strong, evenlJl" rounded, without ridges, decidedly wedge-shaped in lateral view; brain case high, smooth, evenly rounded; nasals short, not overhanging incisors; interparietal moderately developed; anteorbital foramen little expanded above, front edge of its outer plate vertical, not produced forward; palatal foramina very small; posterior nares broad; lower edge of mandible pe.culiarly flattened Just behind symphysis, and pierced with numerous minute foramina; coronoid processes long, strongly curved backward. Incisors I:lluch thro~n forward; only two upper and two lowel" molars in each jaw; no trace of a third molar. Measu?"ements of the type of CeloJnomys silaceus (Thomas). mm.

Head and body, about Tail Hind foot, moistened Skull: Basal length Greatest breadth Nasals, length Nasals, breadth Interorbital breadth Interparietal length Diastema, length Anterior palatine foramina, length Upper molar series, length Mandible, bone only

195 110 33.4 34

19 12 3.3

6.9 2.8 13.1 3.5 3.8 21.5

Remarks.-The type was collected by J. Whitehead on Mount Data, Lepanto, Luzon, February, 1895, at an elevation of approximately 2,460 meters. He says: This curious Mammal at first sight might easily be confounded with Rhynchomys sorricoides, and, like that animal, was also obtained on the table-4;opped summit of Monte Data. It seems rare, only two specimens

404

PHILIPPINE LAND MAMMALS

having been snared in some five weeks. The skull and teeth, instead of being frail as in Rhynchomys, are powerful, and much more nearly allied to Chrotomys. The eye is small as in Rhynchomys, and the ontwanl ~pÂ pearance quite Shrew-like. The habits of this peculiar Mammal I am quite unable even to guess at.

Hollister lists a specimen collected at Haight's Place, Benguct, by Edgar A. Mearns. The three specimens are apparently the only ones known. Genus RHYNCHOMYS Thomas Rhynchomys' THOMAS, Ann. & Mag. Nat. Rist. VI 16 (1895) 160; Trans. Zoo!. Soc. London 14 (1898) 396; HOLLISTER, Philip. J ourn. Sci. Â§ D 7 (1912) 27.

Muzzle enormously elongated like a gigantic shrew; feet normally like the rats, the pollex with a broad nail; tail ratlike, thinly haired; lower jaw low, slender and little curved. Skull and. teeth.-Skull long, brain case broad, smooth and rounded; muzzle narrow and much elongated; nasal long and narrow, terminating behind at level of premaxillre; viewed in profile they show a curious rise at their anterior extremity, the general frontal profile being practically an even slope from crown to a point at end of anterior third of nasals, and then bending upward again; interorbital region smooth, evenly rounded, quite unridged; interparietal large, transversely oblong, unusually variable as to its exact shape; anteorbital foramen typically ratlike but its outer wall very narrow and much slanted backwards, so that anterior edge of upper root-the bridgeis actually posterior to hinder edge of lower root; zygomata slender, low; palate long and narrow, and a distinct incisive fissure present; palatal foramina normal but far forward in skull so that their posterior end is nearly their full length in front of molars; on each side just in front of first upper molar, there is distinct raised ridge about two millimeters long, and there is a somewhat similar ridge behind last molar in lower jaw; these ridges are very possibly used to supplement the minute teeth in eating; posterior bony palate broad and produced far back; posterior nares rather narrow; internal ptergoid processes large, triangular, projecting far downward, even below level of tips of incisors; external processes practically or quib obsolete; no enclosed ptergoid fossre; bul1re small, but not of abnormal structure. Teeth extraordinarily reduced, the dental armature being smaller in this genus than in any other toothed land animal; in-

405

RHYNCHOMYS

cisors white above, pale yellow below; upper ones not grooved, very short, narrow, slender and forming an arc of a very small circle, so that their roots come opposite the anterior end of palatal foramina; two upper and two lower molars so minute that they appear of little use; lower incisors very slender, finely pointed; lower molars similar to size and shape of upper molars. (After Thomas.) One specimen examined by Thomas had only a single lower molar. The type of the genus is Rhynchomys s01'icoides Thomas. The genus appears to be isolated and shows no close relationships. RYNCHOMYS SORICOIDES Thomas

Rhynchomys soricoides THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 160; Trans. Zool. Soc. London 14 (1898) 393, pI. 31, fig. 2; pI. 35, figs. 7 and 10; HOLLISTER, Philip. J ourn. Sci. 搂 D 7 (1912) 27; Proc. U. S. Nat. Mus. 46 (1913) 337.

Description of the species.-Size about that of a common rat; fur very thick, close and velvety, about 14 or 15 millimeters long on back; general color uniform dark olivaceous-gray, without markings anywhere; undersurface dirty gray, not sharply defined; a white patch sometimes present on throat, ears about as in Rattus rattus, their visible surface, when folded, clothed with blackish hairs; metapodials brown, digits whitish; tail shorter than head and body, fairly clothed, but not tufted, blackish above, scarcely paler, below. (After Thom.as.) MeasU1'ements of the type of Rhynchomys soricoides Thomas. mm.

Head and body Tail Hind foot, moistened Skull: Basal length Diastema Length of the two upper molars

215 146 41 4'4 16.8 2.5

Remarks.-Only five specimens of this species have been collected. They were taken on Mount Data, northern Luzon, at an elevation of approximately 2,460 meters. This interesting Shrew-Rat was obtained on the summit of Monte Data, where only five specimens were snared. I am unfortunately unable to give any account of 路t he habits of this extraordinary mammal. Th.e, Igorrotes told me that it lives on grass, which is probably untrue, the teeth apparently being quite unfitted for such food; insects and worms are probably the diet suited for such rudimentary molars. The eye is, compar-

406

PHILIPPINE LAND MAMMALS

atively speaking, small, which leads me to believe that Rhynchomys is a diurnal-feeding Rat, like th~ true Shrews. (Whitehead in Thomas.)

A splendid life-size figure of this species in color is given by Thomas. Genus CHROTOMYS Thomas

Chrotomys THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 161; Trans. Zool. Soc. London 14 (1898) 391; HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 27.

About the size of a common rat but with shorter taiL The form is very similar to the genus Celmnomys Thomas; the formula of mammre 2-2 = 4; molars three above and below; skull otherwise similar to Hydromys; back prominently striped; fur soft and straight; muzzle not cleft, eyes rather small; ears large; pollex with a rounded nail; other digits, including hallux. with well-developed, little-curved claws. Skull and teeth.-In general form the skull is not unlike the skull in Celmnomys, but it is even more wedge-shaped owing to its greater height posteriorly; nasals short, their anterior end level with middle of incisive fissure; interorbital regions similar-· ly rounded and unridged; brain case broader and shorter, its breadth equal to its length; interparietal very small, a mere narrow transverse slip; anterior edge of zygoma plate slightly concave, the plate little developed; incisive fissure large, quite half as large as one of the palatal foramina which are, as usual in this group, very small; posterior nares large and open, hinder edge of palate level with posterior lamina of second upper molar; pterygoids large, proj ecting downward considerably below level of either molars or bullre. Lower jaw as in Cel:;enomys. Incisors pale yellow, thrown forward, simple, rounded in front; three upper and three lower molars, the anterior two very similar in structure to those of Xeromys; third upper molar small, transversely or obliquely oval in section; in size the two posterior molars are barely two-thirds the length of the first. CHROTOMYS WHITEHEAD I

Thomas

Chrotomys whiteheadi THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 161; Trans. Zool. Soc. London 14 (1898) 392, pI. 32, and 35, fig. 8; HOLLISTER, Philip. J ourn. Sci. § D 7 (1912) 27; Proc. U. S. Nat. Mus. 46 (1913) 315.

Description of the species.-Size of Rattus rattus; fur soft and thick; ground color gray-brown, tending in some specimens to rufous; a well-defined bright buff or orange line extending from

407

CRUNOMYS

middle of face nearly to the tail, intensified by a broad shining black band on each side; undersurface slaty buff, not sharply defined; metapodials shining gray; digits white; tail shorter than head and body, thinly haired, blackish above, rather paler below. (After Thomas.) Measurements of the type of Chrotomys whiteheadi Thomas. mm.

Head and body Tail Hind fo()t, moistened Skull: Basal length Basilar length Greatest breadth Nasals, length Nasals, breadth Interorbital breadth Diastema Anterior palatine foramina , length Upper molar series Mandible condyle to incisor t ip

196

111 35 38

35 21.7 13.2

4.1 6.8 15.2 4.4 5.2 31.2

Remarks:-This is another of the distinctive species of the northern Luzon fauna discovered by Whitehead in the mountains of northern Luzon. A specimen was obtained by the Bureau of Science at Irisan, Benguet. Three of Whitehead's specimens that are now in the United States National Museum are from Lepanto Province. The type and cotypes were collected on Mount Data at an elevation of approximately 2,460 meters, February, 1895. According to Whitehead the species feeds on sweet potatoes and grass. Whitehead (in Thomas, loco cit.) states that he saw a specimen in Manila said to be from the forest of Tarlac. If this statement refers to specimens in Manila museums, I am inclined to believe that the locality record should be questioned. The striking coloration and marking easily differentiate this form from all other Philippine Muridre. It is unique. Genus CRUNOMYS Thomas

Crunomys THOMAS, Minutes Proc. Zool. Soc. London for June 15, 1897; Trans. Zool. Soc. London 14 (1898) 393; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 26.

Small; fur thickly mingled with spines; ears short, rounded; hallux with a claw; tail short, thinly haired, apparently flattened, a character which may be due to drying. Skull and teeth.-Skull low, flattened, its front profile rather concave; nasals long, overhanging incisors; interorbital region

408

PHILIPPINE LAND MAMMALS

broad, edges with scarcely a trace of beading; interparietal large; anteorbital foramina similar to Chrctomys. the ou~er plate, however, not produced forward; incisive fissure minute; anterior palatine foramina short; posterior edge of palate in line with hinder edge of third upper molar. The characters of the teeth are in general similar to Apomys. The genus is confined to the Philippines. Key to the species of Crunomys Thomas. a 1. General color grayish, somewhat yellowish on back; belly grayish white; digits white; hind foot, 23 millimeters. (Luzon.) C. fallax Thomas (p. 408). a'. General color bister; belly nearly same as back; digits brown; hind foot, 25 millimeters. (Mindanao.) ........ C. melanius Thomas (p. 409). CRUNOMYS F ALLAX Thomas

Crunomys fallax THOMAS, Minutes Proc. Zool. Soc. London for June 15, 1897; Trans. Zool. Soc. London 14 (1898) 394; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 26.

Description of the species.-Small; fur short and close, profusely mixed with flattened spines; neither hair nor spines on back longer than about 6 millimeters; whiskers long; ears short; fifth hind toe reaching end of first phalanx of fourth; tail about length of body without head, uniformly covered with short hair. Color.-General color pale grayish, somewhat yellowish un back; dorsal spines white, darkening to black at their tips; belly dirty grayish white, not sharply defined, hairs slaty basally, dull whitish terminally; sides of muzzle brown; whiskers mixed bla~k and white; ears uniformly brown; hands and feet grayish brown on metapodials, lightening to white on digits; tail black, lighter along undersurface. Skull and teeth.-Like many aquatic species the skull is low and 'flattened, its frontal profile concave; nasals long, overhanging the incisors in front; interorbital region broad, its edges with scarcely a trace of beading; interparietal large; general shape of anteorbital foramina almost exactly as in Chrotomys whiteheadi; anterior palatine foramina short; posterior edge of palate just level with hinder edge of third upper molar. Teeth related to those of true murines; anterior lamina oblique in first upper molar; second upper molar with a small anterointernal cusp, a long middle lamina, and a mesial circular lamina posteriorly; third molar subcircular with a small anterointernal cusp; lower molars similar to molars in Chrotomys,. save that third molar is bilaminate.

409

CRUNOMYS

Measurement of the type of Crunomys fallax Thomas. mm.

Head and body, length Tail Hind foot

Ear

Skull: Lambda to nasal tip Greatest breadth Nasals, length Nasals, breadth Interorbital breadth Interparietal breadth Anterior zygoma root, length Diastema Anterior palatine foramina, length Upper molar series, length Mandible, incisor tip to condyle

105 79 2'3 10 20.5 12.1 9.2 2.8 4.6 9.1 1.7

7.2 3.5 3.6 16.5

Remarks.-Only a single specimen of the species is known.

Concerning its capt"ure Whitehead (in Thomas, loco cit.) says: In one of my wanderIngs through the parched-up forests of Isabela (in North-Ce~tral Luzon) I noticed a small red Kingfisher (Ceyx melanura) tfiy into the scrub I\ear a small stream. Having only a large gun with me, I sent my servant back to, the village for a small collecting~gun. While seated beside the stream, a small mouse was observed among some large stones on the opposite side, busily searching after food. I opened one of my 16-bore cartridges and picked out all the sman shot (No.6) but four or five pellets, and luckily killed the small animal without much damage being done. Being sure, from its peculiar habits that it must be something interesting, I carefully skinned it and sent it home, and am now rewarded by the add~tion of another new generic form to this already interesting collection.

The type was collected in Isabela Province, Luzon, at an elevation of approximately 307 meters, May, 1894, by J. Whitehead. CRUNOMYS MELANIUS Thomas

Crunomys melanVus THOMAS, Abstr. Proc. Zool. Soc. London (1907) 5 (February 12); Proc. Zool. Soc. London (1907) 141; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 26; Proo. U. S. Nat. Mus. 46 (1913) 337.

Description of the species.-Larger than Crnmomys fallax; fur close and fine, intermingled with numerous flattened spines; hairs on back 6 to 7 millimeters in length; ears very short, closely haired; tail short, very finely scaled (about eighteen to the centimeter) and finely haired, the hairs about one and one-half scales long.

410

PHILIPPINE LAND MAMMALS

Color.-General color above blackish brown (nearest to bistre of Ridgway but darker) ; undersurface nearly as dark as upper without any line of demarcation; individually, hairs of back dull slaty basally, with buffy tips; spines slaty with black tips; ears brown; whole of limbs, including hands and feet, blackish brown; tail uniformly blackish. . Skull and teeth.-Skull less flattened than that of Crunomys fallax (but may be due to youth) ; brain case long, sides almost parallel, in continuation of broad interorbital region; supraorbital edges with scarcely a trace of beading; outer plate of zygoma root cut back as in C. fallax; palatal foramina short, narrow in front, broadened behind, not parallel-sided as in C. fallax. Molars like those of a very simple type-of Rattus, without any supernumerary lateral cusps; first and second lower molars each with a small median posterior cusp as in C. fallax. Measurements of Crunomys melanius Thomas. Type.

Head and body Tail Hind foot, without- claws Ear Skull: Greatest length Basilar length Zygomatic breadth Nasals, length Interorbital brâ&#x201A;Źadth Breadth of brain case Palatilar length Diastema Palatal foramina Length upper molar series Lower molar series Mandible, condyle to' incisor tip

Cotabato

mm.

specimen. mm.

98 68 25 13

122 79 25.5 14.5

28 22 14 9.2 5.@ 12.3

30 23 13 11.2 5.1 12.8

7 3.8X2.3 4.1

7.2 4 4.2 18.2

Remarks.-Thomas expresses some doubt as to the generic identification of his species, and is in still greater doubt as to the relationship of the genus. It is very doubtful that the relationship is with the Hydromyin:e. The form has hitherto been known only through the type, collected on Mount Apo, Davao, Mindanao at an elevation of approximately 924 meters, February 29, 1906, by M. P. Anderson. I collected a specimen of a form evidently of this species at Saub, Cotabato, Mindanao, in April, 1923. It was taken in the forest at sea level. It differs from the type in no important characters, except that it seems to be

411

BULLIMUS

somewhat larger. Mammre, two pairs pectoral, one pair inguinal. Vibrissre on snout mixed black and white, extending well beyond tip of ears; tail somewhat lighter below than above; setre on distal half of tail as long as width of two to three annuli; annuli eighteen to a centimeter; underside of feet purplish brown. The general color above is probably somewhat lighter than in the type, judging from the type description. The specimen is distinctly larger than the type. Genus BULLIMUS Mearns

Bullimus MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 450. Bullilmus HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 30. Mus (part.) THOMAS, Proc. ZooI. Soc. London (1907) 141.

Similar to Rattus rattus but with three additional subsidiary cusps to lower molar series of either side placed externally, one in front of last series of cusps of anterior molar, and one in front of each series of middle molars; lower jaw with a projecting capsule for the accommodation of base of lower incisor tooth; audital buUre collapsed and flattened externally so that audital opening is compressed and directed posteriorly; skull elevated in anteorbital region; rostrum elongate; frontoparietal bead slightly marked, bony palate narrow; size large, pelage coarse, containing a mixture of ordinarily coarse hair and slender spines on upper surface; whiskers long, reaching shoulder; mam-

mary formula: ~ = 8. The type of the genus is Bullimus bagobus Mearns. Thomas is of the opinion that the genus is a synonym of Mus (Rattus), but Hollister agrees with Mearns that the hypsodont molars and the unusual formula for the mammre warrant the retention of the genus. Tl1;ere are two endemic Philippine species in the genus. K ey to the species of Bullimus Mearns. a 1, Tail, basal one-half to t.wo-thirds black, paler below; distal portion white all around .................................. __ B. luzonicus (Thomas) (p. 411). a". Tail purplish brown, paler below.............. B. bagobus Mearns (p. 413). BULLIMUS LUZONICUS (Thomas)

Mus luzonicus THOMAS, Ann. & Mag. Nat. Hist. VI Trans. Zool. Soc. London 14 (1898) 401, pI. 36, fig. Epimys luzonicus HOLLISTER, Philip. Journ. Sci. ยง D Bullimus luzonicus HOLLISTIDR, Proc. U. S. Nat. Mus.

16 (1895) 163;

4' (skull). 7 (1912) 29. 46 (1913) 324.

412

PHILIPPINE LAND MAMMALS

Description of the species.-Similar in size and color to Rattus everetti; fur longer and softer, wool hairs of back about 20 millimeters long; the longer hairs 30 to 40 millimeters; ears of medium length, thinly haired; tail shorter than in R. everetti, well haired, coarsely scaled, eight or nine annulations to a centimeter. Color.-General color coarse grizzly brown, resulting from a mixture of buffy yellow and black; underhairs dark slaty basally, their tips for 4 or 5 millimeters, buff; long outer hairs black, some of them with their extreme tips whitish; underparts dull slaty buff not defined on sides; hairs slaty basally~ buff terminally; head clearer grayish owing to tips of shorter hairs being whitish rather than yellow; eyes with an indistinct blackish ring, most marked posteriorly above; back of ears blackish, finely edged with white; upper surface of hands and feet hairy; some hairs blackish, the majority silvery white; proximal part of tail black; distal part whitish. Skull and teeth.-Skull differs from that of Rattus everetti by reduction of Bupraorbital ridges, which merely form a fine beading along edges of frontal and practically disappear ha1 f路 way along parietals; brain case round, swollen, unusually smooth; posterior nares broad and open; incisors yellow; molars broader than in R. everetti, the laminre more simply transverse, and outer cusp of each lamina less distinctly defined from middle cusp. (After Thomas.) Measurements of Bullimus luzonicus (Thomas). mm.

Head and body Tail Hind foot, moistened Skull: Basal length Basilar length Greatest breadth Nasals, length Nasals, breadth Interorbital breadth Interparietal breadth Interparietal length Diastema Anterior palatine Upper molar series, length Mandible to tip of incisors

240 200 47

44 40.2 25.7 19.8

5.9 7 11.6 5.3 13.5 3.2

9.3 33.2

413

BULLIMUS

Remarks.-The type and three paratypes were collected by .J. Whitehead on Mount Data, February, 1905, at an elevation of approximately 2,460 meters. The types are in the British Museum. Edgar A. Mearns collected a specimen distinctly larger than the type at Haight's Place, Benguet. The following measurements of the skull of this specimen, an adult female with worn teeth, are given: "Greatest length, 56.4 mm.; condylobasallength, 54.4 ; zygomatic breadth, 2'6.4 ; palatal length, 29.6; greatest breadth of rostrum, In; interorbital breadth, 7.7; mastoid breadth, 20.8; length of mandjble, 34.2; maxillary toothrow, 10.3; mandibular toothrow, 10.3." BULLIMUS BAGOBUS Mearns

Bullimus bagobus MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 450; THOMAS, Proc. Zoo!. Soc. London (1907) 141; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 31; Proc. U. S. Nat. Mus. 46 (1913) 324.

Description of the species.-The characters of the species are the same as those given for the genus; tail scantily covered with fine short hairs, nowhere concealing the annuli, which number nine to a centimeter. Color.-Dorsal area from forehead to rump, clove brown many of the hairs tipped or ringed with tawny-olive, and with pelage slate color at base; on sides hair brown, thickly mixed with hairs having tawny-olive tips; pelage smoke gray at base; underparts yellowish white, with gray basal portion of hair appearing copiously along sides; feet sepia above except toes, which are dirty grayish white; claws flesh color at base tipped with horn color; whiskers black, slightly mixed with' grayish white ones; tail purplish brown above, perceptibly paler and faintly rufescent below. Skull and teeth.-Anterior upper molar 5-rooted; middle upper molar 4-rooted; posterior upper molar 2-rooted, the roots exhibiting a tendency to bifurcate (see characters assigned the genus). (After Mearns.) Measu?'ements of the type' of Bullimus bagobus Mearns. mm.

Head and body Tail, mutilated Hind foat Longest whisker

275 128 54'

414

PHILIPPINE LAND MAMMALS

Measu?"f3ments of the type of Bullimus bagobus Mearns-Continued. Skull: Basilar length (Hensel) Occipito-nasal length Greatest interparietal breadth Zygomatic breadth Internasal suture, length Audital bullre, length Audital bullre, width Upper molar row, length Distance between anterior molars Distance between posterior molars

mm.

46 58 21.6 27

23.3

10 7

10 4

4.6

Remarks.-The .type was collected at Todaya, Mount Apo, Davao, Mindanao, by Edgar A. Mearns, at an elevation of approximately 1,203 meters, July 13, 1904. The type is in the United States National Museum, No. 125248. Mearns say: "The unique specimen was snared by a native who brought it and a fine specimen of Mus mindanensis to me on the trail as I was leaving the Bagobo village of Todaya." Malcolm Anderson obtained for the Duke of Bedford's Zoological Expedition a single specim,en at an elevation of approximately 924 meters on Mount Apo. It is in the British Museum. Apparently only these two specimens are known. Genus RATTUS Fischer

Ruttus (sic) G. FISCHER, Das Nat.-Mus. Naturg. Paris 2 (1803) 128 (typ. error). Epimys TROUESSART, Bull. Soc. d'Etudes Sci. d'Angers 10 (1881) 117; MILLER, Proc. Biol. Soc. Wash. 23 (1910) 57; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 28.

Description of the genus.-"External form typically ratlike, the skull and teeth without special modifications; molars slightly graduated in size from first to third, the anterior tooth not tending to disappear; enamel folding of upper molars directly referable to a simple 9-cusped pattern and its reductions, the outer margin of ml and m2 never with more than three cusps, the inner margin of same teeth never with more than 2 cusps; ml usually with 5 roots, its first lamina not distorted by a backward displacement of antero-internal tubercle; upper incisor moderately compressed, set at such an angle that its outer side is worn smoothly away by action of lower tooth." (Miller.) This genus includes the true rats. Thirty-one species and subspecies are recognized in the Philippines. For convenience . the species are arranged in groups as follows:

415

RATTUS PALAWANENSIS GROUP.

Rattus palawanensis. EVERETTI GROUP.

Rattus Rattus Rattus Rattus

everetti. tagulayensis. gala. albigularis.

NORVEGICUS GROUP.

Rattus norvegicus. Rattus tyrannus. RATTUS GROUP.

Rattus magnirostris. Rattus robiginosus. Rattus mindanensis mindanensis. Rattus mindanensis tablasi. Rattus colo'mtus. Rattus 'rattus. Rattus 'ntindorensis.

ALEXANDRINUS GROUP.

Rattus zamboangre. Rattus kelleri. CONCOLOR-EPHIPPIUM GROUP.

Rattus Rattus Ruttus Rattus Rattus Rattus Rattus Rattus Rattus Rattus Ruttus Rattus

querceti. leucophretus. negrinus. todayensis. pantarensis. ornatulus. mayonicus. calcis. vigoratus. vulcani. basilanus. luteiventris.

DATJE GROUP.

Rattus datre. Rattus benguetensis.

These groups va~y in the presence or absence of spines, the mammary formuJ.re, the size, and the shape and characters of the skull. In so large a group of species it is difficult to arrange satisfactory keys. Key to groups of the genus Rattus known in the Philippine Islands. a 1. Tail bicolored. b '. Ears very large; tail black above, white below; fur short, spiny; mammre, 2 pairs pectoral, 1 pail' inguinal; one species (Palawan). palawanensis group (p. 416). b 2. Ears moderate; tail black at base, whitish distally; fur coarse, slightly spiny; mammre, 2 pairs pectoral, 2 pairs inguinal (Mindanao and Luzon) four species .............................. everetti group (p. 419). a 2. Tail unicolored. b '. Very large rats. c'. Skulls heavy; supraorbital beads on skull run nearly parallel on parietals; mamma!, 2 pairs pectoral, 2 pairs abdominal, 1 pair inguinal; two species (widespread) .... n010vegicus group (p. 425). b'. Medium large rats; supraorbital beads curving strongly on parietals (except in magnirostris) ; pelage with or without spines. c '. Mammre, 2 pairs pectoral, 2 pairs inguinal; seven Philippine species (widespread) .......................................... rattus group (p. 429). c 2. Mammre, 2 pairs pectoral, 1 pair abdominal, 1 pair inguinal; two species (Luzon and Mindanao) ........ alexandrinus group (p. 440). c 3 . Mammre, unknown; two species (northern Luzon). datre group (p. 443). b 3. Small rats; supraorbital beads curving or wanting on parietals; pelage more or less spiny; mammre, 2 pairs pectoral, 2 pairs abdominal; twelve species (widespread). concolor-ephippium group (p. 445).

416

PHILIPPINE LAND MAMMALS P ALA W ANENSIS GROUP

This form seems to be characterized by the bieolored tail, the very large ears; mammre, two pairs pectoral, one pair inguinal. Only the species Rattus pala.wanensis is considered in this group. RATTUS PALAWANENSIS ap. nov.

Type.-No 32, E. H. Taylor collection; collected at Sir. J. Brooke Point, Palawan, October, 1923, by E. H. Taylor. Adult female. Dry skin and skull. Diagnosis; description of the type.-Medium-sized rat, with very spiny pelage, very large ears and bicolored tail; snout rather slender with strong vibrissa! extending a considerable distance beyond tip of ears; head with short bristles distinctly appreciable to touch, the longest 3.5 to 4 millimeters; pelage of back and sides with very short soft underfur almost hidden by long stiff spiny bristles that extend over back and to only a somewhat lesser extent on sides, the longest ones approximately 11 millimeters on back, 8 to 9 millimeters on sides; spines on belly short (5 millimeters) and stiff, distinctly softer on neck and breast; spines scarcely n0ticeable on limbs; ears very large, erect and prominent, naked inside, with extremely minute hairs on outside; thumb moderately prominent; tail long with fine setre not or scarcely as long as the width of scales of which there are nine to a centimeter; mammre, two pairs pectoral, two pairs inguinal. Color.-General color of back between wood brown and fawn, lighter on sides, darker dorsally; a dark area in middle of forehead, not very clearly defined, and dark hairs about eyes; an ill-defined darker area on sides of neck and across shoulders, and a second, large, indefinite, darker area across middle of back; color on sides lightens considerably; throat, belly and part under legs cream colored, this color separated from that of sides by a very distinct dividing line running from forefoot to hind foot along side; tail grayish black above, cream white below; forefeet white, hind feet fawn mixed with white; ears dark brownish to bluish (in life). Skull and te.eth.-Skull slender, audital bullre low, widely separated; rostrum slender, tip of nasals extending only slightly more anterior than incisors; supraorbital beads strong, curved on parietals; posterior zygomatic root continued as a ridge; palatine foramina wide and bowed; zygomatic arches relatively

RATTUS

417

slender; skull from parietals forward, nearly level, slightly deflected anteriorly, Teeth typically those of Rattus except that outer cone of anterior lamina of first molar is distinctly reduced or wanting; 118

126

124

122 Balan Is

120

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PHILIPPINE ISLANDS

® Rattus palawanensis ® Rattus everetti © Rattus tagulayensis @

Rattus gala

® Rattus albisularis ® Rattus nOl'vegicus @ Rattus tyrannus

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FIG. 20.

122

124·

126

Distribution of seven species of Rattus in the Philippines.

posterior lower molar with three cones nearly equal in size, forming a triangle; upper incisors rather narrow, orange on anterior surface; lower incisors, slender, elongate, of a distinctly lighter color than upper incisors, 269463-27

418

PHILIPPINE LAND MAMMALS

Measurements of t he type and pa?'atypes 0/ R attus palawanens1s sp, nov . [Measur emen ts in millimeters.]

N o .â&#x20AC;˘ age, and sex.

-3-2-,---'-3-1, ---;--3-0,-,.--29-,------,-2-8-, ----o-~ a dul t fema le.

a dult ma le.

young female.

y oun g female.

y oung fem ale.

------------- --- --- --- --- - - Totallength _________________________ Head and body ______________________ TaiL _______________________________ Forefoot, moistened __________________ Hind (oat, moistened _________________ Height of ear from notch ______________ Width of ear, greatesL _______________ Skull: Totallength _____________________ Zygomatic width _________________ Least interorbital width ___________ W idth of brain case ______________ Length of nasals ________________': Distance between molars ____ ~ ____ Diastema ____________ __________ ~

Upper molars, a lveolar. ___________ Lower molars, alveolar__________ Mandible, incisors to condyle----

402 224 178 18 38 26 21. 5 47 19 6.2 18 19.6 4.8 13 6.1 6.6 28

250 18 40 25.5 20.6 49 20 7 19 20 4.6 13.2 6.1 6.7 28.6

337 19 0 147 18 36 25.5

46 18.4 6.2 18 18 .2 4.8 12.2 6.5 7 27.5

327 173 154 17 35 23 18 38.2 16 5 .9 16 .8 13 4 9.5 6.1 6 .3 22

young female.

---

310 180 130 16 34 23 . 5 18

308 175 133 17 35 22 17.6

40 15 6 16 13.5 3.9 10.1

40 16 5.9 16 14.5 4 10.2

6.4 23 .5

6.3 22

Variation.-The differences in specimens of various ages are remar kable, due somewhat to shedding. No. 31 is a large male with the pelage much worn and of the same general color as the type. The dark spot on the forehead is very distinct and another grayish black spot is evident at the base of the snout. The fine underfur is more mouse colored than in the t ype. The spines, which are heavily gr ooved on the upper surface, vary from wood brown to blackish on their tips. The scales on the tail ar e seven to a centimeter. The scrotal pouch is very large, measuring nearly 50 millimeters in length. The spines continue to and on the scrotal pouch, although here they are slender and less rigid. In No. 30 the general coloring is much darker, a mixed indefinite grayish with the underfur longer and the spines distinctly less prominent but quite as rigid except in the rump region; the spines continue to the base of the tail; the belly spines are very fine. The black spot on the head is very indefinite. In Nos. 29, 28, and 27, the spinous area forms a saddlelike band around the body, beginning some distance behind the shoulders and ending abruptly on the anterior part of the rump; anterior and posterior to this spinous area the body is covered

419

RATTUS

with short soft fur, 5 to 7 millimeters long. The general color is dark grayish in the spinous region, but the spineless regions are gray with a wash of russet brown which also extends on the limbs. All of the six specimens agree in the color of the underside of the body except that the younger specimens are more nearly white than cream. The tail is invariably bicolored, and the ears, which are distinctive in this form, all agree. Remarks.-I am uncertain as to the relationship of this species, but it certainly is a member of none of the Rattus groups known to occur in the Philippines. The six specimens (type and paratypes) were all collected in October, 1923, at Sir J. Brooke Point, southern Palawan, practically at sea level. The forest here showed very numerous burrows, usually about the base of trees, which apparently harbored this species. EVERETTI GROUP

The rats of this group are characterized by rather heavy bodies, long, rather rough pelage, the number of spines variable; long tail annulated, annuU distinctly scaled; tail bicolored, basal part dark brownish black or deep purple, terminal part dull white to yellowish. The type locality of Rattus everetti is no longer known. Most of the mammals listed in the paper * where the type is described are from Dinagat or Mindanao. Probably through an oversight the locality is omitted in the description, but as no other northern forms are listed I conclude that the type is from the south. However, except for Elera's record t for Mindanao and one which is probably not authentic, it appears not to have been found there again if it did originate there. I am of the opinion of Thomas~: that B~~llimus is a synonym of Rattus, but due to my lack of material, I feel that nothing authoritative would be gained by throwing it into the genus Rattus until larger series of the forms are available. If the genus is distinct it is obviously an offshoot (j)f this branch of the genus Rattus, and might easily include others of the everetti group. They have in common the curiously colored tail, the greatly elongated pelage and in the case of R. albigularis the same type of hypsodont molars (variable in albigularis). I doubt that the mammary formula alone is sufficient for separating the genera. Care should be taken not to confuse these forms.

* Gunther, Proc. Zoo!. Soc. London (1879) t Cat. Sisto Fauna Filipinas 1 (1905). :j: Proc. Zoo!. Soc. London (1907) 141.

75.

420

PHILIPPINE LAND MAMMALS

Key to the species of the everetti group of the genus Rattus.

a'. Fur with few spines, chiefly on sli de; feet blackish; tail blackish, terminal third white or yellow. b 1. Hind foot, 49 millimeters; greatest length, 500; upper molar series, 9.1; skull, 51.1; terminal third of tail white. R. eve'i'etti (Gunther) (p. 420). b 2 . Similar to above but teeth larger; hind foot, 48 millimeters; greatest length, 450; upper molar series, 9.4; skull, 48; tip of tail yellow. R. gala (Miller) (p. 421). a". Fur noticeably spiny on back; tail purplish white on distal half. b '. Above yellowish brown; feet bister; hind foot, 52 millimeters; total length, 522; skull, greatest length, 54; upper molar series, 10.1. R. tagulayensis (Mearns) (p. 422). b 2. Feet brown; hind foot, 52 millimeters; length, 496; skull, 54; upper molar series, 10 ............................ R . .albigularis (Mearns) (p. 423). RATTUS EVERETTI (Gunther)

Mus everetti

GUNTHER, Proc. Zoo!. Soc. London (1879) 75; TROUESCat. Mamm. Suppl. (1904) 367; ELERA, Cat. Sisto Fauna Filipinas 1 (1895) 196. Epimys everetti HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 28 i Proc. U. S. Nat. Mus. 46 (1913) 315-316. SART,

Description of the species.-Upper and under parts clothed in rather harsh fur; some of the hairs, especially on sides, slender, flattened, and channelled spines; hair of lower parts shorter and softer than others, longest hair 42 millimeters; ears rather short, rounded and naked; tail almost naked, hairs between annuli being very short. Measurements of Rattus everetti ( GUnther) . [Measurements in millimeters, from Hollister, Proc. U. S. Nat. Mus. 46 (1913) 316.]

I Totallength.โ ข __________ . Head and body __________ Tail ____________________ Hind foot with claws _____ Skull. Condylobasallength __ Zygomatic breadth ___ ~'dibl' ___________ Upper molar series ___ Lower molar series ___

United States National Museum No.

151,500

151,501

151,503

151,502

145,792

151,504

497 239 258 47

450 210 240 49

402 192 210 47

500 235 265 49

383 183 200 45

365 172 193 42

51.1 25.3 30.7 8.8 8.5

47.7 22.4 29.3 8.4 8.5

40.3 19.8 26.3 8.9 9

49.7 28.9 31.3 9.1 9

43.2 21.8 26.2 8.9 8.5

40.8 21 24.1 8.2 8.1

Color.-Underfur gray, shorter hairs brown, longer hairs black towards the extremity, or black with brown tips; lower

421

RATTUS

parts dusky gray; feet blackish; tail blackish with terminal third white. Remarks.-The type, in the British Museum, was collected by Everett, probably about 1878. The locality is unknown, but it would appear to be Dinagat or northern Mindanao. RATTUS GALA

(Miller)

Epimys gala MILLER, Proc. U. S. Nat. Mus. 38 (1911) 398; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 28; Proc. U. S. Nat. Mus. 46 (1913) 316.

Description of the species.-This lallge species is related to the everetti group and might be regarded as a dark local form of Rattus everetti Gunther. Color.-Back and sides a coarse grizzle of black and dull, light ochraceous-buff, the lighter color in excess everywhere except on rump; shoulders with a slight grayish tinge; sides, cheeks, and outer surface of legs somewhat lighter than back; underparts dull ochraceous-buff, not conspicuously contrasted with sides, the line of demarcation scarcely indicated; feet blackish brown, in noticeat>le contrast with legs; tail blackish though somewhat less than basal half; distal portion yellow. Skull and teeth.-In 'all general features the skull agrees with that of Rattus everetti; audital bullre smaller, their longitudinal diameter decidedly less than distance between bases of paraoccipital processes; teeth more r'o bust than in E. everetti, width of first upper molar equal to fully three-quarters the least distance between tooth rows. (After type description.) Measu?'ements of type of Rattus gala (Miller). Type. United States National Museum, No. 144633,

mm .

Head and body Tail Hind foot (dry) with claws Skull: Condylobasal length Zygomatic breadth Interorbital constriction Breadth of brain case, over roots of zygomata Depth of brain case at middle Nasal Diastema Mandible Maxillary molar row, alveoli Mandibular tooth row

209 241 48

Paratype. United States National Museum, No. 144632.

mm.

170 185 44

48 24.8 7.4 19 14.6 20.4 14.2 31.2 9.6 9.4

26.4 9.5 9.4

422

PHILIPPINE LAND MAMMALS

Remarks.-The type, in the United States National Museum, was collected by Edgar A. Mearns, with two other specimens, on Alag River, Mindoro, November 30, 1906. These are apparently the only specimens known. RATTUS TAGULAYENSIS

(Mearns)

Mus tagulayensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 439. Epimys ta,gulayensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 30; Proc. U. S. 'Nat. Mus. 46 (1913) 316.

Desc1'iption of the species.-A member of the Rattus everetti group, resembling Ra,t tus eve1'etti and Bullimus luzonicus, but larger and otherwise different from either; hairs stiff; caudal annuli ten to a centimeter; ears nearly naked; pelage of upper parts mixed throughout with spiny hairs. Color.-Upper parts yellowish brown, pelage gray at base except for a mixture of whitish spines, darkest in vertebral area; sides drab-gray slightly mixed with hairs having raw umber tips; underparts drab-gray, strongly washed with cinnamon and Isabella color on throat, chest, inner aspect of limbs, and genitals; feet bister above, with grayish white tufts at base of nails; claws flesh, tipped with horn color; ears dark drab, flesh color within; tail purplish black all around on basal half, whitish on terminal half. Skull and teeth.-Resembling those of Rattus xanthurus. Measurements of the type of Rattus tagulayensis (Mearns). mm.

Total length Head and body Tail vertebrre Forefoot Hind foot Length of long hair on back Head Ear above crown Ear above notch Longest whiskers Skull: Basilar length Greatest length Occipito-nasal length Interparietal breadth Length of internasal suture Diastema Greatest breadth of rostrum Length of upper t.ooth row

522 262 260 26.4 52 42 64 23 28 88 44 54 54 17.2 21 14.5 9.6 10.1

RATTUS

423

Remarks.-Three specimens are known. The type and paratype were collected at sea level at Tagulaya on Davao Gulf, at the foot of Mount Apo, Mindanao, July 15, 1904, by Edgar A. Mearns. A third specimen was collected in Basilan by J. B. Steere. Hollister states that the Basilan specimen probably represents a new species, but it is too young and in too poor condition to be certain. RATTUS ALBIGULARIS (Mearns)

Mus albigula?-is MEARNS, Pl'OC. U. S. Nat. Mus. 28 (1905) 440. Epimys albigrulwris HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 28; Proc. U. S. Nat. Mus. 46 (1913) 316.

Description of the species.-[From No. 145, Bureau of Science collection (United States National Museum, No. 125261) collected July 6, 1904, by Edgar A. Mearns at an elevation of approximately 2,300 meters on Mount Apo, Davao, Mindanao.] Adult female.-A large long-haired rat; pelage rather coarse, 13 millimeters long on head; on middle of back the longer spiny hair 35; on rump the lORgest hairs 45; on middle and posterior part of body the soft underfur approximately 26; soft fur on belly, about 10; spiny longer hairs not numerous and not noticeable to touch except in middle of back; due to length of hair the ears appear reduced; vibrissre on snout moderately long, rather fine; tail distinctly annulated and scaled; sew on tail very short, not more than 1.5 millimeters long, except toward end where they appear longer and finer; nine annuli to a centimeter; feet large and heavy; mamrrue, four pectoral and four inguinal. Color.-Above mixed yellowish and mummy brown, underpart of pelage gray, growing lighter gray at base, except for the elongate spiny hairs which are silvery to gray white; always darker along vertebral area; underfur whitish on chin, breast, and belly, brownish about anal region; two brownish areas on sides of throat; feet brownish, slightly grizzled; claws horn color; ears drab gray; tufts of hair about claws silver white. Skull and teeth.-Skull massive, with a relatively heavy supraorbital bead, which is very heavy anteriorly, on parietals greatly reduced and scarcely perceptible posteriorly; lachrymal forming a prominent proj ection near anterior attachment of zygoma; a very small median proj ection on posterior edge of palate; bullre large; palatine foramina elongate, somewhat bowed, followed by two rather distinct grooves reaching posterior edge of palate.

424

PHILIPPINE LAND MAMMALS

Teeth heavy, outer anterior cone of first upper molar reduced; a very small basal cone on external anterior part of second upper molar; third upper molar proportionally large; anterior lower molar appears to have three instead of two major cones on anterior lamina; third lower molar has posterior lamina almost as wide as anterior; upper incisors heavy deep ochraceousorange, lower incisors cadmium yellow. Measurements of Rattus albigularis (Mearns). Type. Bureau of Science. United States Nation8!l Museum. No. 145. No. 125258. mm. mm .

Total length Head and body Tail Ear height from notch Ear width Hind foot, with claws Skull: Condylobasal length Occipito-nasal length Zygomatic breadth Interorbital breadth Interparietal breadth Internasal suture Length di~stema Greatest breadth of rostrum Upper molar series Lower molar series Mandible Mandible from incisor tip

481 231 250 22 18.6 49 50.6 54 24'.8 7 15 20.5 14 10 9.3 9.3 30.7 34.2

4'96 260 236 24 52 51.8 54 25.8 16 20 9.8 10 9.8 32.6

Remarks.-The type and seventeen other specimens were collected by Edgar A. Mearns at elevations from approximately 2,075 to 2,430 meters on Mount Apo, Davao, Mindanao, during June and July, 1904. The young are mouse gray above, white below with more or less brown and russet colors added to the upper parts with increasing age. Mearns states that the teeth closely resemble those of Rattus xa.nthurus.. that the teeth have a tendency to develop extra tubercles on the outer borders of both upper and lower molars, but that the subsidiary cusps lack uniformity and are frequently wanting. He found that their food consisted of fruit and insects. The Bagobo name is oom-bough' ka.hwee-lee.

RATTUS

425

NORVEGICUS GROUP

Two species referable to this group occur in the Philippines, Rattus norvegicus (Erxleben), the so-called Norway rat and a larger species, R. tyrannus (Miller). The former probably occurs throughout the entire Archipelago, or at least at all ports. It is only infrequently taken away from human habitation. The second form is known as yet only from the type locality, Ticao Island. The forms of this group are easily differentiated from all other Philippine species by the character of the supraorbital beads which run nearly parallel along the sides of the braincase. One exception is Rattus magnirostris (Mearns) which has similar supraorbital ridges. Key to the Philippine species of the norv egicus gr oup. a \ Supraorbital beads more widely separated posteriorly than at point

where they cross the frontopa r ietal suture; skull, 52 millimeters long .................................................... R. norvegicus .(Erxleben) (p. 425) . a'. Supraorbital beads less widely separated posteriorly than at point where they cross the frontoparietal suture; skull, 66 millimeters long............................................................ R. tyro,nnus (Miller) (p. 427). RATTUS NORVEGICUS (Erxleben)

Mus norvegicus ERXLEBEN, Syst. Regn. Anim., Class I, Mamm. (1777) 381. ? Mus decumanus THOMAS, Trans. Zool. Soc. London 14 (1898) 402. Epimys norvegicus HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 29; Proc. U. S. Nat. Mus. 46 (1913) 317 (Luzon, Panay, Mindoro and Mindanao).

Description of the species.-(From No. 11, Bureau of Science collection; collected August 27, 1913, by Manuela Estacion.) Large rat; body covered with rather short sparse fur, mingled with numerous elongate, thickened, grooved hairs; shorter fur about 10 millimeters long, longer hairs reach a length of about 24 millimeters on rump; fUr on belly short, longest hairs about 10 millimeters in length; vibrissre on snol,1t reaching beyond ears; ears rather large, the upper anterior surface well haired with short wavy hair; tail shorter than head and body, tail thicker and more robust than in other Philippine species (R. tyrannus probably excepted); hairs on tail distinctly longer than width of two annuli; about eight annuli to a centimeter at middle of tail.

426

PHILIPPINE LAND MAMMALS

Color.-Above, a dark indefinite brown resulting from buff tips of shorter fur intermingled with long brownish black hairs; a darker shade on rump; color on sides rather definitely separated from that on ventral surface which is dirty grayish with a faint buffy wash; tail darker above than below, although not "bicolored;" hair above forefeet brownish, lighter than body; hind feet silvery gray. Skull and teeth.-Skull massive; rostrum more or less compressed laterally; nasals not extending as far forward as incisors, and ending posteriorly a short distance behind frontomaxillary suture; zygomatic arches thick, anterior part with a strong groove; lachrymals strongly projecting; supraorbital beads high, forming an angle on frontoparietal suture and continuing backward as two straight, nearly parallel lines (slightly wider posteriorly than anteriorly); palatine foramina large, reaching backward as far as beginning of first molar; first molar narrowed anteriorly, lacking outer anterior cusp; other molars with the typical arrangement of cusps. M eas~wcments

of Rattus n orvegicus (Erxleben ). mm.

Total length Head and body Tail Fore foot Hind foot Skull:" Greatest length Zygomatic width PalaJtine foramina, length Width of brain case Interorbital constriction Length of palate from foramina Upper tooth row, alveoli Lower tooth row, alveoli Mandible, incisors to condyle Mandible, bone

438 230 208 20 35 53 26 9.1 18.2 6.1 11.3 7.6 7

36 31.2

â&#x20AC;˘ Largest skull in my coJlection, No. 691.

Remarks.-This species, now spread over the greater part of the earth, shows considerable variation. I note in Philippine specimens a variation in the thickness and length of the pelage; certain variations in the color; in the character and size of annulations on the tail, and in the character of the supraorbital beads on the skull. Owing to the fact that there has been a more or less constant exchange of specimens from all parts of the world due to cos-

RATTUS

427

mopolitan shipping in Manila Harbor one can never be entirely certain that Manila specimens actually are Philippine. The species is known from most of the larger islands of the Philippines and it probably occurs in all of the islands visited by boats. The type locality is Norway. The species, however, probably originated in southern Asia. RATTUS TYRANNUS (Miller)

Epimys tyrannus MILLER, Proe. U. S. Nat. Mus. 38 (1911) 397; HOLLISTER, Philip. Journ. Sei. ยง D 7 (1912) 30; and Proe. U. S. Nat. Mus. 46 (1913) 317.

Description of the species.-Very large, similar to Ratt~~s imperator Thomas, of the Solomon Islands, the largest member d the genus; skull not differing greatly from Rattus norvegicui) in form; gen,eral form, size ot ear, relative length of tail and body, length of palm ana sole, of digits, and claws, about as in R . norvegicus; tail nearly naked, sparsely sprinkled with stiff hairs about 2 millimeters long; annulations strongly defined, about seven to a centimeter at middle of tail; palm and sole naked, tubercles similar to those of R. norvegicus, but slightly larger; a small but well-developed secondary wart at outer side of plantar tubercle at base of both first and fifth toes; posteroexternal plantar tubercle long, its greatest diameter two and one-half times that of sixth; fur coarse and harsh, very stiff, though nowhere actually spi!l1Y; underfur thin, longer hairs for the most part slender grooved bristles, those of middle ,of back about 15 millimeters long; rump and lumbar regions with rather conspicuous growth of coarse terete hairs 60 to 75 millimeters long. ColoT.-The whole animal a dull indefinite brown somewhat intermediate between wood brown and brocolli brown; median dorsal region from between eyes to base of tail with a darker tinge, sides of shoulders suffused with drab, underparts tinged with dull buff; a faintly indicated dark area around eyes; cheeks and muzzle fawn color; whiskers black in rather noticeable contrast; feet dull indefinite brown not contrasting with body; tail blackish through basal half, then whitish to tip. Skull and teeth.-Except for its much greater size, the skull does not differ greatly from that of Rattus norvegicus; supraorbital beads well developed, perhaps relatively higher than in R. norvegicus, continued as lateral ridges along side of brain case to outer margin of interparietal, where it joins lambdoid crest;

42,g

PHILIPPINE LAND MAMMALS

lateral ridges converging posteriorly, the distance between them at a point where they cross the frontoparietal suture greater than that at outer borders of interparietal, a condition the 118

120

122

124

126

Balan Is

PHILIPPINE ISUNDS

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速 Rattus magnif'>ostr>is @ Rattus rattu8

息 Rattus mindopensis @ Rattus colopatus

速 Rattus pobiginosus 速 Rattus mindanensis @ Rattus m. tablasi

10'

<:>Cagayan Sulu

120'

FIG. 21.

122

124'

126

Dis tribution of seven members of the Rattus rattus group in the Philippines.

reverse of that in R. norvegicus; suture between premaxillary and maxillary relatively farther forward in front of anteorbital foramen than in R. norvegicus. The teeth of the type are worn and their exact relationship is uncertain. In general they represent a stage of reduction of

429

RATTUS

the outer side of upper molars corresponding to R. rattus; fourth tubercle in both first and second upper molars with distinct outer reentrant angle. (After Miller.) Measurements of the type of Rattu8 tymnnus (Miller). mm.

Head and body Tail Hind foot, dry Skull: Condylobasal length Nasal Diastema Zygomatic breadth Mastoid breadth Greatest breadth across lateral ridges Interorbital constriction Depth of brain case at middle Upper molar row, alveoli Lower molar row

310 270 57 62.2 28 19.6

31 24.2 12.8 9.6 15.8 11

Remarks.-The type, in the Philippine Bureau of Science collection on permanept deposit in the United States Nafional Museum, was collected in Ticao Island, May 15, 1902, by R. C. McGregor and A. Celestino. Only the type, an adult male, is known. This is the largest species of Rattus known in the Philippines. RATTUS GROUP

Species of this group are familiar the world over as the common rat. Certain species, especially Rattus rattus, are widely distributed. This species and R. norvegicus probably have southern Asia for their original home. They have been carried throughout the world on ships; few ports are without them. Most of the Philippines species of the rattus group differ from those of the norvegicus group i:q having the supraorbital bead curving along the parietals instead of running nearly parallel along the sides of the brain case. Rattus magnirostris (Mearns), however, is an exception, and Mearns described it originally as belonging in the norvegicus group. Hollister, * however, has pointed out that the teeth are of the rattus type and the species should be classed as an aberrant form of that group.

* Proc.

U. S. Nat. Mus. 46 (1913) 317.

430

PHILIPPINE LAND MAMMALS

Key to Philippine species of the ra.ttus gr011,p. a. 1. Supraorbital beads nearly parallel along the sides of the brain case. (Mindanao.) _________________________ ___________ R. ma.gnirostris (Mearns) (p. 430). a. 2. Supraorbital beads distinctly curved on brain case. b'. General color blackish, above and below.

(Widespread.)

R. ra.ttus (Linnreus) (p. 431). b'. General color brownish above, varied below. c'. Supraorbital beads disappearing on parietals; fur smooth and soft without spines; dark brown above; usually grayish slate to drab below. (Mindoro.) ________ R. mindoremsis (Thomas) (p. 433). c~. Supraorbital beads continuing on parietals to lambdoid crest; generally russet brown above; spines present. d 1. Grayish buff below blending into color of sides; feet with broad bands of blackish brown; upper tooth row, 8.1 millimeters. (Basilan.) ________________________________ R. colo?'atus (Hollister) (p. 434). d '. Cream color below sharply defined from color of sides; feet buffy white; upper tooth row, 7.3 millimeters. R. robiginosus (Hollister) (p. 435), d路. Whitish drab-gray below, washed with cinnamon anteriorly; feet whitish drab; upper tooth row, 8 millimeters; pelage above mixed coarsely with longer black hairs. (Widespread in the Philippines.) ____ R. mindanensis mindanensis (Mearns) (p.437). d '. Similar to R. mindanensis (typical) but with a l'eddish throat and pectoral area. Rattus mindanensis tablasi subsp. nov. (p. 439). RATTUS MAGNIROSTRIS (Mearns)

Mus magnirostris MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 441. Epimys magnit'ostris HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 29; Proc. U. S. Nat. Mus. 46 (1913) 317.

Description of the species.-A large rat belonging to the R.rattus group, but resembling Ratt~~s norvegic'/,~ in general external appearance, except that it has a longer tail, larger ears, and an admixture of grooved spines in pelage of upper surface; eight caudal annuli to a centimeter on middle of tail, two less than in Scandinavian specimens of R. norvegicus,. longest whiskers reaching beyond tip of ears. Color.-Color generally the gray of R. norvegicus but pelage less dense, and a trifle grayer. Skull and teeth.-Skull heavily ossified; temporal fossre smooth; brain case smooth and flattened above, the region included between supraorbital beads narrow and triangular as in R. norvegicus,. anteorbital region broad and elevated; rostrum greatly thickened, nasals broad, obtusely pointed posteriorly, ending opposite frontomaxillary suture; audital bullre flattened

RATTUS

and broadened; teeth similar to those of R. norvegicus. the type description.)

431 (After

Measurements of the type and topotype of Rattus magnVrost1"is (Mearns). Topotype. Type. United States United States National Museum, National Museum. No. 125211. No. 125212. mm. mm. 404 420 Total length 201 195 Tail vertebrre 43 42 Hind foot 56.5 53 Head 18 18 Ear above crown 22 21.5 Ear from notch Skull: Basilar length 39 Occipitonasal length 46 Interparietal breadth 13 Length of internasal suture 18 Greatest breadth of rostJ.\ um 9.3 Length of upper tooth row 7.5

Remarks.-The type and one topotype, in the United States National Museum, were collected at Zamboanga, Mindanao, January 15, 1904, by Edgar A. Mearns. RATTUS RATTUS (Linnreus)

Mus 'rattus LINNlEUS, Syst. Nat. 1 (1766) 83; BLANFORD, Fauna India, Mamm. (1888-91) 406; SANCHEZ, Anal. de la Soc. Espanola de Hist. Nat. 2 (1898) (Luzon, Masbate, Samar); THOMAS, Trans. Zool. Soc. London 14 (1898) 402 (Mindoro and Negros). Epimys 1'atttts HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 30; Bull. U. S. Nat. Mus. 46 (1913) 318 (Luzon, Manila; Panay, Iloilo).

Description of the species.- (From No. 43, E. H. Taylor collection; collected at Manila, P. 1., by E. H. Taylor.) Young female. Moderate-sized rat; body covered with relatively short fur, consisting of a short, soft underfur and numerous longer thickened hairs that are flexible and not spinelike; short fur about 8 millimeters long; the longer hairs about 16 millimeters on rump; on underside the short fur about 4 millimeters; longer hairs about 9 millimeters; vibrissre on snout moderate, reaching tip of ear; ear moderately large, with a dense growth of short hair on inner surface; tail a little longer than head and body, sparsely haired, the hairs as long as width of one and one-half annulations; eleven annulations to a centimeter. Color.-Pelage dull slaty black above, underfur somewhat more grayish black; color below very similar to that of underfur

432

PHILIPPINE LAND MAMMALS

on back; underside of arms somewhat lighter; color on abdomen merges into that of sides; tail uniformly dark; hair on upper surface of feet and hands blackish brown. Skull and teeth.-Rostrum moderate, nasals extending slightly beyond incisors and ending at a point opposite frontomaxillary suture posteriorly; maxillary suture reaches palatine foramina below or slightly anterior to lower notch of ante orbital foramen; palatine foramina short, distinctly in front of a line drawn across anterior edges (alveoli) of first molars; bullre moderately large. Teeth typical of the genus without accessory cusps; tooth rows slightly divergent; first molar with anterior outer cusp (usually well defined in Philippine species) very indistinct. Measurements of Rattus rattus (Linnreus). mm.

Total length Head and body T ail vertebra:! Forefoot Hind foot Skull: Greatest length Zygomatic width Palatine foramina, length Width of brain case Interorbital constriction L.ength of palate from foramina Upper ~ooth row, alveoli Lower tooth row, alveoli Mandible, incisor tip to condyle Mandible bone

337

165 172 18 38

42 19.5 6

16.5 6 9

7.4 7.1 26.5 .

Remarks.-A second specimen in my collection, also from Manila, agrees with the described specimen except that there is a white spot between the forelimbs. This species is comparatively rare in the Islands, but probably occurs in all the larger ports. It is known from Manila, and Iloilo, Panay, from specimens in the United States National Museum; it is reported by Thomas from Mindoro and N egros ; by Sanchez from Masbate and Samar. The species is widely distributed throughout the world and apparently is rivaled only by Rattus norvegicus which seems to dominate it in countries where both occur. It is easily identified by its color.

433

RATTUS

RATTUS MINDORElNSIS

(Thomas)

Mus mindo?'ensis THOMAS, Minutes Proc. Zoo!. Soc. London for June 15 (1897); Trans. Zool. Soc. London 14 (1898) 402; TROUESSART, Cat. Mamm. Suppl. (1904-1905) 369. Epimys mindorensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 29; Proc. U. S. Nat. Mus. 46 (1913) 319.

Description of the species.-(From No. 144608, United States National Museum, Bureau of Science No. 143, collected on Mount Ha1con, Mindoro, November 18, 1906, by Edgar A. Mearns.) Moderate-sized rat, most closely allied to Rattus mindanensis (Mearns) ; fur smooth, soft, and thick, about 12 millimeters long, with a few longer thickened hairs reaching a length of 18 millimeters on rump; longest fur on belly about 8 millimeters long; vibrissre on snout scarcely reaching beyond ears; scrotal pouch large, covered with short fur; ears rather large with extremely fine hairs; tail distinctly shorter than head and body, covered with short sparse setre, the hairs little longer than width of an annulus; nine or ten annulations to a centimeter; forefeet small. Color.-General color of pelage dark grizzled brown resulting from a mixture of the gray slate underfur, each hair of which has a short tip of buff-yellow, and longer, somewhat thickened, brownish black hairs; darker on middle of back and somewhat lighter on sides due to longer buffy tips of hairs; color on sides merges gradually into that of abdomen which is grayish slate with a meager wash of lighter, dirty grayish buff; above foot hairs brownish, the skin below on forefoot dark, still darker on hind feet; scrotal pouch covered with grizzled brownish hair; tail uniformly dark. SkuJ,l and teeth.-Skull somewhat slighter than that of R. mindanensis, the rostrum somewhat shorter; supraorbital beads slight, disappearing on parietals; nasals not extending anterior to incisors, and ending opposite frontomaxillary suture; maxillary suture reaching palatine foramen directly below base of infraorbital foramen; palatine foramina not twice as long as wide, barely reaching level of anterior edge of molars; zygomatic arches slenderer than in R. mindanensis; posterior edge of palate broad, squarish; bullre smaller than in R. rattus. Teeth typical of the genus, without accessory cusps; outer anterior cusp rather strongly defined; tooth rows slightly divergent. 269468-28

434

PHILIPPINE LAND MAMMALS

Measurements of Rattus mindorensis (Thomas). Bureau of Science. No. 143. mm.

Total length Head and body Tail Forefoot, to tip of claws Hind foot, to tip of claws Skull: Greatest length Zygomatic width Nasals, length Palatine foramina Width of brain case Interorbital constriction Length of palate from foramina Upper tooth row, alveoli Lower tooth row, alveoli Mandible to incisor tip Mandible bone

328 180 "148 17 32

Type.

mm.

353 190 163

38 17.8 14.1 6.7 14 6 7.8 6.4 6.3 25 21

32.5 36 19.5 14.5 7 6.1 6.5 26 23.2

.. Tip of tall missing.

Remarks.-The type of the species and four other specimens were taken at an elevation of 1,500 meters on Mount Dulangan, northern Mindoro, by John Whitehead, December, 1895; Edgar A. Mearns collected eleven specimens on Mount Halcon, which touches Mount Dulangan, during November and December, 1906. These I believe are the only recorded specimens. Hollister * has expressed the opinion that this species really inter grades with R. mindanensis (Mearns), which occurs in the lowlands. The specimens mentioned by Whitehead t as being from Canlaon Volcano, N egros, are probably referable to R. mindanensis. The specimen I describe is from the seriâ&#x201A;Źs collected by Mearns, now in the collection of the Bureau of Science, Manila. RATTUS COLORATUS

(Hollister)

Epimys coloratus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 317.

Description of the species.-Related to R. magnirostris, of Zamboanga, but differs in having a smaller, shorter, and broader skull; in being more heavily built, and in having the underparts dark grayish buff; pelage harsh. Color.-General color above and on sides dark russet-brown; hairs gray at base and mixed with numerous brown-tipped

* Proc.

U. S. Nat. Mus. 46 (1913) 319. Trans. Zoo1. Soc. London 14 (1898) 403.

t Thomas,

435

RATTUS

yellowish spines, especially forward; underparts grayish buff, blending with color of upper parts without distinct line of demarcation; pelage below made up of buffy spines and softer hairs of gray; throat and anal regions with numerous, pure ochraceous hairs; hands and feet marked with broad bands of blackish brown, extending along upper surface to base of digits; digits pale buffy white. Skull and teeth,.-Skull smaller than that of R. magnirostris; larger than R. neglectus of Borneo, more "heavily built and more arched, with a relatively wider brain case; supraorbital beads especially heavy, more curved on parietals than in R. magnirostris. (After type description.) MeasU1'ements of the type of Rattus coloratus (Hollister). mm.

Head and body Tail Hind foot without claw Skull: Greatest length Cop.dylobasal length Palatal length Zygomatic breadth Interorbital breadth Mastoid breadth Upper molar row, alveoli Mandible Lower molars

211 222 34 42.8 41.8

24.2 20.4 6.6 16.9 8.1 26.1

7.2

Remarks.-The type, in the United States National Museum, and five cotypes were collected on the Musser plantation near Isabela, Basilan, February 21, 1906, by Edgar A. Mearns. Apparently no specimens have been collected since. RATTUS ROBlGINOSUS (Hollister)

EpiJrnys ?'obiginosus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 318.

Description of the species.-(From No. 32,~ Bureau of Science collection; collected February 7, 1903, in Cagayancillo, by R. C. McGregor and Andres Celestino.) Most nearly related to R. coloratus, of Basilan. Body covered with moderately long fur interspersed with numerous elongate spines that are slender and flexible but can be felt; they appear to be wanting in the rump region where there are numerous elongate hairs reaching a length of 23 to 24 millimeters, while the underfur is 13 to 15 * This specimen is one of the paratype series, but apparently it was not examined by Hollister. The specimen is an adult female.

436

PHILIPPINE LAND MAMMALS

millimeters; fur and spines on middle of back rarely exceed 15 millimeters; belly fur short, spines not or scarcely perceptible to the touch; tail elongate, distinctly annulated (nine to a centimeter), setre of tail short (1.25 millimeters) ; feet normal; ear moderately large; mammary formula, pectoral, 2-2; inguinal, 2-2. Color .-General color above and on sides dark russet-brown, darker TI;ledially, growing lighter laterally; individual hairs mouse gray basally and mixed with slender spines that are brown to blackish distally and grayish cream basally; forehead similar to body in color; ventrally buffy white on throat and chest and somewhat grayish white on abdomen; tail blackbrown, unicolored; forearm and forefeet silvery white with slight brownish admixture; hind feet silvery above with slight brownish admixture. Skull and teeth.-Skull rather massive, the rostrum slightly shortened; supraorbital beads thick, projecting as much horizontally as perpendicularly to skull, curved on parietals forming a pyriform figure; palatine foramina somewhat bowed; bull.re large, somewhat compressed; zygomatic arches heavy, a small ridge continued backward from root of zygoma, above which is another small parallel ridge. Teeth typically murine, similar to but smaller than those of R. coloratus. Measurements of Rattus robiginosus (Hollister) . Bureau of Science No. 82.

mm.

Total length Head and body Tail Height of ear from notch Width of ear Hind foot, dry Skull. Total length Condylobasal length Zygomatic width Width of brain case Nasals, length Interorbital width Mastoid width Diastema Length of upper molar series, alveolar Length of lower molar series Mandible, incisor to condyle

390 215 175 20 16 36 45 43.8 21.2 17 16.5 7 16.8 12 7 7 28

Type.

mm.

404 217 187

43.6 42.4 21.3 7 16.8 7.3 6.3 25.1

437

RATTUS

Remarks.-Thirteen specimens from the type locality are known. They were collected by R. C. McGregor and Andres Celestino on Cagayan Island (Cagayancillo), February 7, 1903. The specimens with one exception are in the Bureau of Science collection on permanent deposit in the United States National Museum. RATTUS MINDANENSIS MINDANENSIS (Mearns)

Mus mindanensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 442. Epimys mdmdanensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 29; Proc. U. S. Nat. Mus. 46 (1913) 318.

Description of the sUbspecies.-(From No. 35, E. H. Taylor collection; collected at Baguio, Benguet, Luzon, July 25, 1923 by E. H. Taylor and John Suarez Wright.) Rather large; fur harsh, rather short, underfur about 8 to 9 millimeters long, longest hairs 22 millimeters long; along middle of back the longer hairs distinctly spinelike; feet normal; tail as long as head and body, setre sparse and short; annuli nine to a centimeter. Color.-Above cinnamon-russet, generally resulting from russet tips of shorter fur and black tips of longer hairs; underpart of fur gray-slate, spinelike hairs white below; fur on belly short, admixed with stiffened hairs which are buff-white; general color white to buff-white, underfur gray-drab but lighter than above; feet generally light buff with a dim, darker stripe; tail uniformly dark. Skull and teeth.-Outer anterior cone of first molar rather well developed with a slight tubercle midway on outer anterior face of molar towards base of median anterior cone; a small outer anterior cone developed on second molar; nasals ending considerably in front of front om axillary suture, extending slightly in advance of incisors; supraorbital beads well developed, continuing to posterior part of parietals; lower anterior molar with two small outer accessory cusps developed between main outer cusps, and one between last two inner cusps; second molar with three small accessory cusps; palatine foramina reaching level of anterior edge of first molars. M eaSU?'ements of Rattus rnindanensis mindanensis (M ea?'ns ) . mm.

Total length Head and body Tail Forefoot Hind foot

400 195 205 19 38

438

PHILIPPINE LAND MAMMALS

Measurements of Rattus mindanensis mindanensis (Mearns)-Continued. Skull: Greatest length Zygomatic width Nasals, length Palat.ine foram~na, length Width of brain case Interorbital constriction Length of palate from foramina Upper tooth row, alveoli Lower tooth row, alveoli Mandible, to incisor tip Mandible, bone

mm.

42 19

14 7

16 5.8 9

7.1 7 27 22.2

Remarks.-The specimen here described is a medium-sized female, and the skull measurements are much less than those for old specimens. The skull of the type measured 47.5 millimeters in basilar length, with the upper tooth row 8 millimeters. The specimens in my collection, ten from Baguio, two from TabIas, and two from Cotabato, vary considerably among themselves. The chief differences noted have to do with the development of accessory molar cusps. Some specimens from the same locality as the described specimen show a total absence of the accessory cones; some only a partial development. The large outer anterior cone, wanting in Rattus pala.wanensis and certain other forms, is invariably present. This species occurs on all the principal islands of the Philippine group and on most of the smaller ones. Hollister (loc. cit.) reports the species from Luzon (numerous localities), Catanduanes, Mindoro, Batan, Romblon, Masbate, Ticao, Panay, Negros, and Mindanao. Hollister * states: I am unable to divide this species satisfactorily intol island races over all this range. In Mindanao and on some of the' southern islands forms are well marked. Doubtless a large series of specimens of comparable age and pelage would make it possible to recognize races on many of the northern islands as well.

It is probable that all references of Rattus neglectus in the

Philippines are really of this form. The type of the species was collected at Todaya, on Mount Apo, Davao, Mindanao, at an altitude of approximately 1,232 meters, by Edgar A. Mearns.

* Proc.

U. S. Nat. Mus. 46 (1913) 318.

439

RATTUS RATTUS MINDANENSIS TABLASI subsp. nov.

Type.-No. 652, E. H. Taylor collection; collected at Odiongan, Tablas, January 18, 1933, by E. H. Taylor; alcoholic skin and skull separate. Diagnosis; description of the type.-Size about that of the usual form of Rattu.s mindanensis; pelage moderately dense, shorter fur about 10 millimeters long; longest hairs about 25 millimeters; tail rather thicker and thumb somewhat more developed than in the typical form. Color.-Pelage above generally russet to cinnamon-russet resulting from admixture of orange-buff tips of shorter fur and longer brown-black hairs; below general color whitish buff with a slight cinnamon wash; throat and pectoral region slightly reddish deep Mikado brown (Ridgway), the hairs being so colored to their bases; color continued on underlips and on underside of snout. Skull and teeth.-Skull and teeth not varying greatly from those of typical Rattu.s mindanensis (Mearns). The following differences seem to prevail: Maxillary suture reaches palatine foramina at a point :rqore anterior, and posterior part of palate shows no small perforations; processes from palate are more nearly parallel than in typ>ical R. mindanensis. MeasUTements of Rattus mimdanensis tablasi subsp. nov. Type. E. H. Taylor E. H. Taylor collection, collection, No. 652. No. 651. rom. mm.

Total length Head and body Tail, tip injured in both Forefoot Hind foot Skull: Greatest length Zygomatic width Nasals, length Palatine foramina, length Width of brain case Interorbital constriction Length of palate from foramina Upper tooth row, alveoli Lower tooth row, alveoli Mandible, to incisor tip Mandible, bone

322 168 154 16 34 42 20.0 15 8 16 7 9 7.2 7.2 27 24.6

325 175 150 18 33 47 22.3 17.2 7.8 17 7.8 9.2 8

7.3 31 36.7

440

PHILIPPINE LAND MAMMALS

Remarks.-The specimens here described were trapped in burrows in a low clay hill near the seashore, some distance from Odiongan, Tablas Island. A single specimen of the typical buffbellied form was captured in the town of Odiongan. This I presumed to be a recent introduction from some other island. The specimens consist of skins (in alcohol) and skulls separate. ALEXANDRINUS GROUP

Two forms are referred to the alexandrinus group (grisiventre group of Bonhote and Trouessart); these are Rattus kelleri (Mearns) and R. zamboangm (Mearns). Specimens of this group are rather difficult to distinguish from members of the rattus group. They are, however, somewhat smaller and the mam:rnre of the Philippine forms are two pairs pectoral, one pair abdominal, one pair inguinal. In Philippine rattus group the arrangement appears to be two pairs pectoral, two pairs inguinal. Key to fJhe species of the alexandrinus group. a '. No spiny hairs; skql1 high in parietall region; skull length, 41 milIi-

meters ____ ___________________, _______ ___________ _______ _____ ___ ___ R_ kelleri (Mearns) (p. 440).

a". Numerous spiny hairs; skull small, narrow with a shorter, narrower rostrum than R. kellM'i; skull length, 38.5- millimeters. R. zamboang33 (Mearns) (p. 442). RA TTUS KELLERI

(~Iearns)

Mus kelleri MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 444. Epimys keU:eri HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 28; Proc. U. S. Nat. Mus. 46 (1913) 319.

Description of the species.-Apparently a member of the alexandrinus group, but without spiny hairs in the pelage; whiskers extending beyond tip of ears; caudal annulations ten to a centimeter on middle of tail; tail uniformly colored, longer than head and body; ears large, scantily haired; mammre four pairs; two thoracic, one abdominal and one inguinal. Color.-Similar to R. mindanensis, but darker and more reddish above; upper parts Mars brown, thickly mixed with black; underparts yellowish white; nose sepia, forearm hair brown; forefoot with a band of fur of hair brown above; narrow band on hind feet, extending to base of toes; tail grayish brown. Skull and teeth.-Skull similar to that of R. mindanensis, but higher in the parietal region, with nasals and rostrum narrower and audital bullre smaller and narrower; teeth not peculiar.

441

RATTUS Measurements of the type of Ratt.U8 kelleri (Mearns). mm.

Head and body, length Tail Hind foot 118

120'

177

198 40 1,4

122

126

Batan I.

PHILIPPINE ISLANDS

ÂŽ

R~ttus kellepi

@ Rattus zamboangre

ÂŠ Rattus benguetensis Rattus datae

4J .c?

'"" .....

"\ 12

~ "~

I.> 0'

'<:

120

FIG. 22.

122

124 0

126

Distribution of four species of Rattus in the Philippines.

Head Ear, above notch Skull: Basilar length Occipitonasal length Interparietal breadth from outer edge of beaded margins

50 22 33,5 41 13.7

442

PHILIPPINE LAND MAMMALS

Measurements of the type of Rattus kelleri (Mearns)-Continued. Skull-Continued. Length of internasal suture Greaites,t breadth of rostrum Length of upper molar row

mm.

7.5

Remarks.-Only the type, in the United States National Museum, is known. It was collected at Davao, southeastern Mindanao, July 20, 1904, by Edgar A. Mearns. The species is named for Mr. Fletcher L. Keller, of Davao. RATTUS ZAMBO ANGlE (Mearns)

Mus zamboang33 MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 4'43. Epimys zamboang33 HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 30; Proc. U. S. Nat. Mus. 46 (1913) 319.

Description of the species.-Pelage of upper parts coarse, spiny; of feet and underparts short, fine, and soft; whiskers long, reaching shoulders; caudal annuli eleven to a centimeter on middle of tail; tail uniform in color, longer than head and body; ears of medium size, nearly naked except on edges. Color.-Upper parts cinnamon-Tusset mixed with gray and black; sides bordered by clear cinnamon; underparts pale drabgray; feet whitish drab, with a strip of hair brown on upper surface; tail grayish brown; whiskers black; anterior face of incisor teeth ochraceous-yellow. Skull and teeth.-Skull small, narrow, with a short narrow rostrum, brain case convex above, and with supraorbital bead bending evenly outward posteriorly without a marked prominence at outer extremity of frontoparietal suture; zygomatic arches very slender; temporal fosS're ridged; audital bullre small, very narrow, and rather high; dentition weak. (After type description. ) Measurements of the type of Rattus zamboa,ng33 (Mearns). mm,

Head and body Tail vertebrre Hind foot Ear above crown Skull: Basilar length Occipitonasal length Interparietal breadth Length of internasal suture Greatest width of rostrum Length of upper tooth row

150 165 36 15 32 38.5 13.6 12.5

6.7

443

RATTUS

Remarks.-The type, No. 125279 in the United states National Museum, was collected at Zamboanga, Mindanao, January 20, 1904, by Edgar A. Mearns. No other specimen is known. DATlE GROUP

Two species are referable to this group, both, so far as is known, confined to the highlands of northern Luzon. These forms differ from other larger rats in the total 31bsence of the supraorbital beads. Key to the species of the datre group. a 1. Smaller; upper parts mixed gray and ochraceous-buff; below light buff, with a strong wash of ochraceous; tail, 130 millimeters; skull, 33.3. R. benguetensis (Hollister) (p. 443). a'. Larger; upper parts darker colored than above; below naphthaline yellow; ta.il, 160 to 175 millimeters; skull, 36 to 37. R. datre (Meyer) (p. 444). RAT.T US BENGUETENSIS (Hollister)

Epitmrys benguetensis HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 323 .

Description of the species.-Like Rattus dat;;e but smaller and lighter colored; under parts conspicuously different, light buff with a strong tendency toward ochraceous. Color.-The color of the type in worn pelage is mixed gray and ochraceous-buff above~ hairs broadly dark gray at bases, tipped with ochraceous-buff; cheeks and sides much lighter, almost pure warm buff, underfur of a lighter shade than on back; entire underparts light buff, with a strong wash of ochraceous, hairs very light gray at bases; feet pinkish buff with a faintly darker streak along upper surface; tail light brown, unicolored; whiskers dark brown and black (in dat;;e these are mixed with white) . Skull and teeth.-Except for its smaller size the skull does not differ appreciably from that of Rattus 'dat;;e. Last upper molars lack anterointernal supplementary cusps, but that this is a somewhat variable character in dat;;e is shown by one specimen in which this cusp is present on one side and absent on the other. (After Hollister.) Measurements of the type of Rattus benguetensis (Hollister). mm.

Head and body, from dry skin Tail Hind foot without claw, moistened

130 130 32

444

PHILIPPINE LAND MAMMALS

Measurements of the type of Rattus benguetensis (Hollister)-Continued. Skull: Condylobasal length Zygomatic breadth Interorbital breadth Breadth of brain case Length of nasals Upper molar row, crowns Mandible Lower tooth row

mm.

33.3

16 5.3 15.3 32

6.2 20.9

6.2

Remarks.-Hollister states that the skull characters of this species do not differ markedly from those of Rattus datre (Meyer), but the color characters are especially different. Only the type, No. 145790 in the United States National Museum, is known. It was collected May 1, 1907, at Camp John Hay, Baguio, Benguet, Luzon, by Edgar A. Mearns. RATTUS DATlE (Meyer)

Mus dattJ3 MEYER, Abhand. und Berichtte des Konig]. Zool. Mus. Dresden 7 (1898-99) 25. Mus chrysocomus THOMAS, Trans. Zool. Soc. London 14 (1898) 403. Epimys dattJ3 HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 28; Proc. U. S. Nat. Mus. 46 (1913) 323.

Description of the species.-Closely related to Rattus callitrichus J entink (= Mus chrysocomus), but differing in having no supraorbital beads present on the skull; the pelage is more softly colored than that of R. callitrichus. Measurements of Rattus dattJ3 (Meyer). [Measurements in millimeters.]

United States National Museum No.

Head and body _________________________________________ _ Tail vertebra! _________________________________________ . _ Hind foot with claws __________________________________ __ _ Skull.: Condylobasallength ________________________________ _ Zygomatic breadth _________________________ .. _______ .

Lยฃ

andible---------- - -------------------------------Maxillary tooth row ________________________________ _ Mandibular tooth row .. ____________________________ __

145809

145797

147798

162 148 36

150 175 41

150 160

34.2 16 21.2 6.1 6.5

37 17.5 23.3 6.7 6.5

36.3 17.3

lU 6.5 6.6

RATTUS

445

Remarks.-The measurements of this species are those published by Hollister. * The color of Rattus dattE differs from that of R. benguetensis in that it is darker, the belly being naphthaline yellow. The type, in the Dresden Museum, was collected on Mount Data, Lepanto, northern Luzon. There are eight specimens of this species in the United states National Museum, collected by Edgar A. Mearns. A single specimen collected by J. Whitehead on Mount Data is in the British Museum. CONCOLOR-EPHIPPIUM 'GROUP

This group is composed of the smaller rats, some of which are scarcely larger than mice. It has been possible to separate island forms based, in several cases, merely on average measurements. The following key probably canpot be wholly depended upon for identifying the forms, as the variations between young and old sp~cimens may be nearly as great as the average differences between species. I~ the locality is considered little difficulty should be experienced in identifying the species. The arrangement in the key does not necessarily show relationships. Key to Rattus species of the concoZor-ephippiwm group. Larger species; total length, 270 to 295 millimeters; tail, 130 to 155. b 1. Wool hairs, 15 millimeters; pelage slightly spiny; teeth relatively small; dark smoky gray to blackish on back. (N egros.) R. negrVrvuso (Thomas) (p. 446). b 2. Wo()l hairs, 10 millimeters; pelage distinctly spiny; teeth larger. c'. Upper molar series, 5.9 millimeters; skull, 34; hind foot, 29; above tawny-olive mixed with black-brown. (Southern Mindanao.) R. todwyensis (Mearns) (p.448). c'. Upper molar series, 5.4 millimeters; skull length, 36.2; foot, 29.6; tawny-olive to russet on back mixed with blackish. (Mindoro .and Tablas.) ............................... R. vigorat~ÂŁs (Hollister) (p. 449). cÂˇ. Upper molar series, 5.2 millimeters; skull length, 32; hind foot, 25.5; eleven to twelve annuli to a centime~er; mandible, 18 millimeters; dark umber above with mixture of cinnamon and black. (Central and northern Luzon.) .... R. q~ÂŁerceti (Hollister) (p. 450). c'. Yellowish rufous mixed with black; skull, 29 (?) millimeters; hind foot, 28; upper molal' series, 5.2. (Palawan.) R. luteivent~'is (Allen) (p. 452). a 2. Medium-sized species; total length, 250 to 260 millimeters; tail, 125 to 130.

0, \

* Proc.

U. S. Nat. Mus. 46 (1913) 323.

446

PHILIPPINE LAND MAMMALS

b '. Above russet mixed with black; thirteen annuli to a centimeter; skull, 29 millimeters; hind foot, 27; upper molar series, 5.2. (Davao, Mindanao.) .................................... R. pantarensis (Mearns) (p. 453). b 2. Central upper parts chocolate; sides drab-gray overlaid by tawnyolive tips; annuli of tail ten to a centimeter; skull, 30 millimetel's; hind foot, 25.5; upper molars, 5.1. (Mount Apo, Mindanao.) R. vulcani (Mearns) (p.454). b 3 . Above tawny-olive mixed with black-brown; skull, 30.8 millimeters; hind foot, 25.5; upper molars, 5.2. (Ba~ilan.) R. basilanus (Hollister) (p. 455). a 3 . Small species; total length, 225 to 235 millimeters; tail, 110 to 125. b 1. Pelage spiny. c'. Dark brownish drab above; rostrum relatively heavy; skull, 28.9 millimeters; hind foot, 24; upper molars, 5.1 millimeters. (Catanduanes.) ............................ R. leucophmatus (Hollister) (p.456). c~. Dark umber with mixture of cinnamon; skull, 30.6 millimeters; hind foot, 23.7; upper molars, 5.6. (Mount Mayon, southern Luzon.) .................................... ........ R. mayonicus (Hollister (p. 457). c 3. Bright Sudan brown, finely lined with blackish; skull, 30 millimetel's; hind foot, 23.5; upper molar row, 5.3. (Cagayancillo, Cagayan Island.) ............................ R. ornatulus (Hollister) (p. 457). b 2. Pelage not spiny; russet above, mixed with black; skull, 30.5 millimeters; hind foot, 25; upper molal' series, 5. (Northern Luzon.) R. calcis (Hollister) (p. 458). RATTUS NEGRINUS (Thomas)

Mus ephippium negrinus THOMAS, Trans. Zool. Soc. London 14 (1898) 403. Epimys negrinus, HOLLISTER, Proc. BioI. Soc. Washington 24 (1911) 90; Philip. Journ. Sci. ยง D 7 (1912) 29; Proc. U. S. Nat. Mus. 46 (1913) 320.

Description of the species.-Similar to Rattus ephippium but rather larger, much longer and softer fur; wool hairs about 15 millimeters and overhair about 18 millimeters in length on back; tail nearly as long as head and body. Color.-General color dark smoky gray, almost blackish along middle of back, lightening to buffy or yellowish on sides; belly color not sharply defined, the hairs slaty at base, yellowish white at tip; hands and feet silvery whitish above; tail uniformly brownish, or slightly paler below. Measu?'ements of the type of Rattus negrinus (Thomas). mm.

Head and body Tail Hind foot, moistened Skull: Length, lambda to nasal tip Greatest breadth

155 135 26.5 30.3 16

447

RATTUS

Measurements of the type of Rattus negrinus (Thomas)-Continued. Skull-Continued. Nasals, length Nasals, breadth Interorbital breadth

rom.

13.1 3.6

5.2

120'

118

124

122

126

Batan Is

PHILIPPINE ISLANDS

® Rattus negpinus @Rattus luteiventr>is

Lubang~

"'.

....

MINDOIlO

Busuan8a~

.2luo ~c .. ; Bupi.., 'ORomblon

ab~5i~ya~~~ao

Culion~

tjA~~~O

aPANAY

Cuyo GOull)apan

Cagayan

'~.

(-/'"')

®/

tsu

ISJ'NEGROS~'O' ~ 12. ~

AALA_N

"Cagayan Sulu

120

FIG. 23.

122

124 0

126

Distribution of six species of Rattus in tbe Philippines.

Palate length Diastema Anwrior palatine foramen Length of upper molar series Lower jaw, bone only

16 9.7

6.5 5 19.7

448

PHILIPPINE LAND MAMMALS

Remarks.-The type was collected in Negros at an elevation of 2,011 meters on Canlaon Volcano, by John Whitehead. One paratype was also collected. There are three specimens in the United States National Museum also taken in Negros, two of which are from Canlaon Volcano. RATTUS TODAYENSIS

(Mearns)

Mus todayensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 445. Epimys todayensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 30; Proc. U. S. Nat. Mus. 46 (1913) 321.

Description of the species.-(From No. 78, E. H. Taylor collection; collected at Saub, Cotabato, April 24, 1923, by E. H. Taylor.) A large species of the group, the pelage consisting largely of moderately stiffened spines, about 9 to 10 millimeters in length on rump, longest hairs reaching a few millimeters farther; ears moderately large; tail as long as head and body with twelve annuli to a centimeter; vibrissre on snout extending well beyond ears. Color.-Upper part tawny-olive finely mixed with blackbrown; somewhat lighter brown on sides; below grayish white with a slight cinnamon wash; tail dark purplish brown; feet drab-gray with a darker stripe. Skull and teeth.-Skull high, with brain case expanded in parietal and contracted in frontal region; supraorbital bead prominent; temporal fossa ridged; rostrum stout; nasals narrowed and bluntly pointed posteriorly; audital bullre high and elongate; postpalatal notch wide; anterior surface of upper incisors ochraceous. Measurements of the type of Rattus todayensis (Mea.rns). mm.

Head and body Tail Hind foot Head Ear above notch Skull: Basilar length Occipitonasal length Interparietal breadth, outer edge of beaded margins Greatest breadth of rostrum Length of upper molar row

138 145 29

4'2 18 27 34

13 6 5.9

Remarks.-The types and four topotypes were collected at Todaya, at an altitude of 4,000 feet on Mount Apo, southeastern Mindanao, July 11, 1904, by Edgar A. Mearns. Mearns states

449

RATTUS

that in general appearance the species suggests the rats of the Rattus surdus group which, however, are much more spiny. My collection contains several specimens of the species from Saub, Cotabato, Mindanao (preserved in alcohol). RATTUS VIGORATUS (Hollister)

Epimys vigoratus

HOLLISTER,

Proc. U. S. Nat. Mus.

(1913) 321.

Description of the species.-(From No. 656, E. H. Taylor collection, collected near Odiongan, Tablas Island, January 15, 1923, by E. H. Taylor.) The largest of the group. Pelage strongly spined, soft underfur being relatively scant; pelage on back about 9 to 10 millimeters long, a few hairs 1 or 2 millimeters longer; foot large; vibrissre on snout reaching slightly beyond ears; ears rather large; tail with setre on middle of tail as long as width of two annuli; about eleven annuli to a centimeter near base of tail; tail longer than head and body. Color.-Upper parts rather tawny-olive with an admixture of darker color on the longer hairs near middle line of back; on sides more brownish; underparts dirty grayish white, perceptibly washed with cinnamon; above, feet silvery white; tail brown, unicolored. Skull and teeth.-Upper profile of skull forming a continuous curve; a slight depression at posterior tips of nasals which end on level with frontomaxillary suture; supraorbital beads prominent, diverging as slightly curved lines, continued only faintly on posterior part of parietals; palatine foramina moderate, reaching backward as far as beginning of first molar; postpalatal notch broad; bullre moderately large. Teeth without distinguishing characteristics; anterior external cusp of first molar smaller than internal; apparently no accessory cusps developed above; on lower molars a tendency for three or four small accessory cusps to develop. ,

Measurements of Rattus vigoratus (HoLLister). mm.

Total length Head and body Tail Hind foot Ear above notch Skull: Greatest length Condylobasal length 269453-29

282 129 153 29.6

15 36.2 32

450

PHILIPPINE LAND MAMMALS

Measurements of Rattus vigoratus (Hollister)-Continued. Skull-Continued. Zygomatic width Palatine foramina Diastema Palate from foramina Interorbital constriction Width of brain case Upper molar series, alveoli Lower molar series, alveoli Mandible, incisor tip to condyle Mandible, bone

mm.

16 6.1 9

7.2 5.4

14 5.4 5.3 23.2 20

Remarks.-I am unable to differentiate the Tablas specimens from Rattus vigoratus described from Mindoro. The large size of the skull of that species is slightly exceeded in these specimens, but in general they agree in essential details. The nine specimens in my collection were obtained by digging into small burrows in the sod along the edges of small ditches in the open fields near Odiongan, Tablas. The species appears to be very numerous if one may judge by the large number of burrows encountered. Hollister states that the species is related to Rattus todayensis (Mearns). The type, in the United States National Museum, was collected on Mount Halcon, Mindoro, about halfway to the summit, November 15, 1906 by Edgar A. Mearns. Four other specimens were obtained at various localities on the island. RATTUS QUERCETI (Hollister)

Epimys querceti

HOLLISTER, Proc. BioI. Soc. Washington 24 (1911) 90; Philip. Journ. Sci. ยง D 7 (1912) 30; Proc. U. S. Nat. Mus. 46 q913) 319.

Description of the 8pecies.-(From No. 22, E. H. Taylor collection; collected on the summit of Mount Santo Tomas, July 19, 1923, by E. H. Taylor ~nd John Suarez Wright.) This is one of the medium-sized species of the conGolor-ephippium group; pelage distinctly spiny and the underfur dense; average length of pelage on rump about 10 millimeters, a few longer hairs reaching a length of 13 to 15 millimeters; much shorter in middle of back and on shoulders; vibrissre on snout barely reaching tip

451

RATTUS

of ear; tail longer than head and body, eleven to twelve annuli to a centimeter; setre on tail not longer than width of two annuli at middle of tail. Color.-General color above dark umber resulting from a mixture of cinnamon and black hairs; sides paler; below, dirty grayish white; feet grayish white above; tail dark brown, slightly lighter below; when a specimen is held, with the tail pointing away, a very unusual green sheen is visible on back and sides. Skull and teeth.-Skull without striking characters to distinguish it clearly from other members of the group; brain case widened; supraorbital beads curving on parietals, continuing to end of parietals; parietal foramen elongate, extending to anterior edge of first upper mo,l ars; teeth not distinctive. M easur ements of Rattus querceti (Holliste?路). mm.

Total length Head and body Tail Hind foot, dry Skull : Greatest length Condylobasal length Zygomatic width Palatine foramina Nasals Diastema Palate from foramina Interorbital constriction VVidth of brain case Upper molar series, alveoH Lower molar series, aveoli Mandible, condyle to incisor tip Mandible, bone only

272 130

142 25.5

32 30.6 15.85 6

11.4 8 6

14 5.2 5

20 17

Remarks.-These measurements agree very well with those of the type. However, there are c.e rtain differences between this and specimens from Manila and from Los Banos, Laguna Province, Luzon. Here the tail averages shorter (135 millimeters) in the largest specimens; there are usually twelve annuli to a centimeter. I am unable to detect the greenish sheen in the latter specimens.

452

PHILIPPINE' LAND MAMMALS

The type and fifty-five other specimens wer:e collected at Haight's Place, Benguet, Luzon, at an elevation of approximately 2,153 meters, August, 1907. Later a large series of specimens was collected by Mearns and Carroll Fox at Manila. Hollister states: The form is readily distinguishable from Epimys ea.lJcis by its larger siz.e and darker coloration. It differs from E. negrinus much as does E. ealeis, having smaller teeth and a somewhat slender skull. It also has much c~arser pelag~. It is mor~ closely related to true E. ephippium than are t.he other described Philippine forms of the group, but its large size and long tail are specific characters of suffici,e nt value. RATTUS LUTEIVENTRIS (Allen)

Mus Luteiventris ALLEN, Bull. Am. Mus, Nat. Hist. 28 (1910) 14. Epimys luteiventr-is HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 29.

Description of the species.-Size medium; pelage spiny, strongly so above, weakly so below. Color.-Above, dull yellowish rufous, slightly varied with black-tipped bristles; below olive-buff, hairs grayish brown at extreme base; bristles on dorsal surface silvery white with blackish tips; on ventral surface yellowish white without black tips; tail pale brown, unicolored, almost naked. Measurements of the type of Rattus luteiventris (Allen). mm.

Total length, from skin Head and body Tail Hind foot Skull: Zygomat.ic blleadth Length of nasals Upper molar series

293 163 130

28 14 12

5.2

Remarks.-The type, No. 29717 in the American Museum of Natural History, was collected at Iwahig, Palawan, by John R. White. Two paratypes were taken, both young adults. They agree perfectly in color. In my collection are two specimens, both adult females, which presumably belong to this species. They are distinctly smaller than the type (according to the few published measurements) and may represent a new species. I append the measurements of the two specimens (No. 54, collected at Sir J. Brooke Point, Palawan, and No. 55, at Puerto Princesa, Palawan).

453

RATTUS

Measurements of Rattus luteiventris (Allen). E. H. Taylor E. H. Taylor collection. collection. No. 54, No. 65, Sir J. Brooke Puerto Point. Princesa. mm. mm.

Total length Tail Head and body Hind foot Ear from notch Skull: Greatest length Condylobasal length Zygomatic breadth Palatine foramen, incisive Diastema Nasals Palate from foramina Interorbital constriction VVidth of brain case Upper molar series, alveoli Lower molar series, alveoli Mandible, incis~H; to condyle Mandible, bone only

210 110 100 23.5 14.5

223 115 113 25.5 16

29 24.5 13 5.2 6.8 10.5? 5.6 5 13.2 4.8 5 18 15.2

The annuli on tail are thirteen to a centimeter. The mammary formula is two pairs pectoral (the second pair considerably behind axilla), and two pairs inguinal. No. 54 was found in a burrow in an old termite hill. Three suckling young were taken at the same time. RATTUS PANTARENSIS

(Mearns)

Mus pantarensis M.EARNs, Proc. U. S. Nat. Mus. 28 (1905) 448. Epimys pantarensis HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 29; Proc. U. S. Nat. Mus. 46 (1913) 32'2.

Description of the species.-Small; tail uniformly colored, as long as head and body, scantily coated with short stiff hairs, with annuli thirteen to a centimeter near base of tail; pelage short, thickly mixed with coarse flattened channeled spines; ears moderate, nearly naked; whiskers short, not reaching tip of ears. Color.-Color of upper parts russet, finely mixed with black; underparts grayish cream-buff; ears and tail pale drab; feet grayish white, with a stripe of sepia on upper surface; whiskers brownish black.

454

PHILIPPINE LAND MAMMALS

Skull and teeth.-Skull considerably arched in parietal region; rostrum short and stout; audital bullre short, broad, high; temporal fossre ridged; nasal bones truncate posteriorly; dentition weak, not otherwise peculiar. (From the type description.) Measurements of the type of Rattus pantarensis (Mea?·ns). mm.

Length, head and body Tail, verlebrre Hind foot Head Ear above noteh Ear above erown Skull: Basilar length Oeeipitonasal length Interparietal breadth, from outer margins Length of internasal suture Greatest breadth of rostrum Length of upper molar row

125 125 27 36 18 14 24.5 29 12.6 10 5 5.2

Rernarks.-The type, in the United States National Museum, was collected by Edgar A. Mearns, at Pantar, Mindanao, at 1,907 feet elevation, September 4, 1903. The type is an adult female. Only the type is known. RATTUS VULCANI

(Mearns)

Mus vulcani MEARNS, Proe. U. S. Nat. Mus. 28 (1905) 446. Mus vulcanLapicis MEARNS, Proe. U. S. Nat. Mus. 28 (1905) 447. Epimys vulcani vulcani HOLLISTER, Philip. Journ. Sei. § D 7 (1912) 30.

Epimys vulcani apicis HOLLISTER, Philip. Journ. Sei. § D 7 (1912) 30.

Epimys vulcani HOLLISTER, Proe. U. S. Nat. Mus. 46 (1913) 321.

Description of the species.-Similar to Rattus concolor but larger; also resembling R. todayensis, but smaller, darker, with less spiny hair; tail not strictly unicolored, but paler on underside; whiskers reaching tip of ears; caudal annuli ten to a centimeter on tail at middle; ears large, nearly naked; tail usually about equal to length of head and body, but frequently longer. Color.-Central area of upper parts dusky chocolate; sides drab-gray overlaid by tawny-olive tips of long hairs; muzzle hair brown; underparts whitish gray, perceptibly washed with cinnamon; tail dusky drab above, drab-gray below; feet whitish drab, without a dusky stripe on upper side; ears drab; whiskers brownish black. Skull and teeth.-Skull light, with prominent superciliary frontoparietal beads; brain case arched above; rostrum <i.e-

455

RATTUS

pressed; temporal fossa slightly ridged; nasal bones rather narrow, pointed posteriorly; audital bullre higher, narrow, and elongate; teeth normal. (After the type description.) Measurements of the type of Rattus vulcani (Mea?Âˇns). mm.

Head and body Tail Hind foot Head Ear, above notch

133 125 25.5 35 14

Skull: Basilar length, Hensel â&#x20AC;˘ 14.3 Occipito nasal length 30 Interparietal breadth, from outer edge of beaded margins . 12.5 Length of internasal suture 11.5 Greatest width of rostrum 6 Length of upper tooth row 5.1 8

Probably a misprint for 24.3 millimeters.

Remarks.-Hollister, who examined Mearns's material, has regarded the form designated R. vulcani apicis as being unworthy of subspecific rank. The type of the species and seven paratypes were collected at 7,600 feet altitude on Mount Apo, Davao, Mindanao, by Edgar A. Mearns, June 26, 1904. The types and paratypes are in the United States National Museum. RATTUS BASILANUS

(Hollister)

Epimys basaa,nus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 322.

Description of the species.-This small rat is related to Rattus todayensis, of Mindanao, but is smaller, with smaller skull and teeth, the underparts less buffy, more yellowish. Measurements of the type of Rat.tus basilanus (Hollister). mm.

Head and body, dry skin Tail Hind foot, without claw

130 130 25.5

Skull: Greatest length Condylobasal length Nasals Interorbital constriction Palatal length Upper molar row, alveoli Mandible Lower tooth row, alveoli

30.8 28.9 10.9

4.9 16.2 5.2 17.2 5.2

456

PHILIPPINE LAND MAMMALS

Remarks.-Hollister states that this is a smaller insular race of Rattus toclayensis. Three specimens are known from Basilan and one from J 010. They were collected February, 1906, by Edgar A. Mearns. The J 010 specimen is represented only by a skull, but according to Hollister it appears inseparable from the Basilan skulls, but suggests that a series of skins might show differences. A very young specimen in my collection from Abung Abung, Basilan, may belong to this species. RATTUS LEUCOPHlEATUS (Hollister)

Epi11'lYs leucophreatus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 320.

Description of the species.-Small rat, with very spiny pelage; it differs from all the Philippine members of the group in its pale grayish brown coloration and relatively heavy skull, with massive rostrum. Color.-Upper parts, from head to rump, dark brownish drab, a mixture of sulphur yellow spines with brown tips and softer hairs with dark gull gray bases and cinnamon-buff tips; cheeks, sides, and limbs paler, more grayish; hands and feet pale buffy white; underparts pale buffy white with brighter yellowish wash on groin and at base of tail; hair of belly and chest faintly gray at base. Skull and teeth.-Skun most resembling that of Rattus querceti of Luzon, but with decidedly heavier, wider rostrum and smaller, more rounded bulke. Teeth essentially those of E. querceti. Measurements of the type of Rattus leucophreatus (Hollister). mm.

Head and body Tail Hind foot, without claws Skull: Greatest length Condylobasal length Zygoma.tic breadth Breadth of brain case Nasals Breadth of rostrum Interorbital breadth Upper molar row, alveoli Mandible Lower tooth row

114 112 24 28.9 27.4 14.2 13.2 10.2 5.4

4.9 5.1 16.5 5

457

RATTUS

Remarks.-The type, an adult female in the United States National Museum, was collected by D. B. Mackie, May 27, 1909, at Bagamanoc, Catanduanes Island. No other specimen is positively known. Two skins in my collection, without skulls, from Polillo Island, may belong to this species. RATTUS MAYONICUS

(Hollister)

Epi1nys mayonieus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 319.

Description of the species.-A medium-sized rat externally like Rattus querceti but slightly smaller, with more yellowish on sides, hips, and underparts; pelage more spiny than that of R. querceti. Skull and teeth.- The skull differs from that of R. querceti in its much narrower, almost parallel-sided, nonspatulate nasals; rounder, higher audital bullre, and larger teeth. Measu~'ements

of the type of Rattus mayonwtGs

(Hollis~er).

mm .

Head and body', dry skin Tail Hind foot, moistened, without claws Skull: Greatest length Condylobasal length Zygomatic breadth Breadth of brain case Interorbital breadth Length of nasals Upper molars, alveoli Mandible Lower molars, alveoli

117 112 23.7 30.6 28.6 14.6 13.2

4.7 11.2

5.6 17.2 5.4

Remarks.-The type, in the United States National Museum, was collected on Mount Mayon, Albay Province, Luzon, at an elevation of approximately 1,230 meters, June 5, 1907, by Edgar A. Mearns. Only the type has been collected. RATTUS ORNATULUS

(Hollister) I

Epimys O?'natulus HOLLISTER, Proc. U. S. 路Nat. Mus. 46 (1913) 322.

Descritption of the species.- A small rat, a member of the concolor-ephippium group; like R. pantarensis (Mearns), of Mindanao, but feet white, not grayish buff, and underparts yellow, not buff; smaller and brighter colored than R. h拢teiventris (Allen), of Palawan.

458

PHILIPPINE LAND MAMMALS

Color.-General color of upper parts bright Sudan brown, finely lined with blackish, made up of brown-tipped yellowish spines and of soft hairs gray at base and ringed and tipped with bright ochraceous and blackish; brighter on rump and hips; sides lighter, yellowish tawny; underparts straw yellow, the hairs of middle breast and belly pale gray at bases; hands and feet thinly covered with white and very pale cream hairs. Skull amd teeth.-Similar to Rattus pantarensis (Mearns), but skull with longer, more slender rostrum and nasals. (After the type description.) Measurements of the type of Rattus ornatulus (Hollister). mm.

Head and body, from dry skin Tail Hind foot, moistened, without claws Skull: Greatest length Condylobasal length Zygomatic breadth Upper molar row, alveoli Mandible Lower molar row

118 119 23.5 30 27.2 14.2 5.3 16.1 4.9

Remarks.-The single type specimen was collected by R. C. McGregor and A. Celestino on Cagayancillo, Cagayan Islands, February 8, 1903. The type, an adult female with slightly worn teeth, is in the collection of the Philippine Bureau of Science now on deposit at the United States National Museum. This is the second species which this small island, isolated in the Sulu Sea, has yielded. RATTUS CALCIS (Hollister)

Epitmys calcis HOLLISTER, Proc. BioI. Soc. Washington 24 (1911) 89; Philip. Journ. Sci. ยง D 7 (1912') 28; Proc. U. S. Nat. Mus. 46 (1913) 320.

Description of the species.-(From No. 142, Bureau of Science collection; collected May 16, 1907, at Baguio, Benguet, Luzon, by Edgar A. Mearns.) (One of the paratype series.) A small rat of the concolor-ephippium group; pelage soft, the spines wanting or at least not thickened or stiffened; on rump, fur generally about 9 millimeters long, the longer hairs 14 or 15 millimeters long; tail a little longer than head and body, about thirteen annuli to a centimeter.

459

APOMYS

Color.-Above, generally russet mixed with black, growing lighter on sides, becoming dark buff low on sides, and gray cream on belly; a faint yellowish wash present ("some specimens almost primrose yellow") ; feet dirty white; tail brownish, generally unicolor. Skull and teeth.-Very similar to those of R(J;ttus querceti; the incisive palatine foramina usually shorter and narrower. Measurements of

Ratt~~s

calcis (Hollister). mm.

Total length Head and body Tail Hind foot Skull: Greatest length Condylobasal length Zygomatic breadth Palatine foramen Diastema Nasals Palate from foramina Interorbital constriction VVidth of brain case Upper molar series, alveoli Lower molar series, alveoH Mandible, incisors to condyle Mandible, bone

235

110 125 25 30.5 27.5 15 5.6 8

11 5.8 4.5 13.5 5 5 19.4 16.2

Remarks.-This species may be separated from R. querceti by its smaller size and the much softer pelage, due to absence of stiffened spines; the spinelike hairs are actually present but are not at all rigid to the touch; the color is lighter than typical R. querceti from Baguio and the north, but not as light as specimens of the latter from Manila and Los Banos. The type of R. c(J;lcis was collected by E. A. Mearns at Baguio, Benguet, Luzon, May 8, 1907, at an elevation of approximately 1,537 meters. Some twenty-two other specimens one of which is described here were collected at the same place. Genus' APOMYS Mearns

Apomys MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 455; Philip. Journ. Sci. ยง D 7 (1912) 33.

HOLLISTER,

General external form as in R(J;ttus; pollex slightly less rudimentary, with a rather broad nail; soles naked; forefoot with

460

PHILIPPINE LAND MAMMALS

five tubercles, four of which are placed at base of digits, the fifth at outer side of base of palm, the two posterior tubercles being relatively much larger than in Rattus,' hind foot with six tubercles; tubercles are not paired as in Rattus but are more widely separated and oblique to each other; eyes relatively smaller; whiskers very long~ reaching to abdomen; body thickly furred above and below, with pelage long and soft; ears large, thin, and naked; tail scantily haired, longer than head and body, sometimes white at end; annuli distinctly visible; mammre, two pairs, inguinal. Skull and teeth.-Skull elongate with a pronounced rostral portion; no superciliary bead; incisive foramen less than half the length of diastema between incisor and lateral teeth; upper ano lower incisors yellow on anterior face, and without groove; first upper molar 6-tuberculate in three series, with three primary and three subsidiary cusps, the anterior series comprising one large anterior tubercle and one small posterointernal tubercle; middle series comprising one large median and two posterolateral subsidiary tubercles, a large anterior tubercle with a prominent anterointernal and two small lateral tubercles, and a single large posterior tubercle joined to the anterolateral tubercles by low lateral ridges, the external one obsolete; third upper molar small, its crown a roughened tubercle with a minute anterointernal groove and tubercle; two anterior lower molars quadrituberculate with a posterior step; posterior lower molar with a notched anterior cusp and rounded posterior tubercle; anterior lower molar with an anterior cusp but shallowly notched; with age an anterior step is developed in first and second lower molars. (A.fter Mearns.) The type of the genus is Apomys hyloccetes Mearns. All the species known are Philippine. Key to the Philippine species of Apomysl Mearns. A white spot on chest. b \ Ears and soles of feet pale flesh; brain case inflated; head and body, 120 millimeters; skull, 22; sixteen caudal annuli to a centimeter. (Mount Apo, Mindanao.) ____________________ A. hylocmtes Mearns (p. 461). b '. Ears and soles of feet purplish slate; brain case more flattened; head and body, 122 millimeters; skull, 24; twelve caudal annuli to a centimeter. (Mount Apo, Mindanao.) A. insignis ins ignis (Mearns) (p. 462). b". Similar to A. insignis but the brain case is narrower and more elongate; white spot may be absent. (Mount Bliss and Mount Malindang, Mindanao.) _______________ , A. ~nsignis bard'l.LS (Miller) (p. 463). 'a 2. Without white spot on chest. a

APOMYS

461

b '. Above russet, below pale fawn; head and body, 114 millimeters; skull, 22; thirteen caudal annuli to a centimeter. (Mount Apo, Mindanao.) ..... _______________ .. __________ . _______________ A. petr:eus Mearns (p. 465). b'. Above wood brown; below paJe smoke-gray with a buffy cast; head and body, 143 millimeters; skull, 35. (Haight's Place, Benguet, Luzon.) ____________________________________________________________ A. major Miller (p. 466).

Body and head ochraceous-buff; below similar but slightly rufous; head and body, 86 millimeters; skull, 22.5. (Baguio, Benguet.) A. musculus Mearns (p. 468). b 路 . Intermediate in size between A. majo?' and A. ?nusculus; upper part cinnamon-brown, below buffy gray; 'washed with pale cinnamon; teeth smaller than in the other species. A. ?nicrodon Hollister (p. 469). b

APOMYS HYLOCtETES Mearns

Apomys lIylocmtes MEARNS, Pl'oc. U. S. Nat. Mus. 28 (1905) 456; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 34; Proc. U. S. Nat. Mus. 26 (1913) 327.

Description of the species.-A forest mouse; general characters those given for the genus; head large and rounded: tail scantily and very finely haired above and below, rings distinctly visible throughout, hairs between verticelli extending to tip of tail, 2 millimeters long; caudal annuli about sixteen to a centimeter; tip of tail white for about 20 millimeters; soles of feet entirely naked; tubercles of hind foot more crowded and the two innermost much larger than in Apomys insignis. Color.-Above, mummy brown; underparts cinnamon-fawn with a white area in center of chest; back and especially top of head finely mixed with black; end of muzzle and throat drabgray; outer surface of forelimbs mouse gray; forefeet whitish; hind feet pale drab-gray above, becoming whitish on toes; toes tufted at base of claws; claws horn color; soles flesh color tinged with drab-gray; tail purplish slate above, pale drab-gray below, white all round at tip; whiskers mixed white and brownish black; naked ears smoke gray; wrists drab-gray below; ankles seal brown; pelage of upper parts blackish slate at base, of underparts (except in white area) slate-gray. (There is some variation recorded in the young adult: Pelage of back sepia mixed with black; underparts smoke gray with a cinnamon-white area on chest.) Skull and teeth.-Brain case inflated; upper profile of skull straight and slightly declining anteriorly, strongly arched posteriorly; audital bullre rounded and very high; rostrum slender, elongate; nasals greatly expanded anteriorly, sharply compressed in the middle, and tapering to a slender point ending at base opposite frontomaxillary suture.

462

PHILIPPINE LAND MAMMALS

Measurements of the type of Apomys hylocmtes Mearns. mm.

Total length Tail vertebra! Hind foot Head Ear above crown Skull: Basilar length Occipitonasal length Interparietal breadth Interorbital breadth Length of internasal sutUl'e Length of upper molar series

250 135 31 35 15 22.3 31 14 5 11.5 5.9

Remarks.-The type and seven paratypes were collected by Edgar A. Mearns at an elevation of approximately 1,846 to 2,150 meters on Mount Apo, southern Mindanao. All are in the United States National Museum. The largest and smallest adult specimens listed by Mearns have head and body measurements of 121 and 115 millimeters, respectively. This species may be distinguished from Apomys insignis by the paler feet and ears; sales of feet flesh color instead of purplish slate; feet broader with larger tubercles. APOMYS INS IGNIS Mearns

Apomys insignis MEARNS, Proc. U. S. Nat. Mus. LISTER, Philip. Journ. Sci. ยง D 7 (1912) 34.

(1905) 4'5 9; HOL-

Two forms of this species are recognized; namely, the typical form from Mount Apo, and Apomys insignis bardus Miller, from northwestern Mindanao. Both are high-mountain forms. The latter was given specific rank by Miller but except for a slenderer rostrum, they agree. Forms of this species have langel' tails than other members of the genus. APOMYS INSIGNIS INSIGNIS (Mearns)

Apomys insignis insignis

HOLLISTER,

Proc. U. S, Nat.

Mus.

46 (1913)

327.

Description of the subspecies.-A medium-sized species; characters those of the genus; caudal rings twelve to a centimeter; tubercles of hind feet small and widely separated, largest less than 5 millimeters long; longest whiskers 60 to 65 millimeters long. Color.-Upper parts mummy brown; underparts cinnamonfawn with a white area in middle of chest; top of head and back very finely mixed with black; end of muzzle and throat

463

APOMYS

drab.gray; foreleg seal brown externally; forefeet white; hind feet whitish drab above; toes white, tufted at base of claws; naked soles and ear purplish s1ate; tail purplish slate, paler below, but not distinctly bicolored; some specimens with white tips to tail; whiskers mixed black and white; wrists clay color below; ankles seal brown; pelage of upper parts slate color at base, of underparts, except in white area, slate路gray; iris dark brown; claws horn color; white area of underparts varies from a small stripe between forelegs to a broad irregular ventral band ending near vent. (The stripe may be interrupted.) The young are mouse gray, becoming brown as they grow older. Skull and teeth.-Brain case flattened as compared with other species of Apomys.. anterior profile of skull flat and level; ros路 trum thickened; nasals evenly tapering to an acute base even with frontomaxillary suture; audital bul1re less inflated and rounded; teeth large. (After Mearns.) Measurements @j the type of Apomys insignis insignis Mea1'ns. mm.

287 176

Total length Tail Hind foot Head Ear from crown Ear from notch Skull: Basilar length Occipitonasal length Interparietal breadth Interorbital breadth Length of internasal suture Length of upper tooth row

37 16 20 24

32 13.5 5 12 6.1

Remarks.-Fourteen specimens were collected on Mount Apo between, approximately, 600 and 1,800 meters elevation, by Edgar A. Mearns, between June 20 and July 12, 1904. Most of the specimens are in the United States National Museum. APOMYS INSIGNIS BARDUS

(Miller)

Apomys ba1'dus MILLER, Proc. U. S. Nat. Mus. 38 (1911) 402; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 34. ApomY8 ifnsignis bardus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 327.

Description of the subspecies.-(From No. 146, Bureau of Science collection; collected May 31, 1906, on the summit of Mount Bliss, Mindanao, by Edgar A. Mearns.} Specimen, old male. Size very similar to that of Apomys insignis insignis, but

464

PHILIPPINE LAND MAMMALS

skull narrower and brain case less inflated; fur long and soft without trace of spines, from 12: to 16 millimeters long on rump, 5 to 7 millimeters on belly; longest whiskers 60 millimeters; ears large, thin, essentially naked; tail very long, 1.43 times length of head and body (1.6 times in insignis but distinctly longer in bardus than in other Philippine species) ; hair on tail very short, with annuli plainly visible, about thirteen to a centimeter. Color.-Above mummy brown to wood brown, somewhat lighter on sides of head, behind ears, and along sides of body; top of head similar to back; underfur slate-gray; belly with a wash of dirty cream, underfur more ashy to drab-slate; no whit2 spot on breast; wrist nearly brown, foot cream white; hind foot dirty whitish drab above, toes cream; tail slaty, slightly lighter below, distinctly so on proximal half but not bicolored as in A. major. Skull and teeth.-Brain. case strongly flattened; anterior part flat and nearly level; nasals widened anteriorly, tapering posteriorly to narrow points; audital bulke moderately inflated; area of skull above posterior part of bullre somewhat inflated; teeth large, posterior molar relatively smaller than in A. major; posterior part of palate fragile, bounded posteriorly by a slight ridge distinctly posterior to third molars. The teeth of this specimen are so worn that the typical generic characters are wanting. Measurements of Apomys insignis bardus (Miller). United States National Museum. Type. mm.

Head and body Tail Hind foot, dry Skull: Condylobasal length Total length Zygomatic breadth Interorbital constriction Nasal Diastema Mandible Maxillary tooth row, alveoli Mandibular tooth row, alveoli

113 162 33 28.8 14.2 4.8 11.8 7.8 16.8 5.8 5.2

Bureau of Science No. 146. mm.

115 161 32 28.2 31 14.3 5 11.6 7.5 16.7 5.6 5.2

465

APOMYS

Remarks.-Mearns collected the type and eight other specimens of this species on Mount Blis.s, Mindanao; the described specimen is one of this lot. Miller points out the following characteristic differences from other forms: Brain case differs in its narrower, more-elongated form; greatest breadth contained about two and one-half times in upper length of the skull instead of about two and one-fourth times or less. Miller states that "the color is so essentially like that of Apomys insignis Mearns as to require no special description." However, the specimen here described is very different in the ventral coloration. Other specimens presumably of this species were taken on Mount Malindang at an elevation of approximately 2,770 meters. The type and paratypes were from an elevation of approximately 1,760 meters on Mount Bliss. The type is in the United States National Museum. I

APOMYS PETRlEUS Mearns

Apomys petrtEus MEARNS, Proc. U. S. Naot. Mus. 28 (1905) 458; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912') 34; Proc. U. S. Nat. Mus. 46 (1913) 327.

Description of the sp'e cies.-A small species of rock mouse; characters those of the genus; tail moderately hairy above and below, dark colored above to naked tip; caudal annuli about thirteen to a centimeter; hairs of tail about length of two caudal rings on upper surface; soles of feet naked except at heel. Color.-Upper parts russet or Mars brown; underparts pale fawn, colors blending on sides; muzzle, throat, and inner aspect of forelimbs smoke gray; outer surface of forelimbs mouse gray; wrists whitish below; feet smoke gray, fading to whitish on toes; claws horn color; tail pale, purplish brown above, flesh color below; whiskers mixed white and brownish black; naked ears drab-gray; iris dark brown; hair of upper parts slate-gray, at base of underparts drab-gray. (There is distinct variation evident in other specimens.) Skull and teeth.-Brain case swollen in parietal region; upper profile of skull straight and .slightly declining anteriorly, strongly arched posteriorly; audital bullre high and rounded; rostrum moderate; nasals expanded apically, tapering gradually to a pointed base opposite frontomaxillary suture. 269463-80

466

PHILIPPINE LAND MAMMALS

Measurements of the type of Apomys petrreus Mearns. mm.

Total length Tai,! Hind foot Length of head Ear above crown Skull: Basilar length Occipitonasal length Interparietal breadth Intero路r bital breadth Lenglth of internasal suture Length of upper molar series

231 120 30 37

16 22 29.5 14 5.1 11.5 5.8

Remarks.-The type and one paratype were collected by Edgar A. Mearns at an elevation of approximately 2,340 meters on Mount Apo, Mindanao, June 30, 1904. The paratype is a female and differs somewhat in color, being markedly darker. The type is No. 125245, in the United States National Museum. APOMYS MAJOR Miller

Apomys ma,jor MILLER, Proc. U. S. Nat. Mus. 38 (1911) 402; HOLLISTER, Philip. Jotltn. Sci. 搂 D 7 (1912) 34; Proc. U. S. Nat. Mus. 46 (1913) 328.

Description of the species.-(From No. 24, E. H. Taylor collection; collected July 19, 1923, at the summit of Mount Santo Tomas, Baguio, Benguet, Luzon, by E. H. Taylor and John Suarez Wright.) Body elongate, moderately slender; tail with annuli distinct, twelve to a centimeter; tail essentially naked, the small hairs from 1 to 1.2 millimeters long on the greater part, slightly longer towards tip; those on tip extending beyond tip 2 millimeters; annuli distinctly visible; fur soft, without spines, longest hairs on rump from 12 to 15 millimeters long; on belly 7 to 8 millimeters; whiskers much elongate, the longest 55 millimeters; ears large, thin, rounded above, naked except for sparse, very short hairs on upper side; hind foot elongate, soles naked; small tufts of hair at base of claws; posterior inner pad on foot greatly elongated, not reaching as far forward as edge of posterior outer tubercle; hallux well developed, claws on this and outer toe only slightly smaller than on other toes; pollex with a well-defined nail. CoZor.-Pelage wood brown above with a slight admixture of dark hairs; distinctly of lighter shade on sides of head and body; top of head darker than back, below creamy buff; under-

467

APOMYS

fur, both above and below, nearly slate-gray; tail black above, underside creamy white, the colors distinctly delimited; wrists and ankles hair brown; forefeet dull whitish with a darker area on back of foot; hind feet whitish mixed with brown above; toes white; whitish below. Skull and teeth.-Skull elongate with a somewhat globular brain case; no lateral ridge, but edge of skull slightly angular laterally in postorbital and supraorbital region; nasals elongate, greatly narrowed posteriorly but not forming points; rostral portion of skull extending beyond incisors a distance greater than distance between incisors and incisive foramen; latter half as long as diastema between incisors and molars; posterior part of palate somewhat concave, delicate, almost lacelike, terminating in a slight, raised, double ridge; anterior part on a level with third molar; bullre inflated but smaller than in similar-sized species of Ra.tt1.ts; area of skull posterior to and above the bullre slightly inflated; a prominent bead anterior to tooth row below anterior edge of infraorbital vacuity. Upper tooth rows diverging somewhat posteriorly; teeth of described specimen much worn and not showing the characters of genus; posterior molar greatly reduced, decidedly smaller than third lower molar. MeasurementJs of Apomys major Miller. rMeasurements in millimeters.] United States

E. H. Taylor collection.

~ational

Museum. Type. Head and body _____ __ __ _ _ __ ____ _ __ __ _ _ __ _ _ _ _ _

Tail_ _______ __ _____ ______ ______ __ ___________ _

143 144

Hind foot , dry_ _____ ______ ___ _________________ 32.2 Skull: Condylobasallength_ ___ _________________ __ 34 . 8 Total length _______ __ ___ ___ _______ ______ ___ __ ___ ___ _ Zygomatic breadth _ __ ____ ____ _ _ __ _ __ ______ 17 . 2 Interorbital constriction _____________ ____ _ _ 6 Depth of brain case at middle _____________ _ 10 __ ____ ___ _____ ____ ___________ _ 15 Diastema __ ___ ______ ____ __ ___________ ___ _ 10.2 andible, excluding teeth _________________ _ 21.4 Maxillary tooth row ____ __________________ _ 6.2 Mandibular t ooth row ____________________ _ 6 ~asall e ngth

~o.

23.

146 142 34 35 .2 38.5

---------5.8 10 14 9. 8 19 7 6.4

~o .

24.

144

- ---- - - --35 30 . 51 35 17 5.8 10 . 2 13 . 3 8 .8 18 . 4 7 6.5

~o.

25.

141 132 31. 5 34 37 17 6 .8 10 .2 1<1.6 9 19

~ 6.2

Remarks.-The three specimens in my collection listed above were captured in traps at night on the summit of Mount Santo

468

PHILIPPINE LAND MAMMALS

Tomas (above the .rest house). The type with three other specimens came from Haights-in-the-oaks, Benguet, Luzon, at an elevation of approximately 2,150 meters, which is apP'r oximately the height of Mount Santo Tomas. The type, collected by E. A. Mearns, is No. 151513 in the United States National Museum. APOMYS MUSCULUS Miller

Apomys musculus MILLER, Proc. U. S. Nat. Mus. 38 (1911) 403; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 34; Proc. U. S. Nat. Mus. 46 (1913) 328.

Description of the species.-Smallest species of the genus. In general characters similar to other members of the genus. Color.-Entire body ochraceous-buff, clear and rather light on sidse; cheeks and underparts tinged with ochraceuos-rufous and slightly clouded with black above; face slightly grayish, feet buffy gray, nOit conspicuously different from underparts; tail obscurely bicolored, brownish above, whitish below. Skull and teeth.-Skull smaller, with a relatively larger and more globular brain ease, more abruptly constricted interorbital region, and a more nearly vertical anterior border ,o f plate forming outer wall of infraorbital canal; teeth similar to other species but smaller. (After Miller.) Measurer/tents of the type of Apomys muscuDus Miller. mm.

Head and body Tail, imperfect Hind foot, dry Skull: Condylobasal length Zygomatic breadth Interorbital constriction Depth of brain case Nasal Diasltema Mandible Maxillary tooth row Mandibular tooth row

86 89 111.6

22.6 13 4'

8 9

6 13.8

4.6 4.2

Remarks.-The type and one paratype were collected at Baguio, Benguet, at an altitude of approximately 1,500 meters, May 6, 1907, by Edgar A. ยฅearns. These are the only specimens known.

469

INSULlEMUS APOMYS MICRODON Hollister

Apomys microdon HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 327.

Desc'ription of the species.-A medium-sized species; differs from most of the known forms in having very small teeth. Other characters those of the genus. Color.-Upper parts cinnamon-brown; sides lighter, almost tawny-olive; ~nderparts buffy gray, strongly washed with pale cinnamon; hands, feet, and tail brown, the tail apparently un icolored. Skull and teeth.-The skull (a broken specimen) is apparently much as in Apomys insignis from Mindanao in general sh3;pe; teeth smaller than in any species of Apomys except the little Apomys musculus of Luzon. (After the type description.) Measurements of the type of ApomY8 microdon Hollister. mm.

Head and body Tail Hind foot Skull: Palatal length Interorbital breadth Upper tooth row Mandible Lower tooth row

106 135 25.5 16.1 4.8 5.2 15.5 4.9

Remarks.-The type, No. 145770, in the United States National Museum, was collected by D. B. Mackie at Biga, Catanduanes, May 8, 1909. This is the only specimen known. Genus INSULlEMUS novum

Resembles a small species of Ratt~ts .. ears rather large; vibrissre reaching beyond shoulders; pollex somewhat more developed than in Rattus, with a broad, slightly curved nail; three anterior plantar tubercles nearly equal and forming a triangle; that below thumb largest, with a very small, rounded tubercle projecting anteriorly; outer tubercle next in size, extending slightly behind the other; hallux much reduced, truncate, ending in 9rounded pad surmounted by a narrow rounded nail about onethird width of tip; digit does not reach as far as plantar tubercles; other toes nearly equal in length, third and fourth slightly longest; three anterior tubercles of hind foot rounded and somewhat flattened, outer one at base of fifth toe extending distinctly farther back than the one at base of second toe; that at base of hallux with a small lateroposterior rounded projection behind which is inserted a very small tubercle which is

470

PHILIPPINE LAND MAMMALS

almost contiguous with large posterior tubercle; latter only slightly longer than broad, the anterior edge of which extends somewhat farther forward than posterior outer tubercle; tail annulated but rat~er indistinctly so, ending in an elongate brush of hair; two pairs of inguinal mammre; no pectoral mammre. Skull fragile, zygomatic arches slender; slight supraorbital beads; coronoid process of mandible practically wanting, represented only by a prominent angle. Three upper and three lower molars; pattern of cusps very complicated; first upper molar elongate, with three major series of cusps separated by longitudinal grooves; outer groove continuous with a similar groove on second tooth; inner groove continuous even to third molar, which lacks the outer groove; two minute anterior cusps on first molar followed by a transverse series of three large cusps; then follow two other transverse series of three, the median cusps largest and in , advance of the two outer, each of which is followed by a small secondary cusp; a small lateroposterior cusp; second molar rounded with two larger median cusps and a series of three smaller on each side; a small posterior cusp; third molar rounded, reduced in size, with three large inner, and three smaller outer cusps; all upper molars three-rooted; lower molars with two elongate grooves, the outer running the length of the series, the inner less deep, continuous on two anterior teeth; first lower molar with about nine maj or and about five secondary cusps or elevations; second tooth with seven major cusps and four secondary cusps; third with three ma)or and three secondary cusps,; upper incisors with a slight lateral groove. I am uncertain as to the exact relationship of this genus. The type is InsultEmus calamianensis sp. nov. INSULlEMUS CALAMIANENSIS sp. nov.

Type.- (No. 104, E. H. Taylor collection; collected at sea level near Minuit, Busuanga Island, Calamianes, April 18, 1918, by E. H. Taylor.) Skin in alcohol; skull separate. Diagnosis; description of the type.-Characters those given for the genus; head typically mouselike; vibrissre especially long, reaching to behind ,s houlders; pelage soft, without spines; most of the hairs averaging about 8 millimeters in length, while a few hairs may be extended 1 or 2 millimeters longer; on belly hair about 4 millimeters long; hands and feet covered above with short, somewhat stiffened, sparse hair which does not lie close

471

CARPOMYS

to skin, but hairs stand erect and free; tufts of hair about claws elongated; ears moderately large, outer surface haired, hairs becoming extremely thin and scattered near tip; a few scattered short hairs on inner surface; pads on tips of toes strongly truncate and rounded; tail with dim scaled annulations, scales and setre so placed as to show strong diagonal arrangement; individual scales rounded and not imbricate; setre near base of tail about 1.5 millimeters long, growing gradually longer toward tip, reaching a length of 4 milljmeters; hair on extreme tip 10 to 12 millimeters in length. Color.-Surface of pelage rich buff-brown, underfur slatedrab; sides of head and snout cream-buff to cream-white; belly creamy white; pads under feet yellowish. Skull and teeth.-As in generic description. Measurements of

Ins~tla3mus

calamianensis 811. nov. mm.

Head and body Head Tail, without hair Hind foot and claw:~ Ear from lower base Length of longest hair in moustache Skull: Greatest length Zygomatic width Breadth of brain case Interorbital constriction Diastema Upper tooth row Lower tooth row Mandible with incisors

30 130 Z2.5 22 46

28.5 16

13.5 6

7.5 4.5 4.5 20

Remarks.-The holotype was found in a tree stump on the seashore at Minuit, Busuanga. It is an adult female, apparently with suckling young judging from the size of the mammre. The specimen is in alcohol, and the pelage probably has become somewhat faded. Genus CARPOMYS Thomas

Carporrvys THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 161; Trans. Zool. Soc. London 14 (1898) 406; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 33.

Form more or less as in arboreal genera of Muridre. Fur thick and woolly; pollex with a large nail; other digits, including

472

PHILIPPINE LAND MAMMALS

the nonopposable hallux, with claws; tail long, well haired; skull with rounded brain case and short face; supraorbital region without .sharp ridges or overhanging ledges; interparietal large; zygoma. root without any forwardly projecting plate, front edge vertical or even concave; anterior palatine foramina fairly long; bullre large; teeth not unusual; incisors smooth in front; first and second molars as in Rattus, but each with an additional lamina, formed, apparently, by the normal posterior lamina being doubled round on itself; last molar normal, so that laminar formula is 4-3-2; both second and third upper molars have well-marked anterointernal supp,l ementary cusps; in lower jaw first molar has an additional lamina in front, and both it and second molar hav:e well-marked posterior supplementary cusps; while the last named has in addition an anteroexternal one. (After T.homas.) The type is Carpomys melanurus Thomas. The genus is not known outside of northern Luzon. The known species are arboreal. Key to the species of Carpomys Thomas.

Tail black; body fur extending some distance on base of tail; teeth large ........................................................ C. melanurus Thomas (p. 472). a ' . Tail brown; body fur extending a shorter distance on base; teeth smaller than in C. melanurus ............................ C. phmurus Thomas (p. 474). 1.

CARPOMYS MELANURUS Thomas

Carpomys melanurus THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 162; Trans. Zoo!. Soc. London 14 (1898) ' 407, pI. 34, fig. 2 (entire animal) ; pI. 36, fig . .6 (skull); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 33; Proc. U. S. Nat. Mus. 46 (1913) 326.

Description of the species.-Size that of the common rat; fur soft, thick, and woolly. Color.-General color deep fulvous, coarsely lined with black; undersurface and undersides of limbs dull yellowish white, bases of hairs slate; ears of medium size, well haired, dark brown, nearly black; limbs to wrists and ankles furred and colored like body; metapodials brown mesially, laterally and on digits white; tail longer than head and body, its basal inch or two thickly furred like the body and of the same color; the rest closely covered with shining black hairs 5 to 7 millimeters long, entirely hiding the scales; not specially tufted at tip. Skull and teeth.-N asal broad in front, abruptly narrowing behind; interorbital region narrow, broader in front than behind;

473

CARPOMYS

traces of ridges mounting on to top, and approaching to within 2 millimeters of each other in middle line; palatal foramina parallel-sided, attaining their greatest width anteriorly; palate ending opposite edge of third upper molar. 120'

118

124

122

126

Balan Is

PHILIPPINE ISLANDS

® Bullimus luzonicus @ Buillimus bagobus

Lub.ng~..

~ iJ

~uque C:\ l " ; Buri.,,,,,

MINDORO

blo

' 6usuangas;~

Culion~.

,'lTic.o

r i AA

'~o

cu~:ANAY~'~ '

~Ouma"an

rnu 0

Cagayan Is.fN [GROS

PALAWAN

C'\

1li!omblon~ •

~ Sibuyan

Deatanduanes ...

"'Cagayan 3ulu

118

FIG. 24.

120

122

124'

126

Distribution of BuUi?n'US, Insulalmu8, CCllTpomY8, and Trypho7Yli/Js in the PhiIippinee.

Teeth broad and heavy; incisors broad, slightly flattened in front in old specimens, dark yellow above, rather more whitish below; molars very broad and large, their combined length exceeding that of palatal foramina. (After Thomas.)

474

PHILIPPINE LAND MAMMALS

Measurements of the type of Carpomys melanurus Thomas. mm.

Head and body Tail Hind foot, moistened Skull: Basal length Basilar length Greatest breadth Nasals, length Nasals, breadth Interorbital breadth Interparietal length Interparietal breadth Anterior zygoma root, length Palate, length from henselion Diastema Anterior palatine foramina, length Anterior palatine foramina, combined width Leng1th of upper molar series Lower jaw, condyle to inciso!l" tip Lower jaw, bone only

197 211 34.2 39.3 36.3 24.5 16 5.6 5.2 6.1 11 5.1 19 12 7.8 2.7 8.8 29.5 27

Remarks.-Whitehead, quoted by Thomas, says: The black-tailed Carpomys differs much from the next species, C. phreurus both in size, colour, and length of fur; in fact it has externally the appearance of Batomys. On Monte Data, where both these new forms were obtained, it was more numerous than either Ba,tomys or the next species.

Thomas figures this species in color. The types are in the British Museum of Natural History. Distribution.-Highlands of central northern Luzon. CARPOMYS PHlEURUS Thomas

Carpomys phreurus THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 162; Trans. Zoo1. Soc. London 14 (1898) 408, pI. 34, fig. 1 (entire animal); pI. 36, fig. 7 (skull); HOLLISTER, Philip. Jour-n. Sci. ยง D 7 (1912) 33; Proc. U. S. Nat. Mus. 46 (1913) 338.

Description of the species.-Smaller than Carpomys melanurus, but quality of fur and general color almost identical; ears rather smaller, less thickly haired, and not prominently black; belly hairs dull buffy white to their roots, not slaty basally; tail with body fur not extending on to its base more than in ordinary rats, more thinly haired than C. melanurus, so that the scales, which average thirteen to a centimeter, are visible through the hairs; in color it is uniformly dark brown, occasionally approaching black, but never the deep shining black of C. melanurus.

475

CARPOMYS

Skull and teeth.-Nasal less expanded anteriorly than in C. melanurus, and less abruptly tapering posteriorly; interorbital space comparatively broad and parallel-sided, the rudimentary ridges not approaching each other on top; palatal foramina pointed in front, gradually broadening backward; palate ending opposite front of third upper molar. Incisors comparatively narrow; teeth in general much smaller than in C. melanurus; molars small in proportion to size of animal, of the same essential structure as those of C. rnelanll1"Us. (After Thomas.) MeaSUTements of the type of Carpomys phamru8 Thomas. mm.

Head and body, stJ:etched Tail Hind foot, moistened Skull: Basal length Basilar length Greatest breadth Nasals, length Nasals, breadtl;l Interorbital breadth Interparietal length Interparietal breadth Anterior zygoma root, length Palate length from henselion Diastema Anterior palatine foramina, length Foramina, combined width Length of upper molar series Lower jaw, condyle to incisoT tip Lower jaw, bone only

195

178 31 36

33 23 14.5

4.6 6.1

4.8 13 3.8 16.4 11

7.7 2.8 6.1 26.6 24

Remarks.-Whitehead, quoted by Thomas, says: The brown-tailed Ca1'pornys was somewhat raTe on Monte Data. The Igorrotes used to hunt for the various Rats on Monte Data in small parties accompanied by their dogs, and spent most of the day at this-to them--curious occupation. The animals that I saw captured weTe dug out from among the roots of trees by the aid of spears and choppers. The flat table-top of Monte Data is much burrowed by various species of Rodents; the Igorrotes, unlike the Kina Balu Dusans, not trapping Rats for their food.

Dist1路ibution.-Highlands of central northern Luzon. The type, in the British Museum, was collected by Whitehead on MoU):it Data, northern Luzon.

476

PHILIPPINE LAND MAMMALS

Genus TRYPHOMYS Miller

Tryphom;ys MILLER, Proc. U. S. Nat. Mus. 38 (1911) 399; HOLLISTER. Philip. Journ. Sci. .ยง D 7 (1912) 31.

External form murine, tail about as long as body without head; hind foot with outer digits so reduced that neither extends beyond level of base of three median digits; fur of back coarse and harsh, tips of shorter hairs tending to curve forward, giving pelage a peculiar scorched aspect. Skull and teeth.-Skull rather short and broad, with widely spreading zygomata and unusually large anteorbital foramina; outer alveolar wall swollen into a noticeable protuberance at level of first upper molars; bony palate terminating in a broad median ridge bounded by lateral vacuities, teeth robust, their structure not differing materially from those of Rattus. (After Miller.) TRYPHOMYS ADUSTUS Miller

Tryphorntys adustus MILLER, Proc. U. S. Nat. Mus. 38 (1911) 399. pI. 20; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 31; Proc. U. S. Nat. Mus. 46 (1913) 325.

Description of the specie/;,.-Size of a small rat; tail about as long as body without head, coarsely and conspicuously annulated, eight annuli to a centimeter at middle; thumb with a small adpressed nail; sole naked throughout, the five tubercles well developed, fifth large; inner toe extending to base of three middle digits, outer toe about to middle of inner; claws well developed,. those on hind foot largest; formula of mammre: Pectoral, 2-2; ventral, 1-1 i inguinal, 2-2 = 10. Color.-Back and sides a coarse grizzle of wood brown and black, the brown everywhere in 'excess, particularly on sides; median dorsal region abruptly more grizzled than sides, the recurved hairs ,s howing grayish in certain light; bases of hairs slate-gray streaked by the lighter gray of very slender grooved bristles; head and face as back but with a grayish cast; underparts and cheeks buffy white, dulled by the slate-gray undercolor i feet like upper parts but with a decided tinge of broccoli brown; tail uniform dark brown, the minute hairs brownish to silvery. Skull and teeth.-Skull heavily built; outer wall of anteorbital foramen ,so greatly developed that orifice is more conspicuous in dorsal view than from front; margin of interorbital region beaded, the ridge continued back to near middle of parietal; palate much narrower between anterior molars than posteriorly; incisive foramina long and narrow, their hinder borders reach-

477

MUS

ing level of root of first upper molar; audital bullre moderate, evenly inflated; mandible deep and short, the coronoid , process rising noticeably above condyle. Teeth are so worn in the type that their true characters are scarcely discernible. They do not differ greatly from those of Rattus; upper tooth rows strongly converging. Measurements of the type of Tryphomys rulustus Miller. mm.

Read and body Tail Rind foot, dry Skull: Condylobasal length Zygomatic breadth Interorbital constriction Mastoid breadth Depth of brain case Nasal length Diastema Mandible Maxillary tooth row, alveoli Mandibular tooth row, alveoli

174 150 33 37.2 20.4 5.2 15 13 15.2 10.4 24 7.6 7.6

Remarks.-The type, in the United States National Museum, is an old female with worn teeth. No others are known. The specimen was collected by Edgar A. Mearns at Haight's Place, Benguet, Luzon, July 26, 1917, at an elevation of approximately 2,460 meters. Genus MUS Linnreus Mus LINNJEUS, Syst. Nat. 10th ed. 1 (1758) 59 (part.); MILLER, Proc. BioI. Soc. Washington 23 (1910) 59; HOLLISTER, Proc. BioI. Soc. Washington 7 (1912) 31. Musculus RAFINESQUE, Proces. des Decoul'. Somat. (1814) 13. Leggada GRAY, Charlsworth's Mag. Nat. Rist. 1 (1837) 586. Drymomys TSCHUDI, Fauna Peruana (184'4 ) 178. Nannomys PETERS, Monatsber. K. Preuss. Akad. Wiss. Berlin (1876) 480. Acromys TROUESSART, Bull. Soc. D'Etudes Sci. d' Angers 10 (1881) 133. Pseudoconorruys RHOADS, Proc. Acad. Nat. Sci. (1896) 531. Dryomys PHILIPPI, An. Mus. Nac. de Chile 14 (1900) 20.

By many authors this name has been used for the combined group now recognized under the two genera Mus and Rattus. The genus as now accepted is defined by Miller as follows: In general like Epimys (Rattus) but mechanical scheme of molars modified by the elongation of crown of anterior tooth until it forms the main portion of tooth row; m l with three roots, its crown decidedly

478

PHILIPPINE LAND MAMMALS

longer than those of the two succeeding teeth combined, its first lamina much distorted by displacement backward of inner tubercle into line with outer and middle tubercles of second lamina; m S small and tending to disappear, in some species wiJthout trace of first lamina; upper incisors much compressed, each set at such an angle that a subapical notch is normally cut in its outer side by action of lower tooth.

The type is Mus musculus Linnreus. Two species are recognized in the Philippine fauna. Key to the Philippine species of Mus Linna3us. a'. Rich chestnut.-brown; feet and tail uniform in color with body. M. castaneus Waterhouse (p. 478). a 2. Generally slate-gray, with upper tips of hair brownish; tail brownish above, whitish below........................ M. commissarius Mearns (p. 479). MUS CASTANEUS Waterhouse

Mus castaneus WATERHOUSE, Ann. & Mag. 'Nat. Hist. I 12 (1843) 134; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 32; Proc. U. S. Nat. Mus. 46 (1913) 337.

Description of the species.-Head long, rostrum slender; ears medium; smaller than Mus musculus, rostrum slenderer, tail proportionately longer. Color.-Rich chestnut-brown above, body paler below; feet and tail uniform in color with body (these colorsl noted in spirits). (After Waterhouse.) Measurements of the type of Mus castaneus Waterhouse. mm.

Head and body Tip of snout to ear Length of tarsus and digits Ear Tail

65.5 16.8 15.8

9.5 76.2

Remarks.-The type was collected in the Philippines, exact locality unknown, by H. Cuming. Elera reports specimens from Luzon and Cebu. Mearns failed to obtain .specimens of the species. In my collection, and in that of the College of Agriculture, Los Banos, are specimens, preserved in alcohol, which appear to belong to the Waterhouse species. They have the "rich chestnut brown" color, somewhat paler below. To what extent the alcohol is responsible for the "rich chestnut" color of the type, I cannot say; but in the case of certain Pol ill 0 specimens of which I have skins and alcoholic specimens the alcohol has changed the color of the latter from a "hair brown" of the typical Mus commissarius to a color one may define as chestnut.

479

MUS

Hollister * suggested the possibility of Mus casta.neus Waterhouse and M. commissanus Mearns being identical. I am of the same opinion. Many of my specimens lack the third molars. In Nos. 99 and 100, from Polillo, third molars are present above and below. These are young specimens; in 726, from Los Banos, the third molar is absent in the right mandible; in 71.9, from Los Banos, the third molar is present in only the right upper molar series. The third molar is lacking in some specimens in the United States National Museum. t MUS COMMISSARIUS Mearns

Mus commissarius MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 449; MILLER, Proc. BioI. Soc. Washington 24 (1911) 38; HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 32; Proc. U. S. Nat. Mus. 46 (1913) 325; ELERA, Contr. Fauna Filipina (1915).

Description of the species.-Smaller than Mus musculus Linmeus; color similar but with underparts darker; whiskers not reaching tips of ears; tail bicolored. Color.-Upper parts hair brown, strongly washed with raw umber; underparts drab, faintly washed with cinnamon; tail dusky brownish above, soiled grayish white below. Skull and teeth.-Similar to those of Mus musculus, but smaller, with relatively smaller audital bullre; nasals ending obtusely, considerably in advance of base of premaxillaries, instead of acutely and even with premaxillaries. Measurement8 of the type of Mus commis8arius Mea?·ns. mm.

Total length Head and body Tail Hind foot Head Ear above notch Ear above crown Skull: Occipitonasal length Basilar length Greatest parietal breadth Greatest zygomatic breadth Length of audital bulla! Length of internasal suture Length upper molar series

* Philip.

Journ. Sci. § D 7 (1912) 32. 46 (1913) 326.

t Hollister, Proc. U. S. Nat. Mus.

167 84 83 17 24 11 13.5 19.5 ~5.5

8.9 10.2 3

6.2 3.1

480

PHILIPPINE LAND MAMMALS

Remarks.-I have tentatively retained this species in the Philippine list although I am of the opinion that it is a synonym of PrIus castaneus 路Waterhouse. The latter species was based on a specimen ~pparently discolored by alcohol. Unfortunately, only a few measurements of the type were published. The bicolored tail of this species is scarcely noticeable, particularly if specimens have been kept in alcohol. It is highly probable that Mus musculus Linn:reus will eventually be taken (Elera has already listed the species, but his record is doubtful) in the Philippines, since it is found in most of the ports of the world together with various widespread species of Rattus. Genus BATOMYS Thomas

Batomys THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 162; Trans. Zool. Soc. London 14 (1898) 405; HOLLISTER, Philip. J ourn. Sci. 搂 D 7 (1912) 32; TROUESSART, Cat. Marnrn. (1898-99) 501.

Large, heavily furred rats, similar to Carpomys but with a shorter, though similarly hairy, tail; eyes either surrounded by a seminaked or finely haired ring (B. grantii) or normally furred (B. dentatus); ears br0ad and rounded; forefeet rather elongate; pollex with a nail; hind feet broad; six large sole pads, unstriated; whole 'o f heel hairy to level of hinder end of last sole pad; fifth hind toe reaching base of third phalanx of fourth; hallux reaching base of second toe. Skull and teeth.-Skull more elongate than in Carpomys; brain case sm~ll, face long; interorbital space narrow, its edges with only 'the slightest indication of ridges; interparietal moderately large; anterior edge of zygoma root not projected forwards; anterior palatine foramina large; bullre small. Incisors narrow, smooth in front. Molars instead of being brachyodont are more or less hypsodont, crown at least as high above bifurcation of roots as it is broad; molar laminre as in Mus .. laminre transverse, not oblique; second and third upper molars with well-defined anterointernal supplementary cusps; first and second lower molars with distinct posterior medial supplementary cusps. The type of the genus is Batomys grantii Thomas. Two species of this genus are known, both from the highlands of northern Luzon.

481

BATOMYS

Key to the species of Batomys Thomas.

a'>.

A bare area around eye; tail uniformly col()red and covered with hair, concealing scales --____________________ .________ .________ ._ B. grantiti Thomas (p. 482). /18

120

PHILIPPINE ISLANDS

122 Batan I.

I 4

126

速 Mus commissar>ius (s> Batomys gr>antii

息 Batomys dentatus @ limnomys picinus

速 limnomys sibuanus 速 limnomys mear>nsi @ Tar>somys apoensis

'-i ~

....

..... :):: ~

"'Cagayan Sulu

120

FIG. 25.

122

124'

126

Distribution of Mus, BatomY8, L-imvnIYml//B. and Tar8O'rTll/lB in the Philippines.

a '. No bare area around eye; tail thinly covered with hair, not concealing

scales; lateral half of tail white; teeth larger than in B. grantii. B. dentatus Miller (p. 483). 269468--81

482

PHILIPPINE LAND MAMMALS BATOMYS GRANTII Thomas

Batomys grantii THOMAS, Ann. & Mag. Nat. Hist. VI 16 (1895) 162; Trans. Zoo!. Soc. London 14 (1898) 405, p!. 33, fig. 2 (animal); p!. 34, figs. 6 and 8 (teeth and skull); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 32.

Description of the species.-Size of a large rat; fur thick, close and rather coarse; eyes surrounded by naked or finely haired ring; tail with a moderate growth of hairs which hides scaly annulations; ears of medium length, rather thinly haired. Color.-General color coarsely grizzled fulvous and black on back and sides, face more grayish; posterior part of back and rump tending toward rufous; back of ears black or dark brown, undersurface dirty buff, not sharply defined; base of hair slatecolored throughout, though an indistinct mesial whitish line is sometimes present; metacarpals and metatarsals brownish mesially, whitish laterally and on digits; tail thickly and uniformly covered, except on basal centimeter, with dark brown or black hairs 7 to 9 millimeters long. Skull and teeth.-Skull long, rounded, the supraorbital edges not ridged; zygomatic plate slightly concave anteriorly; bullre small; third upper molar as long as second. (After Thomas.) Measurements of the type of Batomys grantii Thomas. Male. mm.

Head and body Tail, doubtfully perfect Hind foot, moistened Skull: Basal length Greatest breadth Nasals, length Interorbital breadth Diastema Palatal foramina Upper molar series Lower jaw, condyle to incisor tip

204 121 35.5 40.5 22.2 19.5 5.5 12.8 8.5

7.8 30.2

Remarks.-The type and paratypes were collected on Mount Data by Whitehead, at an elevation of approximately 2,154 meters. Whitehead, quoted by Thomas, says:

BATOMYS

483

This interesting new Rat was captured for me by the Igorrotes, with the aid of their small terriers; it seems rare, only three specimens being obtained. Batomys grantii is at first sight the same animal as the next species, Carpomys melanurus, but has a bare ring about the eye and when alirve is easily separated from Carpomys by 't his character. The two animals, however, which externally appear almost identical, have the teeth so different that they have been separated by Mr. Thomas into different genera. Found at 7,000 feet on Monte Data. BATOMYS DENTATUS Miller

Batomys dentatus MILLER, Proc. U. S. Nat. Mus. 38 (1911) 400, pI. 20 (5 unnumbered figs.); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 32; Proc. U. S. Nat. Mus. 46 (1913) 326.

Description of the species.-Similar to the preceding species in size and general appearance, but underparts more buffy and terminal half of tail white; no bare area around eye, this area furred as in other rats; tail very thinly covered with hair so that annulations are plainly marked; twelve rings to a centimeter. Color.-Entire upper parts uniform light brown, slightly more yellow than wood brown; face tinged with drab, lumbar region and rump russet, entire area finely and inconspicuously varied by blackish tips of longer hairs and subterminal annulations on those of underfur; sides and flanks suffused with ochraceous-buff, this color covering entire underparts except chin and throat, which are pale cream-buff much darkened by slate-gray undercolor; general effect in this region dull buffy-drab; feet dull buffy-gray, the metapodials clouded with sepia; ears an indefinite dark brown; whiskers black; tail uniform blackish brown through a little more than basal half, then abruptly white to tip. Skull and teeth.-Skull agrees largely with that of Batomys grantii; in lateral view it differs in being of greater depth through brain case over front of parietals. Teeth disproportionately large; width of crowns ,s lightly exceeds that of palate at its narrowest point; length of tooth row, measured along alveolar surfaces, exceeds distance from alveolus of first upper molar to front of incisive foramen by about 1.5 millimeters; enamel pattern of teeth agrees with that of B. grantii except in certain unessentials.

484

PHILIPPINE LAND MAMMALS

Measurement8 of the type of Batomys dentatus Miller. mm.

Head and body Tail Hind foot, dry Skull: Condylobasal length Zygomatic breadth Interorbita.l constriction Mastoid breadth Greateslt depth of brain case Nasal length Diastema Mandible Maxillary tooth row, alveoH Mandibular tooth row, alveoli Width of crown of first upper molar Least width of bony palate

195 185 36 42.2 23 6 15.4 12.4 18.8 12.2 28.2 9.6 9.8 2.8 2.8

Remarks.-Only the type of this species is known. It was collected by Edgar A. Mearns, at an altitude of approximately 2,154 meters, at Haight's Place, Benguet, Luzon, July 31, 1907. Genus LIMNOMYS Mearns

Limnomys MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 450; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 31.

A marsh rat, resembling Rattus.. skull relatively broad and high, with flattened audital bull:re; supraorbital beading slight, becoming obsolete on front of parietals; teeth apparently not to be distingui.shed from those of Rattus except that incisors are very much narrower; pelage long and dense; that of under~ parts resembling that of Oryzomys.. tail thinly haired, more hairy near end where annuli are. concealed; tail longer than head and body; ears moderately coated on edge and outer surface with short hair; feet broad, their underside smooth and naked, with tubercles flattened; sixth (or posteroexternal) tubercle obsole~e. (After Mearns.) The type of the 'genus is Limnomys sibuanus Mearns. So far as is known the genus is confined to the Philippines. Three species, all from high mountains, have been described. Key to the species of Limnorruys Mearns. a

Tail longer tha.n head and body. . b'. Raw umber above, washed with tawny-olive; feet banded with seal brown .__ ... __ ... __ . ____ .. __ ........... __ .__ ........... __ ..... L. sibuanus Mearns (p. 485).

485

LIMNOMYS

b '. Grayish brown above, sides paler; hands whitish; feet blackish, edged with white .................................. L. mearnsi Hollister (p. 487). a '. Tail shorter than head and body, blackish above. L. picinu s Hollister (p. 486). LIMNOMYS SIBUANUS Mearns

Limnomys sibuanus MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 452; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 31; Proc. U. S. Nat. Mus. 46 (1913) 324.

Description of the species.-Characters those of the genus; tail uniformly colored with sixteen annuli to a centimeter at the middle, extreme base smooth and naked below; whiskers long, extending to shoulders; belly so densely furred that mammre cannot be detected; longest hairs of rump 35 millimeters long. Color.~General color of upper parts raw umber, washed with tawny-olive on sides, darkest on middle area of upper parts, where black hairs are intermixed; entire underparts, except tail, cream-buff; washed with clay color on chest and around vent; tail grayish brown; upper surface of feet broadly banded with seal brown, toes and sides of feet brownish white; nail tufts grayish white; claws flesh color tipped with horn. color; ears dark drab, with pale yellowish tufts of hair at anterior base; muzzle grayish drab; whiskers mixed with black and white; pelage of upper parts gray at base, of underparts creambuff to base. Skull and teeth.-Same as for genus. Measurements of the type of Limnomys sibuanus Mearns. mm.

Total length Tail vertebra! Tail to end of hairs Hind foot Head Ear, above crown Ear, above notch Skull: Basilar length (Hensel) Occipitonasal length Interparietal breadth Length of internasal suture Greatest breadth of rostrum Length of upper molar row

275 150 153 30 40 15 21

26 32 14.5 12 5.5 5

486

PHILIPPINE LAND MAMMALS

Remarks.-The type specimen was collected by Edgar A. Mearns at an elevation of approximately 2,030 meters on Mount Apo, Davao, June 30, 19Q4. Mearns says: The unique type was trapped on a wet, mossy growth of vegetation, on the left bank of a little stream which in its lowm- course becomes the Sibuan River, iflowing from Mount Apo. In spite of persistent efforts we failed to procure other specimens. LIMNOMYS PICINUS Hollister

Limnomys picinus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 325.

Description of the species.-Medium-sized rat; tail and ears short; coloration wholly different from the other known members of the group, blackish above, dusky below; pelage long, very soft. Color.-Above, blackish; head, forward part of body, and sides finely lined with ochraceous-buff; hairs uniformly dark neutral gray, with narrow tips of ochraceous-buff; scattered through the pelage of back are numerous overlying hairs of pure black, which greatly reduce the effect of the ochraceous tips on the ordinary hairs; center of back anrl rump almost pure blackish; cheeks lighter, more grayish buff; underparts dark grayish drab, not sharply marked from color of sides, hairs gray at base, tipped with drab; hands and feet brownish black, digits with long, yellowish white hairs; tail brownish black above, slightly lighter and more brownish below. Skull and teeth.-Skull smaller than that of Limnomys sibuanus, larger than that of L. mearnsi; in general more like that of L. mearnsi, with wide rounded brain case and indistinct beading; teeth slightly larger than in L. mearnsi, much larger than in L. sibuanus. (After Hollister.) Measurements of the type of Limnvmys picinu8 Hollister. mm.

Total length Tail vertebrm Hind foot Skull: Condylobasal length Zygomatic breadth Breadth of brain case Length of nasals Upper tooth row, crowns Mandible

205 100

28 26.7 14.9 14.4 10.1 5.4 16.2

487

TARSOMYS

Remarks.-The type, in the United States National Museum, was collected by Edgar A. Mearns at an elevation of approximately 1,380 meters on Mount H.alcon, Mindoro, November 16, 1906. The type, an adult female, is the only specimen known. LIMNOMYS MEARNSI Hollister

Litmnomys mearnsi

HOLLISTER,

Proc. U. S. Nat. Mus. 46 (1913) 324.

Description of the species.-Small; coloration darker than in Limnomys sibuanus, upper parts more slate-gray, less reddish; underparts whitish, not buffy. Color.-Upper parts an indefinite dull grayish brown, pelage long and soft, deep neutral gray, tipped with dull brown; sides paler; underparts entirely whitish, hairs unicolored; hands thinly haired, white; feet blackish, edged with white; toes white. Skull and teeth.-Skull much smaller than that of L. sibuanus, ,with relatively wider and much more rounded brain case; supraorbital beading very slight; teeth larger, relatively wider and much more :r:ounded ; first upper molar especially wider and less narrowed anteriorly. (After Hollister.) Measurem,e nts of tAe type of Limnomys mea?'"nsi Holliste?'. mm.

Total length Tail vertebrre Hind foot Skull: Condylobasal length Zygomatic breadth Breadth of brain case Interorbital constriction Length of nasals Upper tooth row, crowns Mandible Lower tooth row

22'0 133 27.5

25.3 14.4 14 4.5 9.6 5.2 15.8 5.4

Rerrw;rks.-The type and two paratypes were collected by Edgar A. Mearns near the summit of Grand Malindang Peak, Mindanao, at an elevation of approximately 2,770 meters, June 7, 1906. The type is in the United States National Museum. Genus TARSOMYS Mearns

Tarsomys MEARNS, Proc. U. S. Nat. Mus, 28 (1905) 453; TER, Philip. Journ. Sci. ยง D 7 (1912) 32.

HOLLIS-

488

PHILIPPINE LAND MAMMALS

Pelage long, rather coarse, but not spiny, with hair of two lengths, overhair about twice the length of the ordinary coat; ear prominent, its height from crown nearly one-third length of head; skin of ear roughened and coated on both surfaces with short thick hairs; whiskers long, surpassing ears; eyes of medium size; feet naked below; forefeet with five metapodials; hind feet with six, those at base of inner and outer digit two~ lobed, and with several small additional tubercles; tail hairy. Skull and teeth.-Skull and teeth resembling Batomys Thomas, but relatively broader, with a more inflated brain case, and large vacuities -above auditory meatus; lateral teeth extremely oblique in their position, upper rotated outward and lower correspondingly rotated inward; anterior upper molar resembles that of Mus, but differs in that the anterior series comprises but two cusps, a small inner and a large outer one. (After Mearns.) Only one species is known. TARSOMYS APOENSIS Mearns

Tarsomys ClJpoensis MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 453; HOLLISTER, Philip. Journ. Sci. ยง P 7 (1912) - 32; Proc. U. S. Nat. Mus. 46 (1913) 326.

Description of the species.-Pelage coarse, longest on rump; longest underhairs 20 millimeters long, stiff black overhairs 40 millimeters long; palms and soles naked, but with a hairy tuft on inner side at base of heel and a few long hairs at outer border of wrist; toes tufted at base of claw.s, hairs exceeding length of claws; forefoot 5-tuberculate; three small anterior tubercles forming a triangle, the front one placed at j unction of third and fourth digits; the other two, at junction of second and third, and fourth and fifth digits, respectively, the outer one being slightly posterior to other two; hind foot with doublelobed tubercles at base of inner and outer digits; -small rounded tubercles at j unction of second and third and third and fourth digits, a small rounded tubercle near outer margin of naked sole anteriorly and an elongated one near inner margin posteriorly; several additional minute tubercles; tail thickly coated with stiff brownish hair obscuring the scaly annuli; thirteen annuli to a centimeter on middle of tail; pollex with a small, round, flattened nail. Color.-Upper parts brownish slate, with hairs of undercoat lightened with yellow-brown tips; muzzle blackish; underparts

489

HYSTRICIDJE

grizzled yellow-brown; tail dark purplish brown throughout; hairs on top of feet slightly grizzled drab-brown; ears dark purplish brown, coated with blackish hairs; whiskers black with a few grayish white ones; pelage of upper parts of body slate at base; on underside paler. Skull and teeth.-Brain case swollen; rostrum elongated, similar to that of Apomys, but audital bullre and anterior palatal foramen very different; rostrum elongate; nasals narrow, truncate; skull light, with a large semicircular vacuity between temporal and parietal bones above auditory opening; bullre similar to those of Rattus, but more flattened and compressed externally and the palate broader. (After Mearns.) Measurements of the type of Tarsomys apoensis Mearns. nun.

Head and body Tail Forefoot Hind foot Head Ear from crown Skull: Basilar length OccipitonasaI length Greatest inlterparietal breadth Least interorbital bFeadth Length of internasal suture Breadth of palate between last molars Length of upper molar row Mandible Length of mandibular molar row

135 120

16 415 14 32 28.5

37.5 16 6

15.2

5.1 6.2 21

6.2

Rernmrks.-The type, No. 125280, in the United States National Museum, was collected by Edgar A. Mearns at an elevation of approximately 2,075 meters, on Mount Apo, Davao, Mindanao, July 5, 1904. The National Museum also has two specimens from Mount Malindang and two from Mount Bliss. No other specimens are known. Family HYSTRICIDlE Stoutly built animals; limbs of nearly equal length; tail elongate and prehensile or short and nearly absent; facial portion of .skull short and broad, jugal witnout an inferior angle; molars with external and internal enamel folds; molars completely or partly rooted; body skin with numerous spines.

490

PHILIPPINE LAND MAMMALS

Three subfamilies, the Synetherinre, the Atherurinre, and the Hystricinre, are recognized; the last is represented in the Philippines by one genus and species. Subfamily HYSTRICIN..-E

Tail :short, without a hairless scaly portion between base and apex; terminal hairs of tail modified into hollow capsulelike structures, mostly open at ends; well-developed quills on back and sides; four sacral vertebrre; molars rootless, hypsodont. There are three genera in this subfamily; namely, Hystrix, Acanthion, and Thecurus. The last is represented in the Philippines by one species, which is confined to the Palawan and Calamian groups of islands and, perhaps, Cagayan Sulu. * Genus THECURUS Lyon

Thecurus LYON, Proc. U. S. Nat. Mus. 32 (1907) 582; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 34; Proc. U. S. Nat. Mus. 46 (1913) 338.

Similar to Acanthion, but capsulelike ends of caudal hairs smaller and shorter, often closed at ends; quills ,smaller and replaced on lower rUrbp by grooved spines similar to those on upper back; no bristly hairs on head or neck but merely soft flattened spines; large heavy quills occupy about the third fourth of back; less numerous and shorter than in Acanthion, the largest not exceeding 150 millimeters; a few long stiff bristles scattered among quills; some short quills on base of tail, terminal portion with capsulelike hairs; sides of head, underparts, and legs generally clothed with rather soft, flattened, grooved spines; cervical vertebrre, 7; dorsal, 14; lumbar, 5; sacral, 4; caudal, 17. . 1 0 1 3 Dental formula: 1. c. 0' pm. I' m. 3 ==20.

THECURUS PUMILUS

(Gunther)

HY8triX pumila GUNTHER, Ann. & Mag. Nat. Hist. V 4 (1879) 106 (Paragua); EVERETT, Proc. Zool. Soc. London (1893) 495; STEERE, List Birds and Mamm. Steere Exp. Philipp,i nes (1890) 30; BOURNS and WORCESTER, Occ. Papers Minn. Acad. Nat. Sci. 1 (1894) 1-64; SANCHEZ, Anal. Soc. Espanola de Hist. Nat. 2 29 (1900) 177-290

* Capt. Francis Link, Philippine Constabulary, has prepared a list of native names of Ithe animals occurring in the Sulu Archipelago and Cagayan Sulu. His note follows: "Batak-Porcupines. None on this island, (Jolo), but they live on Cagayan Sulu, where they are not popular on account of the damage they do tA:> coconut trees."

THECURUS

491

(Palawan and Balabac); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 34; Proc. U. S. Nat. Mus. 46 (1913) 338. Acanthion pumila ELERA, Contr. Fauna Filipina (1915) 151.

Description of the species.-(From No. 18, E. H. Taylor collection; collected at Iwahig, Palawan, November, 1923, by E. H. Taylor.) Body densely covered with flattened spines, each deeply grooved and somewhat flexible; on posterior half of body especially on rump they increase to heavy, nearly rigid quills, all except the largest, with a strong groove; these grow smaller and shorter towards tip of tail; at tip of tail a few bristles which terminate in small capsulelike cups either open or closed and terminating in a long bristle tip; a few elongate stiffened vibrissre emerge from spines and reach a length nearly double that of spines in the same locality; an occasional bristle on back and side; very fine small hairs appear between spines; vibrissre on snout greatly elongated; a prominent group below anterior base of ear; nose covered with fine short hair; belly with grooved spines, but more flexible than on back; feet clothed with coarse bristlelike hairs. Color.-Spines on anterior part of back dark drab-brown, hidden basal portion dirty whitish and extreme tip likewise colored; quills on posterior part of back darker, nearly black; the white tips more prominent; on sides the amount of white on tips varies and a flecked appearance results; lighter brown (flecked with buff on sides); belly with much buff-white, especially on breast; on head spines more brownish as are spines and hairs on limbs and feet; outer surface of incisors orange. SkuU arod teeth.-Width of interorbital constriction but little less than brain case; nasals short; a distinct crest on posterior portion of skull; zygomatic arch much thickened where process arises; anteorbital foramen large, somewhat rounded; bullre large; palate ending at posterior end of second molar in a Vshaped notch; palatal foramina very small. Teeth somewhat cylindrical, with internal enamel folds which in worn teeth appear as small cavities on nearly level surface of tooth; upper premolar of greater diameter than other cheek teeth; vertical groove, caused by infolding of enamel, visible on inner side of premolar (probably present in other teeth not so greatly worn) ; in lower jaw the teeth show these folds on premolar and last molar. Posterior edge of mandible slopes downward and forward from condyle.

492

PHILIPPINE LAND MAMMALS

Measurements of Thecurus pumilus (Gunther). mm.

Total length, estimated Tail, estimated Spines on neck Quills on back, longest Vibrissre on snout V ibrissre on posterior body Length of hind foot, dry Height of ear, dry Skull: Greatest. length, estimated Nasal bone Interorbital constriction Zygomatic width Width of brain case Length of upper molar series Diastema Lower molar series Diastema Mandible, bone only

480 39 36 65

120 130 55

98 24

52 36

20 24

18.8

17 58

Remarks.-The skull measurements agree almost exactly with the type. The specimen described is from a place near the type locality, which is Puerto Princesa, Palawan. GUnther gives the length of the longest quills as about 200 millimeters, which is much longer than any now present on my specimen. I have examined two specimens belonging to the Bureau of Science (No. 138) which show very remarkable variation and may represent a different species. The specimens are from Busuanga Island. Both were captured alive and were kept alive at the Bureau of Science, Manila. One of the specimens now consists of a dry skin and skull, the second the skull only. The following differences obtain: Color above distinctly lighter brown with indefinite darker line along middle of back from neck; quills blackish, in marked distinction from surrounding spines; white tips of spines indistinct or wanting; belly generally drabbrown, rather buffy on neck and breast; hair and spines on feet blackish instead of reddish brown; longest spines somewhat longer and ears seem to be distinctly larger than in the Palawan specimen. Skull shorter, teeth somewhat smaller, and posterior edge of the mandible sloping downward and slightly backward from condyle; the ascending ramus of the mandible less than half the bulk of that on the Palawan specimen.

493

ARTIODACTYLA

Measurements of Thecurus pumilus from Busuanga. Skull: Greatest length Zygomatic width Interorbital width Length of nasals Length of mandible Upper tooth row Lower tooth row

nun.

Mm.

93 53 31 23 56 17.5 19

89 4'9 31 22 55 18 19

The type, in the British Museum, was collected at Puerto Princesa, Palawan, by Everett. Order ARTIODACTYLA As originally planned this work was to include only the .smaller Philippine mammals, and it is with considerable hesitancy that I have included the larger forms belonging to the Ungulata. However, I feel that, while I am unable to straighten out satisfactorily the tangled messes which obtain in the families Cervidre and Suidre and perhaps to a lesser extent in the Bovidre, it will be of value to record the present state of our knowledge of these families. The chief confusion in the Cervidre and in the Suidre is the result of the work of the Reverend Pere Heude on the deer and the pigs of the Philippines. He says: II sembI a que depuis Cette revision de Sir V. Brooke o~ ait considere l'Hude des cerfs comme terminke. II est pourtant loin d'en etre ainsi. On'a pris au moins les type generiques pour des especes; on n'a pas compare piece a piece, soit faute de materiaux suffisants Oili convenables, soit pour toute autre raison. L'etude du type RQussa, tel qu'il se montre aux Philippines, fait voir clairement que si Ie nombre des formes specifiques n'est pas aussi considerable que je Ie pense, du moins faut il en admettre une quantite bien superieure a celle qu'on a admise jusqua ce jour.

With this premise the good father has given specific names to no less than twenty-nine deer species within a radius of 100 miles of the City of Manila and to a total of forty-one species in the Philippines. In a footnote on page one of Heude's papers, * appended to the title, one notes the following: Les personnes qui ont les onze premieres pIanches de se cahier sont de se rappeler que Ie titre n'en etait que provisoire ainsi que les

pril~es

* Etude sur Les Ruminants de l'Asie oriental Ceris des Philippines et de I'Indo Chine. Mem. Rist. Nat. Emp. Chinois Tome II Premier Cahier avec 21 planches. Chang-hai (1888).

494

PHILIPPINE LAND MAMMALS

Iegendes. Des materiaux plus abondants et des etudes plus approfondies ont engage I'auteur du Memoire a introduire un titre different et une nouvelle explication des planches. Ce que a precede doit donc etre considere comme suprime.

In a later revision of the Philippine deer of the northern islands he reduces the number somewhat and' corrects the explanations of the plates. Some of the forms described are certainly valid species, while others are merely synonyms. It is doubtful if it will be possible to determine positively the status of the names given, for even in the revised explanation of the plates there are certain obvious errors, such as the same drawing serving for two, presumably distinct, species. Owing to the fact that I had not originally intended to include this group, my own collection contains only a few specimens of these larger forms; and there is but a very meager representation in the collection of the United States National Museum. The revision of this group must necessarily await large collections from all parts of the Philippines, particularly from the type localities of the species described. It is obvious that there must be considerable variation in the deer species as well as in those of the pigs, probably due to crossing of species. The Artiodactyla are represented in the Philippines by the families Suidre, Tragulidre, Cervidre, and Bovidre. Key to PhilJippine families of Artiodactyla. ((. '. Without horns. b', Larger; snout elongated, with upper incisor teeth.

(Pigs.) SUIDJE (p. 494). b". Smaller; deer like ruminants without horns but with upper canines tusklike. (Chevrotain.) ...................................... TRAGULIDJE (p. 505). a'. With horns. b '. Solid horns, only in males; size variable; upper canines present or absent. (Typical deer.) ............................................ CERVIDJE (p. 507). b". Horns in both males and females. (Buffalo.) ........ BOVIDJE (p. 526).

Family SUIDlE This family is represented in the Philippines by the genus Sus. The numerous genera and subgenera that have been proposed appear to be unwarranted as the relationship among most of the forms is specific rather than generic. The species all belong to the single subfamily Suinre. Genus SUS Linnreus

Sus LINNJEUS, Systema Naturre 10th ed. 1 (1758); 12th ed. 1 (1776) 102-104; BRISSON, Regnum Animale in Classes IX distrib. 2d ed.

sus

495

12 (1762) 73-78; HAY, Bull. U. S. Geol. Surv. 179 (1902) 661; TROUESSART, Cat. Mamm., Suppl. (1904-05) 660. EU8US (subgenus) GRAY, Proc. Zool. Soc. London (1868) 32 (type Sus barbatus Mtiller, Borneo). Euhys (subgenus) GRAY, Cat. Carniv., Pachyderm., and Edentate Mamm. Brit. Mus. (1869) 339 (later made genus, type Sus barbatus Muller). Nesosus (subgenus) HEUDE, Mem. l' Hist. Nat. Emp. Chinois 2 pt. 2 (1892) 85, 92, 106 (includes S. calamianensis, S. arietinus, S. minutus, and S. cebifrons from the. Philippines with several exJtraPhilippine species). Verrusus HEYDE, Mem. l'Hist. Nat. Emp. Chinois 2 4 (1894) 213222 (includes Sus inconstans, S. megalodonflus, S. effrenus, and S. arietinus) . Rhinosus (subgenus) HEUDE, Mem. l'Hist. Nat. Emp. Chinois 2 4 (1894) 213, 222 (includes Sus longirostris Nehring and Sus calamianensis Heude, (type).

Most, if not all, writers regard all of the Oriental and East Indian wild boar under th~ genus Sus with but little regard for 8ubgeneric designations. The group may be defined as typical wild hogs with the following dental formula:

. B

1 4 3 44 c. I' pm. 4' m. 3 = . Upper incisors diminishing rapidly in size from first to third; canines strongly developed with persistent pulp, partly covered with enamel, those of upper jaw curving outward, upwards and inwards; upper molar-premolar teeth in a continuous series, growing more complex from first to last, usually contiguous; the same is true of lower molar-premolar series except that first premolar is usually separated from the rest of the series; lower canines grow upward, outward, and backward, usually describing an arc of a circle. The milk dentition consists of 28 teeth. 路31 3 F ormu1a: 1. 3' c. I' pm. 3 = 28\ . 1.

Skull with axis of face bent down on basicranial axis; broad flat frontals with small supraorbital processes which do not join zygomata, leaving orbits. open behind; nasals long and narrow; premaxillre send up long nasal processes, stopping short of frontals; a prenasal bone developed at anterior extremity of mesothymoid, serving to strengthen snout for rooting; palate long and narrow and extends behind last molar tooth. The number of species occurring in the Ph-i lippines is still a matter of doubt and much work remains to be done in this

496

PHILIPPINE LAND MAMMALS

group. I shall include here species which appear to be worthy of recognition based on published works and the meager material I have been able to examine. Heude, * in his several papers, lists Sus calatmianensis, S. cebifrons, S. minutus, S. effrenus, S. arietinus, S. microtis, and S. frenatus, with descriptions and with references to plates and figures. Figures are given of ,several other forms which are presumably Philippine; namely, Sus megailodontus, S. 'fIULinitensis, S. conchyvorous, and S. jalaensis; two names given as Phi1ippine~ Sus militaris and S. basilanensis, are,. so far as I can find, nomena nuda. ~us inconstans bears a very meager description. Hollister t recognizes as valid species Sus calamianensis, S. cebifrons, S. minutus, and S. inconstans, while S. e/fre'YIJUs, S. micro tis, S. frenatus, and S. a,rietinus are placed as synonyms of Sus philippinensis Nehring. Besides the four supposedly valid species of Heude, Hollister recognizes Sus ahamobarbus Huet, S. barbatus balab'acensis Major, S. barbatus palavensis Nehring; S. mindanensis Major, and S. philippensis Nehring. The three specimens of Philippine wild boar in the United States National Museum are identified as Sus philippinus (Luzon), S. inconsta;ns (Mindanao), and S. calamianensis (Culion) . sus P'HI1'..IPPENSIS

Nehring

Sus ce~ebensis philippens1Js NEHRING, Sitz-Ber. Ges. Nat. Freunde Berlin No.5 (1886) 83, May, (Luzon); (1888) 10 fig.; Abhandl. undo Ber. Konigl. ZooI. Mus. Dresden (1888-89) 14-17, 24'-26 pI. 1, figs. 3 and 4, pI. 2 fig. 4; Zool. Ann. (1891) 457 (Mindoro). Sus mar chei RUET, Le Naturaliste 2 No. 20 (1888) 6 (Laguna Prov). ? Sus arietinus REUDE, Mem. Rist. Nat. Emp. Chinois 2 (1892) 114 pI. 20, fig. 1, pI. 20 B, fig. 5, pI. 27, fig. 8, pI. 29, figs. 1, 2 and 3, pI. .29 C, fig. 4 (Manila ?); (1894) 219. ? Sus effrenus REUDE, Mem. Rist. Nat. Emp. Chinois 2 (1888) (unnumbered page following pI. 19) pI. 17 fig. 7; 2 (1892) 114, pI. 20 B, fig. 2, pI. 27, figs. 4, 5, 6, pI. 29, figs. 3, 7; pI. 29 C fig. 3 (J alajala, Laguna de Bay); 2 (1894) 215.:/: Sus micro tis R.IDUDE, Mem. Rist. Nat. Emp. Chinois 2 (1888) (unnumbered page following pI. 19) pI. 17 fig. 10; 2 (1892) 114, 115, pl.

* Mem. l'Rist. Nat. Emp. Chinois 2 1 (1888) 52-64, pI. 17; 2 2 (1892) 85-111, 115, pIs. 20, 20B, 27, 28, 29, 29C; 2 4 (1894) 212-222 pI. 40, figs. 1, 2, and 5. t Philip. Journ. Sci. ยง D 7 (1912) 38. :/: Rere the following plates are said to refer to this species: pI. 17, fig. 11; pI. 27, figs. 1, 2, 8. 8; pI. 19 C, fig. 3.

sus

497

20 B fig. 6, pI. 28, figs. 5, 6, 7 (Jala-jala, Laguna de Bay); 2 (1894) 217. ? * Sus frenatus Heude, Mem. Hist. Nat. Emp. Chinois (unnumbered page following pi. 19) pl. 17, fig. 11; 2 (1892) 114, 115, pI. 27 figs. 1, 2, 3, 7 (Jala-jala, Laguna de Bay); (1894') 216 pI. 20 B, fig. 2. ? Sus joloensis HEUDE, Mem. Hist. Nat. Emp. Chinois 2 (1888) (unnumbered page following pI. 19) pi. 17, fig. 9. ? Sus conchyvorus HEUDE, Mem. Rist. Nat. Emp. Chinois 2 (1888) (unnumbered page following pI. 19) pI. 17, fig. 8. Sus celebensis philippinensis THOMAS, Trans. Zooi. Soc. London 14 (1898) 411 -(Cape Engafio, Luzon); BOURNS and WORCESTER, Minn. Acad. Nat. Sci.; Occ. Papers 1 (1894) 63 (v~ry numerous ,l ocalities; probably part of these belong to other species. Lists Tawitawi, Sulu, Basilan, Mindanao, Samar, Negros, Panay, Mindoro, Sibuyan, Luzon, and Masbate).

Description of the species.-Closely related to Sus celebensis but skull lower and more drawn out, rostrum narrower and proportionally longer than appears to be the case in S. celebensis; bony area above upper tusk lower and not so thickened posteriorly; many other small differences occur which are difficult to expre~s. The variations in the teeth from ,those of S. celebensis are relatively small and are ,scarcely greater than obtain between various specimens of S. philippensis. Measurements of Sus philippensis Nehring. mm_

Basal length of skull Profile length of skull, median Greatest zygomatic widlth Greatest width of occipital wings Least width between occipital beading (Scheitelleisten) Greatest length of lower jaw

259 315 136 86 30 230

Remarks.-N ehring identified all of the Philippine pig material, examined by him, under this species except forms occurring in Palawan. It is not improbable that he had more than a single speci-es in his material. Hollister does not comment on the Bourns and Worcester records which report specimens from numerous southern localities. They report the form from Mindanao, Negros, Panay, Samar, Sibuyan, Sulu, and Tawitawi. Steere lists specimens from Basilan and Mindoro. I believe that many of these records should be questioned, at least until *Reude, page 216, gives the following plate references to this form: pi. 27, figs. 5, 6, 4; pI. 29, fig. 7; pI. 17, fig. 9; pI. 20 B, fig. 2, which elsewhere are supposed to refer to effrenu8. 259453-32

498

PHILIPPINE LAND MAMMALS

further evidence is brought to prove the presence of the species in the southern islands. SUS CALAMIANENSIS

Heude

Sus calamianensis HEUDE, Mem. l'Hist. Nat. Emp. Chinois 2 (1888) 53 (and unnumbered page following p. 64) pI. 17, fig. 4, (teeth); 2 (1892) 114, pI. 20B, fig. 3; 2 (1898) 221, pI. 40, figs. 1, 2, and 5. Elera, Cont. Fauna Filipina (1915) 258 (name, no data). Sus ba?'batus var. calamianensis NEHRING, Sitz.-Ber. Ges. Nat. Freunde Berlin 8 (1894) 191 (Culion) and ZooI. Gart. (1895) 46; TROUESSART, Cat. Mamm. Suppl. (1904-05) 662; FORSYTHE-MAJOR, Ann. & Mag. Nat. Hist. 19 (1897) 535. Sus calamianensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 38; Proc. U. S. Nat. Mus. 46 (1913) 331 (Culion).

Description of the species.-This form is probably specifically distinct from the wild boar found on Palawan and Balabac, which lie between the Calamianes and Borneo. It is presumed that only a single species exists in the Calamianes. The literature gives rather meager details for descriptions. The following notes are from Heude's drawings and published notes. The upper canines are very divergent; the internal radius of the arc of the circle described by the lower tusks is 56 millimeters; they have a length above the alveoli of 65 millimeters; the lachrymal bone is small, rectangular below; the depression is much sunken under the much overhanging cornice of the frontal; the orbital cornice is lacking, but the orbital edge is thickened; the two molar protuberances are sharp, elongated, and remarkably close to the flattening of the jugal; the forehead is broadly concave, while the frontal region is slightly convex; the typical peculiarity of this species and perhaps of the entire group is the posterior elongation of the palatines and the great develop~ent of the ptergoid wings. The teeth are rather weak for so large an animal; the fourth upper premolar shows the suppression of the internal rugosities of the cones; the internal face of the lower tusk is strongly arched for three-fourths of its extent; the anterior fourth has a marginal groove; the groove is larger upon the internal face; the posterior face of the lower tusk is entirely channelled. The length of the skull (four specimens) is 360 to 390 millimeters. Remarks.-Heude states that the four male skulls on which he founded the species were a gift from C. Sandoval who obtained

sus

499

the animals in the Calamianes. The exact locality is no longer known. The species has been reported from Culion-a single specimen collected there by Lanson-Scribner is in the United States National Museum. Whether or not this is a valid species or subspecies has not been determined. Hollister accepted it, at least tentatively, and identified a museum specimen as belonging to this species. However, he had no specimens from Palawan or Balabac for comparison. SUS MINUTUS Head.

Sus minutus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 (1888) (unnumbered page preceding p. 65) pI. 15, fig. 6; 2 (1892) 114, 129, pI. 20 B, fig. 1 (molar teeth); pI. 29, figs. 4, 5, 6 (cranium); 2 (1894) 218, pI. 17, figs. 6, 7, 8 (teeth) (here considered under a subgenus Nesosus.) Nesosus minutus HEUDE, Mem. Hist. Nat. Emp. Chinois 4 (1899) 116, 127 (footnote). Sus minutus TROUESSART, Cat. Mamm.; Suppl. (1904-05) 662 (Mindanao ?); HOLLISTER, Pl1ilip. Journ. Sci. ยง D 7 (1912) 39; Proc. U. S. Nat. Mus. 46 (1913) 339.

The first reference to this form is merely a name on a page of plate explanations pointing to a figure of two back molar teeth; later it appears on the same page referring to plate 19 where three views of a skull are given; in another paper (1894) a short description without measurements is given. Description of the species.-(After Heude.) This is the smallest form known. Although small, the premolars are larger and stronger than those of certain larger species; lower canines differ from those of other Philippine species; on posterior face a simple groove; lateral face with posterior half strongly swollen; anterior half with a depression bordered by a light raised beading which limits anterior depression and also depression of internal surface; superalveolar height of canine 40 millimeters; internal radius 31; upper tooth row continuous and includes canine; interval between first two premolars is very slight; skull has a remarkable swelling of orbital portion of jugal; malar protuberance rather slight, permitting large postmaxillary tubercle to be clearly seen; forehead slightly convex anteriorly; lachrymal bone nearly square; expansions above and behind upper canines on maxillary elongate. Remarks.-The small size of this form seems to warrant its being regarded a very distinct species. The type locality, how-

500

PHILIPPINE LAND MAMMALS

ever, seems to be open to question. Heude says: * "par example, celles du S. minutus et a autres Sus de l'ile de Mindanao." In the description t no locality is given. However, later t he states: "Le S. minutus qu'on truove de la Laguna it (a) Tarlac, on il abonde est la plus petite espece peut-etre du groupe luconnien." Consequently, until further correction is made we must consider this a Luzon species. SUS CEBIFRONS Heude

Sus cebifrons.

HEUDE, Mem. Rist. Nat. Emp. Chinois 2 (1888) (unnumbered page preceding p. 65), pI. 17, fig. 5 (teeth); (1892) 106, 114, pI. 28, figs. 1-4 (skull); (1894) 218 (description); FORSYTHEMAJOR, Ann. & Mag. Nat. Rist. VI 19 (1897) 527 (hybrid?); TROUESSART, Cat. Mamm. 2 (1898-99) 826; RO!LLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 38; Proc. U. S. Nat. Mus. 46 (1913) 338.

Description of the species.-The teeth of this species are probably smaller than those of any other Philippine form; lower canine 45 millimeters, internal radius 42; posterior face wide, and divided into three equal bands by two threadlike raised areas; internal face has likewise three depressed areas of unequal width; exterior face is plane except for slight striations; it is bordered posteriorly by a ~ather strong striation; profile of skull more .strongly concave and forehead more proclivous than in any other species; forehead rounded; lacrimal bone small, fossa extending backward under the projection of the frontal; maxillary projections short and elevated, not extending farther forward than posterior edge of upper canine. This species (if the condition of the skull is normal) appears to be di.stinct. Heude says: Nous avons ice un crane etrange; mais tout bossue, excave qu'il 8Oit, je crois qu'il possede la des n()tes naturelles. La raison en est qu'un second crane d'une laie que n'a pas encore toutes ses dents inddque deja dans son ossature et 路s es con/t()urs ce qu'est devenu Ie crane du vieux male.

Forsythe-Major states: The skull of "S. cebifrons" from Cebu conveys the impression of being that of a more or less domesticated animal, possibly a cross with a domestic pig-a not infrequent occurrence in those islands according to Pere Reude's statement.

The type locality is Cebu.

* Op. top.

cit. II 3 (1894) 129. cit. 2 (1894) 218. tOp. cit. 4 (1898) 127, footnote.

sus

501

SUS INCONSTANS Heude

Sus inconstans HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 2 (1892) 67, pI. 29C (canine); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 39; Proc. U. S. Nat. Mus. 46 (1913) 331.

Description of the species.-I find in Heude's work only the following descriptiorul of this form. Speaking of the character of the molar he says: "Chez Ie Sus inconstans, de l'ile de Mindanao, on ne voit plus qu'une trace du parietal externe." On plate 29C there are three figures representing the three faces of the lower canine. Hollister identifies a specimen from Catagan, Mindanao, in the United States National Museum, as belonging to this species. SUS MINDANENSIS Major

Sus verrucosus mindanensis MAJOR, Ann. & Mag. Nat. Hist. VI 19 (1897) 524. Sus mindanensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 39.

Description of the species.-HolIister fails to recognize this species in his second list. Whether it is merely an omis.sion or whether he considers the form identical with Heude's Sus inconstans, a form so meagerly described as to be practically a nomen nudum, I cannot say. It is significant, however, that he identifies a specimen collected by E. A. Mearns at Catagan, Mindanao, as Sus inconstans Heude. Rostral portion of skull long; nasals narrow in the middle of their length, but whole of nasal region still remains remarkably broad on account of overlapping of intermaxil1re on upper surface of nasal region. Measurements of Sus mindanensis Major. mm.

Basal length of skull Length of occipital crest to tip of intermaxillary Length from foramen magnum to posterior tip of vomer Length from foramen magnum to back of median of palate Length from back of palate t.o anterior end of intermaxillary Length of molar region of palate Length of incisor region of palate Length of intermaxillary at alveolar margin Width of zygomatic arches Frontal breadth between postorbital processes Narrowest frontal breadth, beitween upper lachrymal sutures

269 319 43 75 194 124.5 72 71 130 88 59

502

PHILIPPINE LAND MAMMALS

Measurements of Su.s mindanensis Major-Continued. mm.

Greatest breadth of occipital wings Width of nasals between posterior ends of intermaxillary Palatal breadth between anterior part of third molars Palatal breadth between anterior part of first molars Breadth of snout above third premolar Height of occiput from lower margin of fo'r amen magnum Height of skull resting on mandible Nasal length on middle line Length of palatine bone behind third molar (middle line) Length from nasals to a line drawn between postorbital processes Length to middle of occipital crest from line drawn between postorbital processes Length of upper molar-premolar series Length of upper third molar Length of horiz&ntal mandibular ramus from posterior end of third molar Breadth bffi;ween eondyles Length of lower third m01lar

71 25.5 23 26 46 107.5 170 164 4.5 65.5 86 93.5 24 165 110 26.5

Remarks.-This type (British Museum No. 91. 11. 28.3) was collected at Ayala, Mindanao. SUS AHOENOBARBUS Huet

Sus ahoenobarbus HUET, Le Naturaliste XX 2 (1888) 5, (January) text fig. 1 (drawing of animal) and 2 skull, mandible, and molars (probably an error, see Trouessart loco cit.); EVERETT, Proc. Zool. Soc. London (1893) 4'9 6; BOUR.NS and WORCESTER, Minn. Acad. Nat. Sci. Occ. Papers 1 (1894) 63 (Palawan); NEHRING, Sitz.Ber. Goo. Nat. Freunde Berlin No.8 (1894) 190 (Oct. 16) (Believes this form synonymous with Sus barbatus palavensis) ; HEuDE, Mem. Rist. Nat. Emp. Chinois II 4 (1892) 221 (compared with calamianensis); ELLIOT, Field Col. Mus. Zool. Series 1 (1896) 67; 8 (1907) 32: HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 38; Proc. U. S. Nat. Mus. 46 (1913) 338. Sus ba~'batus palavensis NEHRING, Abhandl. und Berichte des Konig!. Zool. Mus. Dresden (1888-89) 22, 32 (Puerto Prinoesa, Palawan); Sitz.-Ber. Ges. Nat. Freunde Berlin No. 10 (1889) 196, December 17. Sus barbatus ahoenobarbus TR.OUESSART, Cat. Mamm. Suppl. (190405) 662.

Description of the species.-Huet compares a male specimen with a female specimen of Sus barbatus and notes a remarkable difference in the condition of the lateral wings of the maxillary.

sus

503

Nose, upper lips, and chin clothed with short, black hair; below and in front of eye a band of very large, harsh hairs, blackish at base, then ocher to tip; this band becomes narrow on sides Jf nose, forming a line which passes under eyes where it widens litnd spreads out over jaw up to posterior angle of mandible; in front of yellow band and a little behind snout there are two tufts of harsh bristles curly or frizzled behind; throat, region about eyes, forehead, top of head, neck, body, and limbs covered with black hairs; beginning at top of head a sort of mane formed of longer hairs; for the greater part of their length the hairs black, but on median line of neck some among them with their tips yellowish; beginning between shoulders and continuing on back the mane is black to tips of hairs; tail rather large, with elongate, black hair, especially on median line, and becoming more elongated towaI:d extremity, where hair forms a veritable flattened brush; ears small, terminating in a point, above covered with black hair, below almost nude. Colo1".-Black, except for an ocher lateral band on .side of head, and yellowish tips on median hairs on anterior part of mane. Skull and teeth.-Skull elongated, but presumably less so than in S. barbatus; frontal region broad, very slightly arched, parietal region greatly narrowed, leaving little more than a widened crest. Last upper molar narrowed behind, widened in front, its crown, formed by numerous folds, shows in irregular spaces remains of enamel; posterior inferior molar almost as wide anteriorly as posteriorly, with three transverse cone ridges separated by grooves; posterior heel not strongly developed. Measurements of Sus ahoenobarbus Huet. mm.

Head and body Tail vel'tebrre Tip of snout to anterior part of eye Tip of snout to base of ear Length of hind foot Tot.al length of skull Length from tip of nasal to edge of orbit Width between orbits Width of zygomatic arches Width of nasals Length of upper molars

1,420 120 200 300 190 360 230 70

160 40 60

Remarks.-The status of this species is apparently not fully settled. Heude regarded it apparently as a synonym of S. cala-

504

PHILIPPINE LAND MAMMALS

mianenS1:S while Nehring considers it a synonym of S. barbatu8 palavensis. Hollister recognizes all three of these forms. The type is from Palawan ("Palauan") and was collected by M. Marche. Elliot gives the following description of a female specimen; which, together with a male, was collected in Palawan, presumably by Bourns and Worcester. Nose, 'flesh color; a br.oad band .of black hairs crosses the nose from margin of upper lip, remainder of nose and sides of head tawny ochracoous. Space between eyes, t.oP .of head, and a ,t ine down back of head, black. Hairs 'on back of neck very long, rufous with black bases. Whiskers quite long, yellowish white, with black hairs intermingled. Skin .of body tawny ochraceous (which gives :the general color to the animal) and all of it covered sparsely with black hairs, longest and thickest in the middle .of the back. The hairs between the shoulders are rufous fQr a third of their apical length. Tail covered along the center on top with short black hairs and the end tufted with the same. Legs covered with short black hairs. Ears not tufted. Hoofs narrow, black, pointed. The head is very long and narrow, with the nasa'l portion greatly elQngated.

sus

BARBATUS BALABACENSIS Major

Sus barbatus balabacensis MAJOR, Ann. & Mag. 'Nat. Hist. VI 19 (1897) 534; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 38; Proc. U. S. Nat. Mus. 46 (1913) 338.

DescriJption of the species.-The type description of this form consists chiefly of a detailed series of measurements from a series of skulls from Balabac. It is said to be related to Sus barbat'Lts, S. ahoenobarbus Huet, 'a nd S. calamianensis Heude. Measurements of Sus barbatus balabacensis Major. British Museum, No. 94688.

mm.

313 Basal length of cranium Profile lengith, occipital crest toO anteri.or end of 360 irutermaxillary Length from foramen magnum to poOsterior 43.5 end of vomer Length from foramen magnum toO back oOf 68 palate, middle line Length frQm back of palabe to anterior end .of 245 intermaxillary 177.5 Length of molar region of palate 69 Length of incisor region of palate 69 Length of intermaxilla at alveolar margin 125 Zygomaltic width 90.5 Frontal breadth between postorbital processes

British Museum. No. 94687.

mm.

307 353 42 68.5 238.5 170.5 68.5 71 134.5 93

505

TRAGULIDJE

Measurements of Sus barbatus balabacensis Major-Continued.

Frontal breadth, narrowest, between lachrymal sutures Greatest breadth of occipital wings Width of nasals betwoon posterior end of intermaxillary Palatal breadlth between anterior part of third molar Palatal breadth between anteriQr part of first molar falatal breadth between anterior part of third premolar ' Breadth of snout directly above third premolar Height of occiput from lower margin of foramen magnum Heighit of skull resting on mandibula Length of nasals Length of nasals to a line drawn between postorbital processes From occipital creSt to line between postorbital processes Upper molar-premolar series Length of upper third molar Length of horizontal mandibular ramus from posterior end of third lower molar Breadth between mandibular condyles Length of third ,lower molar

British Museum.

No. 94688.

No. 94687.

mm.

61.5 60

64.2 66,

24.5

29.5

30.5

35.5

48.5

104 180 166

103.5 177

95 92 113 29

91 112.5 30

190 104.5 33

185 109.5 37.5

Remarks.-The types of this form are from Balabac Island, and were collected by A. H. Everett. The measurements here given are of a young adult male and a very old female. Probably the mal_es reach a la.rger size than that given in the measuremeLts for the skull of a young adult male. Family TRAGULIDJE TragulidlB A.

MILNE EDWARDS,

Ann. des Sci.' Nat. Series V Zool. et

Pal. 2 (1864) 157.

This family comprises a group of diminutive deerlike animals, lacking horns, and having rather enlarged upper canine teeth. Its species are confined to the Oriental and West African Regions. The group, in spite of general appearances, is not a close relative of the Cervidre. It differs in the character of the axis vertebra which bears a simple conical peg; the fibula is complete; it has four complete digits, the ,s tomach has three instead of four compartments.

506

PHILIPPINE LAND MAMMALS

Two genera are commonly recognized. Tra,gulus is represented by many species ranging from India through southeastern Asia to the western Malaysian Islands. This genus has a representative in the Palawan group. Dorca,therium occurs on the west African coast. Genus TRAGULUS Brisson

Tragulus BRISSON, Regnum Anim. 2 ed. (1762) 65; PALLAS, Spicil. Zool. 13 (1779) 27.

Small deerlike forms without horns in either sex; canines present in both jaws, especially well developed in upper; feet with four fully developed toes; middle metacarpals fused; molars selenodont, premolars more adapted for cutting. 1 3-,m.-3 =34 Dental formula...1. 0-,c.-,pm. 3 1 3 3 . The tympanic bullre are usually filled with loose bony tissue. In the characters of four well-developed toes the family shows a greater relationship with the pigs than with the deer. The placenta is diffuse, differing thus markedly from the tufted cotyledons of the typical ruminants. Tra,gulus nigrica,n8' Thomas is the only Philippine species. Some authors have confused this with Tra,gw.lus ja,va.nicus. Elera lists T. ja,va.nicus, T. fulviventer, T. pygma,eus, and T. nigrica,n8 from Balabac. It is improbable that more than one species exists in Balabac. TRAGULUS NIGRICANS

Thomas

Tra[J'Ulus nigrricans THOMAS, Ann. & Mag. Nat. Hist. VI 9 (1892) 250; NEHRING, Sitz-Ber. Nat. Ges. Freunde (1894) 223, fig. 224. (Balabac); ELElRA, Contr. Fauna Filipina (1915) 247; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 39; Proc. U. S. Nat. Mus. 46 (T913) 339; ELLIOT, Field Col. Mus. Pub. Zoo!. Ser. 1, No. 3 (1896) 76.

Tragulus javanicus BOURNS and WORCESTER, Minn. Acad. Nat. Sci. Occ. Papers 1 (1894) 63 (Balabac); EVERETT, Proc. Zoo!. Soc. London (1889) 223.

Description of the species.-{From the type description.) Allied to and apparently about the size of T. napu F. Cuv.; arrangement of colors above much as in that species, but the whole of dorsal and lateral surfaces broadly washed with jetblack hairs white at their bases, then dull orange and broadly tipped with black; nape with indistinct blackish line as in T. napu; face and sides of neck mixed black and dull fulvous; chin with the usual na~d gland~lar patch between rami of lower jaw;

507

CERVIDJE

arrangement of white throat bands quite different from that of allied species, perhaps similar to that of T. stChnleyanus .. all stripes very narrow, sharply defined; anteriorly, on each side of naked space, a short, pure white stripe, which ends abruptly at about level of posterior canthus of eye; these short stripes commonly separated from each other and from posterior stripes by a dark browln space, the break in their continuity with the latter not less than 1 inch in length; posteriorly the median white stripe, which is very narrow and scarcely broader behind, is bounded on each side, between lateral white stripes, by deep jet-black, fur strikingly different from fur in corresponding position in other species; behind, the stripes separated from the white patch between the forelimbs, by a broad blackish band; belly hairs broadly tipped with black, but inner sides of thighs, as usual, white. The specimen is young with milk teeth; the second upper premolar is much smaller than in a similar specimen of T. napu of corresponding size; third upper premolar slightly smaller, while fourth and first molar are of about the same anteroposterior diameter. SimHarly, below, second and third premolars each much smaller than, in T. napu, while fourth premolar and first molar are of about the same size. Measurements of Tragulus nigricans Thomas." mm.

Second upper premolar, anteroposterior Third upper premolar anteroposterior Fourth upper premolar, anteroposterior First upper molar, anteroposterior Second lower premolar, anteroposterior Third lower premolar, anteroposterior Fourth lower premolar, anteroposterior First lower molar, anteroposterior

7.0

7.5 5.8 6.5

5.7 6.6 8.0 7.2

â&#x20AC;˘ The type is immature and the skull is in such a state that no measurements are given. except those for teeth.

Remarks.-The species is known definitely from Balabac only, the type locality. I have seen a live specimen purporting to come from southern Palawan. Specimens of the species have been taken to Palawan and will likely become established there. The type, No. 91. 11. 28. 2. of the British Museum, was collected by the Steere expedition to the Philippines in 1887-88. Family CERVIDJE Antlers present in the male deer usually; when absent, canine teeth greatly enlarged; a facial gland below eye in the

508

PHILIPPINE LAND MAMMALS

antlered forms, absent in others; lateral hoofs retained, representing the lateral metacarpal bones. Usually three Joints supporting lateral hoof; canine teeth present or absent; liver usually without a gall bladder except in the musk deer. It is highly probable that a family distinction will come to be generally recognized between the musk deer, of the subfamily Moschime, and the subfamily Cervime. In the Philippines the genus Rusa alone is present. Genus RUSA H. Smith

Rusa H. SMITH, In Griffith's Animal Kingdom subgenus).

(1827) 309 (as a

Generally resembling Cervus,. antlers usually simply forke.d at extremities with second and third tines wanting; bases and tines usually rounded. Pelage generally uniform or spotted at all seasons and all ages, or intermediate between the two, without light-colored caudal disk or black-bordered white area on buttocks; throat and neck frequently heavily maned; ears large; tail relatively long; naked portion of muzzle large, extending to upper surface of face and not constricted above its junction with upper lip; face gland frequently very large and capable of eversion; upper canines small or wanting; upper molars tall, crowned with a small additional column on inner side; size from small to large. The number of actual species in the Philippines is uncertain and large collections must be made before any complete interpretation of the forms described can be made. Hollister * recognizes eight species, as follows: Rusa philippinus (Smith), R. barandanus (Heude), R. francianus Heude, R. basilanensis Heude, R. nigellus Hollister, R. alfredi ' (Sclater), R. calamianensw (Heude), and R. nigricans (Brooke). It is highly probable that several of the species of deer described by Heude from the Philippines are referable to Rusa. philippinus (Smith). However, the descriptions as well as the illustrations of these forms named by Heude show, in many cases, distinct variations in size and shape of the horns, as well as in the dentition. Some of the skulls may be from long-captive specimens, and probably others are crosses between species from various islands. The people in the Islands are fond of keeping live deer and frequently one sees deer being transported

* Proc. U. S. Nat. Mus. 46 (1913) 331-333 and 339-340.

RUSA

509

from one island to another on interisland boats. That some deer are able to cross is proved by a statement made by Brooke* that Cervus alfredi, female, and Cervus' nigricans, male, produced an offspring which resembled the mother more than the father. The following species described by Heude purport to come from Luzon, but it is probable that due to mixed locality data, and errors in labeling plates, some of these nominal forms are from other isla:Q.ds. USSA GORRICHANUS Heude. Ussa gorrichanus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 21, pI. 1, fig. 3; pI. 2, fig. 6; pI. 6, fig. 4. Described from a living cap;tive specimen from Batangas, apparently a typical Rusa philippinus. USSA NUBLANUS Heude. USS(1) nublanus HEU9E, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 24, pI. 2, figs. 7, 7 bis; pI. 7, figs. 3, 3'. This species is suppressed in a later work and the figures here listed are given under the name U ssa tuasoninus Heude. It purports to come from Laguna Province. USSA TUASONINUS Heude. Ussa tuasoninus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt 1 (1888) 25, pI. 1, fig. 9; pI. 9, fig. 2; 4 pt. 3 (1899) 137. See under U. nublanus and U. longicuspis. Th~ figures given for U. nublanus, U. longicuspis, and U. tuasoninus agree in general conformation of the skull. The horns of the first two are thickened and much heavier than those in the figure of the third. USSA LONGIOUSPIS Heude. Ussa longicuspis . HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 34, pI. 2, fig. 2; pI. 9, figs. 1, 1'. This species in a later paper is regarded as a synonym of U8sa tua.soninus and the name regarded suppressed. USSA SPATHARlUS Heude. Ussa spatharius HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 25, pI. 2, fig. 6; pI. 10, figs. 3, 3';. 4 pt. 3 (1899) 137 pI. 1', figs. 2, 3; pI. 14, fig. 3; pt 15, fig. 1. This species purports to come from La Laguna province. The second paper listed regards the' following species U. brachyceros as a synonym. USSA BRACHYCEROS Heude. Ussa brachyceros HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 36, pI. 1 8 , fig. 5; pI. 15, fig. 1. This form is later regarded a synonym of Ussa spatharius (vide supra). Type locality Batangas.

* Op.

cit. 59.

510

PHILIPPINE LAND MAMMALS

USSA RAMOSIANUS Reude. Us sa 1"amosianus REUDE, Mem. Rist. 'Nat. Emp. Chinois 2 pt. 1 (1888) 36, pI. 1 fig. 4; pI. 14, figs. 1, 2; 4 pt. 3 (1899) 135, pI. 7, fig. 4' (1); pI. 1 8 , fig. 3. The porti-on of the type description as regards color is suppressed in the later paper. Th.e type locality of this form is probably doubtful. Reude says: "Ce cer! paralt commun a la Nueva Ecija. Je la tiens des j ardiniers du Jardin Botanique de ManiUe et de Mariano Ramos, ouvrier a l'Ateneo MunicipaL" 8

USSA AMBROSIANUS REUDE. Ussa ambrosianus REUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 'l.7, pI. 3, fig. 5; pI. 11, fig. 2; 4 pt. 3 (1899) " 135, pI. 7, fig. 1. This form, which Reude states is the smallest on Luzon, recalling Mindanao forms, was sent to him in China from Nueva Ecija. Judging by the description and figures this species may represent a form differing from Rusa philippinus (Smith) but without additional material this point cannot be determined. USSA MACARIANUS Reude. U ssa macarianus REUDE, Mem. Rist.. Nat. Emp. Chinois 2 pt. 1 (1888) 28, pI. 1, fig. 6; pI. 10, fig. 1; 4 pt. 3 (1899) 135. The type locality of this species is Nueva Ecija, Luzon. The figure shows horns much elongated, bending inward at the tips considerably, their length double the length of the skull. USSA ElLORZANUS Reude. Us sa elorzanus REUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 28, pI. 2, fig. 5; pI. 7, figs. 4, 4'; 4 pt. 3 (1899) 135, pI. 15, fig. 4. The second paper suppresses the text in the Itype description regarding the color of the animal. The type locality is Bataan. The figures represent an animal with rather slender and slightly divergent, horns. U SSA GARCIANUS Reude. Ussa garcianus REUDE, Mem. Rist. Nat. Emp. Chino is 2 pt. 1 (18.88) 29, pI. 1, fig. 2; pI. 10, fig. 2'; pL 14, fig. 4; 4 pt. 3 (1899) 135, pI. 1', fig. 4 (in vol. 2). Reude 4 pt. 3 (1899) 134 gives pI. 10, fig. 2 (1) female as U. verzosanus! The type locality of this form is Nueva Ecija. U SSA GUIDOTEANUS Reude. U ssa guidotean~ÂŁs RElUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 2'9, pI. 2, fig. 1; pI. 15, fig. 3; 4 pt. 3 (1899) 138. The type locality is Batangas. USSA ROSARIANUS Reude. Ussa rosarianus REUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 29", pI. 1 n, fig. 1; pI. 14, fig. 6. This form is not listed in the second paper but the plate references are given as belonging to U ssa verzosanus Reude. The type locality is Nueva Ecija.

RUSA

511

USSA NIGRICANS Heude. Us sa nigricans HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 31, pI. 2, fig. 4; pI. 6, fig. 3; pI. 13, 'figs. 5, 6. Heude believes the specimens here described to be identical with Brooke's Cervus nigricans. It appears to be fairly well established by Lydekker, Deer of All Lands (1898) 159-161, that the type locality of Ce1'vus nigricans is very probably Basilan and ~he distriBu.tion is given as this island only. Heude's specimens were from J alajala, Rizal which is in Luzon. He states: "Le pelage de ce cerf est noir uniform." It was probably on the basis of color that he connected the specimen with Brooke's species. He makes no mention of the small size. U SSA MARAISIANUS Heude. Ussa ma'raisianus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 31, pL 6, fig. 2; 4 pt. 3 (1899) 138. Heude regards this form as separable from U. nig1-icans only on the basis of color. Both forms are from the same immediate locality, J alajala, Rizal Province. USSA DAILLIARDIANUS Heude. Ussa dailliardianus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 32, pI. 1, fig. 10; pI. 7, fig. 2; pI. 13, fig. 3; 4 pt. 3 (-1899) 138. Heude in the type description states thalt the type specimen is from Jalajala, Rizal, but in the second paper states that it is not from Jalaj ala, Rizal, but certainly is from Luzon. According to Heude it is one of the largest species. USSA MARZANINUS Heude. Ussa marzianus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 33, pI. Z, fig. 3; pI. 13, fig. 4; 4 pt. 3 (1899) 138. The type locality of this species is Infanta (Tayabas?) not Laguna and Nueva Ecija as given by Hollister. USSA ROXASIANUS Heude. U ssa 1'oxasianus REUDE, Mem. Rist. 'N at. Emp. Chinois 2 pt. 1 (1888) 33, pI. 2, fig. 8; pI. 13, figs. 1, 2; 4 pt. 3 (1899) 138. Heude in the second paper suppresses, pI. 2, fig. 8. "Supprimer Ie texte relatif au pelage qui m'est inconnu, ainsi que Ie dents. Celles qui ont ete figurer sous ce nom ne lui appartiennent pas." Type locality Batangas, Luzon. USSA MICRODONTUS Heude. U ssa microdontus REUDE, Mem. Rist. Nat. 'Emp. Chinois 2 pt. 1 (1888) 34, pI. 1 n, fig. 8. This species was established on a lower jaw of a specimen believed to have been taken in Batangas. Heude's second paper does not menltion this species so one presumes that the name and description were suppressed. USSA GONZALINUS Heude . â&#x20AC;˘ Ussa {lonzalinus REUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 35, pI. 1, fig. 7; pI. 9, fig. 3; 4 pt. 3 (1899) 138.

512

PHILIPPINE LAND MAMMALS

This form is founded on a single skull said to have much the same characters as U. gorrichanus. The type locality is not given. One presumes that iJt is from Luzon. USSA TELESFORIANUS Heude. Ussa telesforianus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 36, pI. 1, fig. 8. _, This species is Jl!ot mentioned in the later paper and it may be that Heude no longer recognized it as a distinct species. It was originally founded on a lower jaw preswmably from Batangas, Luzon. USSA CORTEANUS Heude. Ussa corteanus HIDUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 37, pI. 1·, fig. 3; pI. 10, fig. 2; pI. 11, fig. 1; 4 pt. 3 (1899) 134 (and in vol. 2, pIs. 6, fig. 1; pil. 1, fig. 1). This species is founded on specimens from Bataan and Nueva Ecija. USSA RUBIGINOSUS Heude. Ussa rubiginosus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 37, pI. 1, fig. 1; pI. 6. fig. 1. This form is not recognized in Heude's second p'a per but the figures here given are referred to Ussa cortea,nus. The type l'ocality is Bataan and Nueva Ecija. U SSA HIPOLITIANUS Heude. U ssa hipolitianus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 39, pI. 8, fig. 1; 4 pt. 3 (1899) 138. This species is from Laguna and Baltangas. USSA CHRYSOTRICHOS Heude. Us sa chrysotrichos HIDUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 39, pI. 15, fig. 2. In Heude's second paper this form ,appears to be regarded as a synonym of Ussa atheneensts since he relegates the figures given [PI. 15, fig. 2 (See page 141 of second paper)] to the latter species. The type locality is Laguna and Batangas. USSA GUEVARANUS Heude. Ussa guevaranus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 40, pI. 15, fig. 4'. Heude in his second paper does not re.tain this species and the figure given is referred to Ussa elorzanus. The type locality is Mariquina, Rizal Province. USSA VERZOSANUS Heude. Ussa verzosanus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 37, pI. 1·, fig. 2: pI. 14, fig. 3: 4 pt. 3 (1899) In vol. 2 pt. 1 (1888) pI. 14, fig. 6; pI. 1·, fig. 1; plI. 10, fig. 2. The type locality of this form is "Nueva-Ecija, au nord." The text of the type description is suppressed in :the second paper. , USSA MICHAELINUS Heude. • 3 Ussa michaelinus HEUDE, Mem. Rist. Nat. Emp. -Chinois 4 pt. (1899) pI. 32 A, figs. 1-5. The type locality is Tarlac, Luzon.

RUSA

513

USSA VILLEMERIANUS Reude. Ussa villemeriamts REUOID, Mem. Rist. Nat. Emp. Chinois 4 pt. 3 (1899) pI. 32 A, fig-so 4, 6, 7, 10. The type locality is San Mig-uel de Tarlac, Luzon. USSA VIDALINUS Reude. Ussa vidalinus REUDE, Mem. Rist. Nat. Emp. Chinois 4 pt. 3 (1899) pI. 32 A, figs. 8, 9; pI. 32 B, fig-. 9. Type locality, San Miguel de Tarlac, Luzon. USSA BARYCEROS Reude. Ussa bar.yceros REUDE, Mem. Rist. Nat. Emp. Chinois 4 pt. 3 (1899) 139, pI. 32b, fig. 1. Type locality is Laguna and Batangas, Luzon. SIKELAPHUS SOLOIDNSIS Reude. Sikelaphus soloensis REUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 3 (1894) 147. Type locality Jolo. This form, of which I have seen only living specimens, suggests relationship with some Asiatic species. It is known that gifts of animals have been made in the past to former Sultans of Sulu by rulers of other Asiatic states. r am of the opinion that the Jol0 deer is a direct descendant of some Asiatic species.

The following forms may be synonyms of Rusa alfredi Sclater; of this, however, there .is no certainty. MELANAXIS MASBATENSIS Heude. Melanaxis masbatensis HEUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 47, pI. 3, figs. 1, 1', 4. This species, the rtype locality of which is Masbate, is founded on the cranium of a female kept in captivity. MELANAXIS BREVlCEPS Reude. Melanaxis b?'eviceps REUDE, Mem. Rist. Nat. Emp, Chinois 2 pt. 1 (1888) 47, pI. 1", fig. 6; pI. 14, fig. 5. This species is likewise founded on female skulls, two specimens said to be from Masbate. MELANAXIS ELEGANS Reude, Melanarcis? elegans REUDE, Mem. Rist. Nat. Emp. Chinois 2 pt. 1 (1888) 49. Reude founds this species on a pair of antlers and a frontal fragment of unknown origin in the collection of the Ateneo de Manila. USSA CINEREUS Reude. Ussa cinereus REUDE, Mem. Rist. Nat, Emp. Chinois 4 pt. 3 (1899) 14'0. Ussa crassico1'nis REUDE, op. cit. 2 pt . .1 (1888) 23, may represent a recognizable species. Matschie, Sitz-Ber. Ges. Nat, Freunde Berlin No.5 (1898) 43, identifies a Samar specimen as C. c?"assicornis. RUSA PHILIPPINUS (Smith)

Cerf des Philippines DESMAREST, Mamm. (1820) 442 (Luzon); CuVIER, Oss. Foss. ed. 2 4 (1821-24) 46. 2511463-83

514

PHILIPPINE LAND MAMMALS

Cervus philippinus S'MITH, Griffith's Anim. Kingdom 4 (1827) 147; 5 (1827) 33 and 319; PuCHERAN, Rev. et Mag. Zooi. II 7 (1855) 49; II 9 (1857) 481, pI. 15 (Manila); BROOKE, Proc. Zool. Soc. London (1877) 51, pI. 8. Rusa philippi-nus GRAY. Knowsley Menag. (1850) 63. ? Cervus mariannus FRASER, Proe. Zoo!. Soc. London (1866) 367; HOLLISTER, Phtlip. Journ. Sci. § D 7 (1912) 40. Ce?'vus philippinensis MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde Berlin No.5 (1898) 43.

Description of the species.-Small deer; hair harsh, of moderate length; tarsal gland plainly marked; ear of moderate size, covered externally by short, close-set hair; horns remarkably rough and massive; a powerful long brow antler and one short tine directed inwards. Color.-General color rich, ruddy brown, darkest on back and lightest on neck; forehead and cheeks rufous-fawn color; dark, almost black, str€aks commence over each eye, and, blending together, form a band running down the cent€r of the face, which is separated from the dark moustachelike mark bordering the muzzle by a narrow track of pale fawn; breast and belly uniform brown; tarsal gland marked with a rufous spot. Shdl.-Anteorbital fossa very deep, suddenly pressed in anteriorly to orbit and occupying the entire facial plate of malar; facial portion of skull compressed laterally and deeply from above downwards, canines present. M easurements.-Height at shoulder, 684 millirpeters; length of skull, about 2.87. Remarks.-The type was collected in Luzon by Dussumier in 1820. It was taken alive to Paris and was preserved in the Musee d'Histoire Naturelle. Sir Victor Brooke gives a full page drawing of the species in color. His specimen, of unknown origin, was obtained from Rotterdam, presumably having been taken there from the Philippines . • Whether or not Cervus mariannus Desmarest is identical with this species appears never to have been fully decided. Quoy and Gaimard, * Zoologists of the Uranie, write concerning the deer on Guam: Une petite espece de eerfaxis, qui a ete apporwe des Philippines, a tellement multiplie, que l'on ne eonnoit pas de lieu qui en eontienne porportionellement davantage; car il existe a Guam plus de mille de ees animaux.

* Zoo!.

Voy dey l'Uranie (1824) 33.

515

RUSA

On nourrit de leur chair les equipages des navires qui touehent a eette ile, et Ie notre n'eut presque pas d'autres vivres pendant Ie temps que nous y demeuriames. . . . Cet arehiple n'a qu'um rnarnrnifere que ne lui ait pas ete apporte.

It has been pointed out that the specimens from Guam do not have canine teeth. While unquestionably a Philippine species, it is impossible to be certain whether it is identical with Ru,sa philippinensis or some other Philippine deer. It is, how~ ever, the opinion of Sir Victor Brooke and Lydekker* that the species are identical. Hollister lists specimens from Abra Province, lIocos Sur Province, and Manila. It appears to be confined to Luzon. RUSA ALFREDI

(Sclater)

Cervus alfredi SCLATER, Proe. Zool. Soc. London (1870) 381 pI. 28; (1871) 478; (1872) 24; BROOKE, Proe. Zoo I. Soc. London (1877) 59; (1878) 902; MElYER, Proe. Zool. Soc. London (1879) 666; LYDEKKER, Deer of all Lands (1898) 172. Rusa alfredi HOLLISTER, Philip. Journ. Sei. ยง D 7 (1912) 40; Proe. U. S. Nat. Mus. 46 (1913) 339. Melanaxis alfredi HEUDE, Mem. Rist. Nat. Ernp. Chinois 2 pt. 1 (1888) 47, pI. 3, figs. 2 and 3.

Desc'r iption of the species.-SmalI; hair soft, of moderate length; ears and tail short, the former almost naked externally; horns not greatly exceeding head in length; a short brow antler, and a very short tine directed inwards. Height at shoulder about 643 millimeters. Color.-General color rich, dark brown with scattered ruddy tints; cheek and occiput pale brown; lower jaw and throat dirty white; neck and shoulders anteriorly uniform dark brown; posterior parts of shoulders, back, and sides distinctly marked at all seasons of the year by yellowish white spots; breast and belly yellowish white ;.limbs externally, above carpal and tarsal joints, uniform brown, internally dirty white; below carpal and tarsal joints pale brown; adult female resembles male in color; young spotted at birth. Remarks.-The type specimen was formerly the property of H. R. H. the Duke of Edinburgh who received it from a Spanish gentleman in Manila. The skin and skeleton of the type are now in the British Museum. The skull is said to resemble that of Rusa nigr'icans more than that of R. philippinus. * Deer of All Lands 158.

516

PHILIPPINE LAND MAMMALS

Hollister notes specimens from Cebu, Guimaras, Leyte, Masbate, Negros, Panay, and Samar. The exact type locality is unknown. RUSA BARANDANUS (Henne)

Ussa barandanus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 22, pI. 1, fig. 5; pI. 8, fig. 3; 4 pt. 3 (1899) 138 (Mindoro) . Rusa bumndanus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1912) 331 (Mindoro) .

Description of the species.-(After Heude.) The horns measure (following the curve) one-fourth longer than base of cranium; horns and their basilar branches symmetrically curved on the inside; grooving of horns very distinct, deep, and decorated here and there with tubercles; color ashy white; horns as well as peduncles thick and robust; peduncles diverge remarkably, distance between them under crown of burs is 90 millimeters (not 90 centimeters as Heude states). The cranium is remarkable for the swelling in the frontonasal region which resembles the forehead of a sheep; forehead widened between peduncles; wide and flattened in postorbital region; fronto-occipital angle 30째 more open than in ordinary species; nasals project strongly anteric>rly; lachrymal fossa very high and deep; anterior part of palate wide, deepened and bordered by high edges; basilare straight, with a median keel. Canines persistent; upper premolars very high and show considerable lateral compression; length of molar-premolar series, 84 millimeters; premolars alone, 35 millimeters. Remarks.-This species was founded on two specimens, 'Presumably from Mindoro. There are seven specimens which Hollister refers to this form in the United States National Museum collected by J. C. Le Hardy in Mindoro. I have examined a pair of antlers of this form from Calapan, Mindoro (specimen in the University of the Philippines) having the following measurements: Measurements of Rusa bU7'andanus (Heude). mm.

Base of bur to tip of first tine Circumference at bur Bur to tip of horn Length of outer tine Length of inner tine Height of peduncle, inner measurement Circumference of peduncle Length of frontal suture

160 260 118 31 24

100 95

517

RUSA Meastu'ements of Rusa bamndanus

(Heude)~Continued.

rom.

Int.erorbital width Width between supraorbital foramina Frontal suture to back of occipital Distance between tips of first tines Distance between tips of second outer tines Distance between tips of third inrier tines

44 70 155 202 152

The horns are relatively short, the "beading" very rough, extending to the upper branch, The channeling shows to the tip. RUSA FRANCIANUS (JIeude)

Us sa francianus HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 24, pI. la, fig. 7 (3 premO'lar teeth). Rusa francianus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 331.

Description of the species.-This was established by Reude on a specimen purporting to have come from Mati, a town on the Pacific coast of Mindanao. The specimen consisted of "une peau avariee, les maxillaires, et un frontal, portant seulment des pedoncles." The characters published offer the following facts: Color uniform chocolate brown; peduncles large, rounded, somewhat flattened above temple and narrowed toward orbit; incisive foramina large, well developed; anterior portion of palate large and hollowed out; superior milk canines have left no trace, if they have ever been present; total lengths of upper tooth row, 78 millimeters; premolar series, 31; from first premolar to tip of maxillary, 82. There are twenty-three specimens in the United States National Museum from Mindanao, which are listed by Hollister un'd er the species. He states: There are apparently two forms of deer represented in the above series, or the variation in the size of the teeth in the common Mindanao Rusa is much greater than is usual in animals (j)f this family. Without more material from other parts of the island, and! especially topotypes of fr,ancianus, from Matai, Gulf of Davao, it seems undesirable to attempt to divide the species into local l'aces.

I have three specimens of deer from Saub on the Cotabato coast (two skulls and one pair of antlers with frontals), which may belong to this species. Hollister examined the specimens. After comparison with the National Museum specimens he was not prepared to say tD what species they belonged. He suggested that while they might be new to science, he left that the material

518

PHILIPPINE LAND MAMMALS

was inadequate to warrant a description. Then again they might be merely forms of franciarvus. For the time being I shall consider them as belonging to this species. RUSA BASILANENSIS (Rende)

Mela,naxis ? ba,sila,nensis HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 49. Rusa, ba,sila,nensis HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 332.

Heude founded this species on a mutilated cranium from Basilan. Hollister has identified a pair of antlers collected by Mearns in Basilan, as belonging to this species. The 'd escription given by Heude is meager and it is probable that this form is identical with Rusa alfredi, or only a subspecific form of that species. Hollister states: There are evidently two species of deer on Basilan. One, Rusa, wigricans (Cervus steerii Elliot), is fairly well known. It is a much smaller species than is represented by the frontals and antlers collected by Doctor Mearns at Isabela. The species described by Heude was based on a mutilated skull, which was doubtfully supposed to represent a species related to R. a,lfredi of the more nortlilern islands. From the excellent figure of this skull and the antlers, I judge Doctor Mearns's specimen to be the same. If this is so, the species is a member of the group of larger Philippine deer, including philippinus of Luzon and fra,ncianu8 of Mindanao. RUSA

NIGELLUS

Hollister

Rusa, nigellus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 332, pI. 29, 3 figs. (skull and teeth).

Description of the species.-A small deer, apparently related to Rusa nigricans, but with prominent facial markings, smaller skull, and much smaller teeth; much smaller than the common deer of Mindanao, Rusa francianus. Color.-Face brownish black, marked by two transverse bands of golden brown, the first over muzzle, 10 millimeters above nose pad; the second across forehead, bowing to each horn; lower lip whitish; chin blackish; throat dark brown with median longitudinal stripe of pale cinnamon; nape and back of ears black; lining of ears buffy white; neck and body blackish brown (mummy brown of Ridgway), mixed with numerous blackish hairs; middle of belly darker, more blackish; pectoral and inguinal regions ochraceous-buff, sharply marked from general color of middle underparts, legs, and sides; outer sides of forelegs blackish, a narrow stripe of bright ochraceous down inner side to hoofs; hind legs blackish above, with a narrow blackish

519

RUSA

stripe in front extending to near hoofs; a wide area of ochraceous on back side from region of knee to lateral hoof. Skull and teeth.-Skull apparently much as in R. nigricans but smaller, the adult male slightly smaller than the female of R. nigric(}'n8; antler pedicles very long, even longer than usual in this group of rusine deer; teeth much smaller than in R. nigric(}'n8; the antlers of the type are small with very short brow tines; a shed antler from Mount Malindang apparently of this species, is much the same; it measures 115 millimeters long from the burr, without fork, an'd with a short brow tine of only 21 millimeters length on upper surface. (After the type description.) Measurements of the type of Rusa nigellus Hollister. mm.

Head and body, from skin Tail Ear, moistened Skull: Total length, estimated Zygomatic breadth Palatal length Length of median frontal suture GreateSit length of nasals Greatest width of nasals Least width of nasals Anterior point of nasals to end of premaxilJre Orbit to end of premaxillre Length of horn pedicle from orbit Upper molar-premolar row Upper molar row Upper premolar row Greatest width of second upper molar Lower molar-premolar row

1,000 80 80 214 92.5 124 78 67 34.5 13.8 29.5 108.5 72.5 66 41 27.5 13.6 72

Remarks.-The type and one shed antler were collected on Mount Malindang, Mindanao, by Edgar A. Mearns, at an elevation of 8,000 feet, May 30, 1906. The great extension of the horn pedicle suggests an approach to the condition which obtains in the Muntjac of India, otherwise there appears to be no relation. The canine is apparently absent. RUSA CALAMIANENSIS ( Reude)

Hyelaphus calamianensis HEUDE, Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 49 (CaIJamian, P. L) pI. 4, figs. 1, 2. Cet'vus ____ _____ ___________ ELLIOT, Field Col. Mus. Pub. 1 (1896) 68 pIs. 6 and 7 (Culion). Cervus culionensis ELLIOT, Field Col. Mus. Pub. (1897) 159 (Culion), LYDEKKER, Deer of All Lands (1898) 173.

520

PHILIPPINE LAND MAMMALS

R usa culi onensis HOLLISTER, Philip. Journ. .Sci. ยง D 7 (1912) 40. Rusa ca,lamianensis HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 339.

Desoription of the speoies.-A small, brown deer ; hair rather coarse and stiff, longest and coarsest on flanks; body rather stout , legs slender, hind ones longest; ears rather long, slightly r ounded at tip; head slender, nose rather long and pointed; hoofs narrow and pointed; ears hairy, hairs on inside long and thick. Color of male.-Nose and sides of face in front and around eyes pale yellowish brown; horn pedicles ochraceous; back of neck, back and sides mummy brown, hairs being whitish at base, t hen a ring of ochraceous, and tip black, sometimes tip ochraceous; general color of body mummy brown with a wash of ochraceous ; from between shoulders to root of tail a black dorsal stripe about 25 millimeters in width; end of upper lip yellowish white surrounded by a band of pale brown, and space between jaws beneath yellowish whi'te; ears Mars brown, hairs tipped with ochraceous; front of neck pale yellowish brown like sides of face; chest, shoulders, underparts, except belly and outside of thighs, purplish brown, with many long white hairs intermingled with those of other color; belly and inside of thighs white; forelegs blackish brown, inclining to ferruginous behind; hind legs blackish brown with a long narrow whitish stripe in fron't of knees; hoofs black; horns in velvet whitish brown; end of nose and center of upper lip naked, jet-black; tail rather bushy, mummy brown above, white beneath. Color of female.-General color throughout pale yellowish brown, darkest in center of back and top of head; forelegs Prout's brown on outside, inside russet; hind leg russet; stripe in front of knee dark buff; center of belly yellowish white; tail bushy, russet above, white beneath; hoofs black; ears dark brown outside, hairs tipped with buff; inside white. Skull and teeth of male.-Roof of cranial cavity arched, rising but slightly at forward portion above plane of face, but declining sharply from junction of frontals, parietals, and supraoccipitals; general shape rather narrow, with a moderately long face, diminishing rapidly in width from anterior edge of first premolar; premaxillre very long, from their upper posterior borders to extreme anterior edge equal to length of nasals; distance from anterior end of nasals to end of premaxillre 45 millimeters; anteorbitaI fossa occupies nearly all lachrymal; it is deep and measures in length about seven-eighths the diameter of orbit; anteorbital vacuity rather large, reaching about three-fourths

521

RUSA

the length of upper border of anteorbital fossa and is 9 millimeters wide in its broadest part; there is a slight sagittal crest commencing at junction of frontals and parietals and extending forward on frontal plateau to middle of orbits; nasals much rounded posteriorly, becoming flatter at their anterior extremity; parietals much longer than broad in its widest part; occipital condyles large; paroccipital processes flat, broad above, tapering rapidly to a point, and curving inward; auditory bullre large and prominent; mesopterygoid fossa short in proportion to length of skull, and auditory bullre extend in width about half its length; teeth of moderate size, with small supplementary columns on molar series; horn pedicles long commencing at upper border of orbit and rising 22 millimeters above cranium, and extending upward and backward from orbit (horns in velvet and probably not full size); burr of moderate size and only slightly exceeding beam in diameter; horns flattened at base, and extend upward and curve, outward for 140 millimeters on left side and 157 millimeters on :right side, both turned inward at tip; a straight brow antler branches off about 25 millimeters above burr and rises for about 75 millimeters, the right being somewhat longer than the left; extreme width between horns at apex 228 millimeters; width between burrs 45 millimeters; coronoid process of mandible greatly curved, and its point extends behind condyle; ramus also greatly curved on its lower border from angle to anterior edge of first lower premolar; mental foramina large, oblique, with exterior borders spread outward. Female skull.-Slenderer than in male, with narrower frontals and slightly larger nasals; cranium arched in a similar manner, with the same sharp decline of parietals to supraoccipital, and with the small sagittal elevation anterior to parietal suture; the depression less in posterior portion of frontal; anteorbital fossa neither so wide nor so deep as that of the male, but anteorbital vacuity is very extensive; audital bullre very large. (Mter Elliot.) Measurements of Rusa calamianensis (Heude)." Total length of skull from tip of nose to lambdoidal suture in straight line Point of premaxillre to tip of nasals Length of premaxilla Length of nasals Anterior rim of orbit to free end of premaxilla â&#x20AC;˘ Measurements reduced from inches to millimeters.

nun.

216 44 64 69 110

522

PHILIPPINE LAND MAMMALS

Measur ements of Rusa calamianensis (Heude)-Continued. mm.

Upper end of nasals Ito frontoparietal suture Across maxilla 1 inch in front of rim of orbit Across maxilla 2 inches in front of rim of orbit Extent of upper molars Length of lower premolar series Length of lower molar series Circumference of burr (error)? b Length of humerus from head Length of ulna, extreme Length of radius Length of metacarpal cannon bone Length of femur from upper border of head to lower extremity of internal condyle Length of metatarsal cannon bone b

75 64 37 36 23 41 12 142 163 125 104 178 124

This circumference stated 0.60 inch.

Remarks.-This species is known only from Culion. The type locality, as given by Heude, is Calamian, one of the Calamianes islands. It is remat;kable that a species of deer should exist in the Calamianes when the genus is wanting in the Palawan group.* RUSA NIGRICANS (Brooke)

Cervus wigricans BROOKE, Proc. Zool. Soc. London (1877) 57, pIs. 9 and 10; (1878) 90~ ? Cervus mariannus SCLATER, Proc. Zool. Soc. London (1870) 279. Ussa nigricans HEUDE, Mem. Hist. Nalt. Emp. Chinois 2 (1888) 31. Cer,vu8 steerei ELLIOT, Field. Col. Mus. Zool. Series 1 (1895) 72, pIs. 8-10; (1897) 157. R usa steeri HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 40. Rusa nigricans HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 4·0; Proc. U. S. Nat. Mus. 46 (1913) 339.

Description of the species.-Small; hair coarse and harsh, spreading out from a circle on top of shoulders; body stout, heavy, with stout, rather short legs, hind legs longer than front legs; ears small, narrow, covered with hair within and without; head long and narrow, especially in nasal region, which is sud'd enly contracted in front of upper tooth row; nose pointed; hoofs broad for their length and heavy wide spread and rounded, instead of pointed at tip. ---- -

- - - - - - -- -- - - -

* At Sir J. Brooke Point, southern Palawan, a number of Borneo deer have been turned loose, with the hope that they might gain a foothold on the island. None has been seen since.

RUSA

523

Color of male.-Nose and sides of face Vandyke brown, tips of hairs ferruginous; a small space above nostrils blackish brown without ferruginous; bare end of nose and center of upper lip brownish black; no evidence of a mustache; forehead, nostrils, sides of upper lip, back of head to anterior border of eyes, and inner horn pedicles black, each hair largely tipped with ferruginous; beneath eyes, cheeks, and outer side of horn pedicles bright ferruginous; ears black behind and white inside; back of neck and line to center of shoulders Vandyke brown; no dorsa~ streak; sides of neck, back, sides of body, outside of legs to ankles, and tail above and below Prout's brown; chin and sides of underlip for half its apical length Vandyke brown; lower sides and between jaws whitish; chest between forelegs, entire belly, and insides of thighs yellowish white with a strong tinge of buff on belly; inside of legs light brown, hoofs blackish brown at base, remainder pale yellow; horns roughened for their entire length, and black except tip of tines; tip of tines and edges of roughened portion of beam yellowish white. Skull of male.-Roof of cranial cavity convex from middle of orbits to supraoccipital, slope being most marked from between pedicles to base of skull; facial portion flat, broad, retaining its width to second premolar, where it narrows rapidly, and is depressed markedly at junction of nasals and maxillre anterior to anteorbital vacuity; anteorbital fossa large and deep, being nearly equal in length to diameter of orbit and occupying nearly all of" lachrymal; nasals broad at their j unction, with frontals narrowing rapidly for one-third their length, then continuing with equal width beyond junction with premaxillre, when they narrow to a point; anteorbital vacuity moderate, longer than wide, being a moderate slit at junction of frontal, nasal, lachrymal, and maxilla; pedicles of moderate height, rather stout, rising behind orbit just in front of parietal suture, inclined backwards, and directed above plane of face and convexity of frontals; audital bullre very small; condyles large; paroccipital process broad at top, compressed laterally, curving inwards, and graduating to a point; premaxilla rather broad, considerably exceeding in length the upper molar series. Teeth large, the molars possessing prominent supplementary columns; canines small, slightly curved and pointed, on upper jaw only; length of parietal suture about equal to its greatest width; coronoid process of mandible long, of about an even width and curving backwards,

524

PHILIPPINE LAND MAMMALS

its upper extremity extending beyond posterior edge of condyle; this last transverse proj ection is broadest and flattest on its anterior border tapering to a sharp edge on exterior; inferior dental foramina pyriform,' pointed end upwards, while mental foramina are oblong, slit lengthwise with mandible; when jaws are closed the first lower premolar lies beyond and in front of first upper premolar. Skull of female.-Cranial cavity like that of male, decidedly convex above, slope most abrupt anteriorly to between orbits, differing in this respect from the skull of the male which has slope most pronounced to supraoccipital. Facial portion rather flat and narrow, and contraction of maxillre at first premolar not so conspicuously apparent as in male; nasals broad posteriorly, narrowing sharply about midway their length, and continue with equal width beyond posterior border of premaxillre; audital bullre quite small; supplementary columns not so prominent as in male; canines short and blunt at point; general appearance of skull compared with male is long and narrow. MeasU1'e~ents

of Rusa 'fI,igricans (Brooke)." mm.

Height at shoulder Height at rump Length from nose to I\;ail Length of ear outside Antler from burr to tip Brow antler outside Brow antler inside Inner tine, inside Circumference of burr Circumference of beam at base Skull of male: Total length Tip of premaxilla to tip of nasal Length of nasals Anterior rim of orbit to tip or premaxilla Upper end of nasals to frontoparietal suture Width of skull one inch in front of orbit Width of skull two inches from orbit Extent of upper pre'm olars Extent of upper molal'S Extent of lower premolars Extent of lower molars Length of premaxilla

570 818 1,118

89 228 64 41.5

51 99 86.5

222 30 72 114.4 83

72 30.2 32

44 33.2

47 53

â&#x20AC;˘ Measurements and description from EDiot's description of CeTvus ateerei.

525

RUSA

Measurements of Rusa nig?'icans (Bt路ooke)-Continued. Skull of female: Total length Tip of premaxilla to tip -of nasals Length of nasals Anterior rim of orbit to top of premaxilla Upper end of nasals to frontoparietal suture Width of skull one inch from orbit WidJth of skull two inches from orbit Extent of upper premolars Extent of upper molars Extent of lower premolars Extent of lower molars Length of premaxillre

rom.

228.4 34 76

120 76 70 48 28 35

28 43 46

Re1na1~ks.-The

type of R. nigricans was received in London from the Philippines in 1870, the collector and exact locality being unknown. It was kept alive. A male hybrid was produced between this animal and a male Rusa aZfredi. In external appearance the hybrid was said to resemble the mother more than the father. Sclater doubtfully referred this specimen to R. rnarianna. while it was still living. Rusa nigricans is said to resemble R. alfredi of Mindanao, more than any other known form, particularly in its short naked ears, flattened and wide face; it differs, however, in being lighter in build and uniform dark brown. Elliot's specimen which has furnished this description is compared and certain differences are pointed out, particularly the shape of the anteorbital vacuity Which differs, being quite sharply pointed at its lower end, and the anteorbital fossa is larger and deeper; the color of the coat is said to be much brighter than Brooke's nigricams. With the known material, it does not seem feasible to separate the two forms. The type of Elliot's R. steerii is from Basilan, while the type locality of R. rvigricans is merely "Philippines." Elliot's specimen was collected by Steere who writes of its presence in Basilan as follows: The Basilan Cervus was found occupying the higher parts of the island, in a country of steep, rocky ridges, covered thickly with timber and thick undergrowth. In the valleys were shallow streams. I took it rather for a mountaineer than a swamp inhabiter, though the whole country

* Elliot, op. cit. p. 75-76.

526

PHILIPPINE LAND MAMMALS

was dripping with moisture at ithe time of my visit, and the higher lands in the Philippines are always the swampiest. We have one male mounted in the museum of the University of Michigan. I noticed that all the deer procured in Basilan were closely alike in color, size and character of horns. I believe the skin and skull you have, with a skin of a doe and the buck mounted here, were all the specimens we procured from Basilan. I have an impression that the same differentiation of species of Cervus exists in the Philippines. as I found so prominent in many genera of birds.

Family BOVIDJE Incisors persistent only in the lower jaw; horns persistent in both 路sexes, formed by a long bony projection covered with horny material; hind legs longer than forelegs; stomach with four compartments. Genus BUBALUS Smith Bubalus H. SMITH, Griffith's Cnvier's Animal Kingdom 5 (1827) 371 (subgenus of Bos); HOLLISTER, Proc. BioI. Soc. Washington 24 (1911) 191.

Skull much less massive than in Syncerus, narrower and more elongated; facial profile nearly straight; rostrum relatively long and slender. Nasal bones elongated and slender, narrowed in the middle, and projecting much beyond end of median suture; thus longest laterally. Vomer fused with palatine bones and palatine plate of maxilla the entire length of median suture. A,udital bullre very small, reaching to about plane of ventral surface of basioccipital. Molars short and high, crowns almost square. Hair of dorsum reversed, directed forward from haunches to head; ears comparatively small, without conspicuous fringes. (Hollister.) In the Philippines there are four recognized species of this genus. B~~balus bubalis (Linnreus), Buaalus mainitensis Heude, and Bubalus moeUendorffi Nehring are large, Bubalus mindorensis Heude is a dwarf species. Buba,lus b~/;balis is the domestic carabao of the Philippines generally. Just when importations of this form began is unknown. The species is present on practically all of the inhabited islands. That certain changes have taken place in the species is evident. The forms one sees in the Visayan islands seem to have developed much shorter horns, while the wild carabao of northern Luzon has very long horns which seem to form a different angle with the skull than in the case of the domestic animals. Persons who have seen the wild carabao on the Calamianes and the domestic carabao about Lake Mainit

527

BUBALUS

are inclined to regard them as recent variations from the typical species. [BUB.A.LUS

MAlN1TJ!:NSIS

Reade]

Bubalus 'l'lULinitensis HEUDE, Mem. Hist. Nat. Emp. Chino is 2 (1888) 205 (Mindanao); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 45; Proc. U. S. Nat. Mus. 46 (1913) 340.

Description of the species.-Heude in a footnote gives only a very meager description of this form. Que ce Buffle soit ice une race locale, une espece representative, il importe assez pen. Ce qu'il y a de certain, c'est qu'il differe notablement du Buffle de Lucon, que ce dernier soit indigene on bien qu'il ait He importe. Le Buffle du lac de Mainit est petit de taille, ales jambes greles; la tete est courte et relativement efflee. Les cornes sont courtes, grosses epaisses peu annelees. Elles se recourbent assez brusquement. Celles des bufflones sont bien plus greles que celles des males, et dans une proporcion que je n'ai pas observee ailleurs. Ce Buffle est actuellement entierement domestique; on Ie reccontre dans Ie district de Surigao, Nord de l'ile de Mindanao; son type Ie plus characterise. se trouve aux bords du lac de Mainit.

It is doubtful whether or not this domestic animal warrants retention as a distinct species. It appears that no specimen of this form has reached any American or European museum to this date, and until such material is available, one can do nothing more than recognize the species tentatively. Trouessart places the form as a synonym of B. bubalus. BUBALUS M(ELLENDORFFI Nehring

Bubalu8 moellendorffi NEHRING, Sitz.-Ber. Ges. Nat. Freunde Berlin No. 8 (1894) 185 (Oct. 1&.) Busuanga r. Calamianes Is. figs. 1-3; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 45; Proc. U. S. Nat. Mus. 46 (1913) 340. BUbalus mollendo?'jJi MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde Berlin No.5 (1898) 43 (Busuanga). Buffelus moellendorffi TROUESSART, Cat. Mam. Suppl. (1904-05) 743.

Description of the species.-(After the type description.) This species is intermediate between Bubalus mindorensis and the wild carabao (cimma'ron of Luzon) and seems to indicate a near relationship with the former; this is chiefly evident in the formation of the lower molars which appear to be merely heavier specimens of the corresponding teeth of B. mindorensis (an exception is found in the third molar which shows a triangular "dentine island"). The teeth of both forms agree in that the lower premolars are almost without cement cover, and the molars also show only a meager cement cover. Very striking and different is the form of the last upper premolar, which

528

PHILIPPINE LAND MAMMALS

bear s a strongly developed accessory cusp, which on the grinding surface appears as an accessory lamina. MeaSU1"em en ts of th e type Bubalus moellendorffi Neh'ring. mm.

Basal length of skull T ota l length of skull Widlth of posterior part of head Width at zygoma Circumference of horn at base Length of horn Length of upper tooth row Le ngth of lower tooth row Leng-th of three upper molars Length of three lower molars Total length of mandible

438 469 211 204 320 500 141 151 88 96 398

Remarks.-Nehring's figure shows a skull that has the general appearance of the typical domestic carabao of the Visayan Islands, and shows but little resemblance to the timarau, Bubalus mindorensis. The horns emerge from the skull laterally nearly at right angles to the long axis of the skull, while in the timarau the horns arise from the frontal region nearly parallel to the long axis. The skull presumably belongs to a wild rather than to a domestic animal, according to von Moellendorff who collected the specimen. BUBALUS MINDORENSIS

Rende

Bubalus 1rl!inroorensis HEUDE, Mem. Hist. NaJt. Emp. Chinois 2 (1888) 4 and 50 (August) (Mindoro); HELLER, Abh. Mus. Dresden 2 (1890) 3 and 31; NEHRING, Zoo1. Anz. (1890) 448; Sitz.-Ber. Ges. Nat. Freunde Berlin (1890) 101; JENTINK, Notes Leyden Mus. (1894) 199, pIs. 8-11; HEUDE, Mem. Hist. Nat. Emp. Chinois (1894) 204, pI. 19E, fig. 19; MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde Berlin No. 5 (1898) 43; MEYER, Abh. Zoo1. Mus. Dresden 6 (1896) 12 pIs. 7 and 8; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 45; Proc. U. S. Natt. Mus. 46 (1913) 333; ELERA, Cont. Fauna Filipinas (1915) 229 pIs. facing pp. '228 and 230. Anoa mindorensis STEERE, in Sclater, Nature 38 (1888) 364 (Aug. 16) Mindoro; OUSTETET Bull. Mus. Paris (1895) 202. Probubalus mindorensis STEERE, Proc. Zoo I. Soc. London (1888) 415 (No. 20); Zool. Gart. (1889) 221; LYDEKKER, Wild Oxen (1898) 128, pI. 10, fig. 1. Buffelus mindorensis TROUESSART, Cat. Mamm. 2 (1898-1899) 988, supp1. 743; THOMAS, Trans. ZooL Soc. London (1898) 410.

Description of the species.-A small buffalo with relatively short legs, horns short somewhat triangular at base, showing a series of r.ough grooves on anterior, posterior, and lateral sides; f

529

"BUBALUS

inner edge very rough, suggesting a beginning of a branch or tine; distal portion rounded, rather sharply pointed and converging; frontal region slightly bulging; ears moderate in size. Measurements of Bubalus mindorensis Heude. mm.

Basal length of skull Total length of skuII Width of lunder part of skull Width of zygoma Circumference of horn at base Length of horn Length of upper molar-premolar series Length of lower molar-premolar series Length of upper molars Length of lower molars Total length of lower jaw

353 381 171 163 290 380 106 117 62 63 320

mm.

352 380 165 270 400 107 121

327

Remarks.-This form appears to have been recognized almost simultaneously by Reude and Steere, and, strikingly enough, each published on the form under the specific name of mindoremis. The descriptions were both published in August, 1888, that of Steere's August 16, while the actual date of publication of Reude's paper is unknown. Reude is usually given credit for the species. This buffalo had been known ~ny years before this time, but it had been confused with the anoa of Celebes and appeared in earlier works under the name of Anoa depressicornis. The measurements given are of specimens in Berlin and Dresden published by Nehring. [BUBALUS BUBALIS (Linnreus)]

Bubalu.s bubalis LINN)EUS, Syst. Nat. 10 ed. 1: 72; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 45; Proc. U. S. Nat. Mus. 46 (1913) 333. Bubalus keraba1~ terus MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde Berlin No.5 (1898) 43 (Nueva Ecija and Cagayan).

Thia form is an imported domestic animal and nee'd not be considered in lists of Philippine mammals. 26~463-84

530

PHILIPPINE LAND MAMMALS

â&#x20AC;˘

ADDENDUM

Angel Cabrera, Genera mammaIium, Insectivora Galeopithecia, Madrid (1925), was received after the manuscript was sent to press, and his references are not included in the synonomies. He has shown that the proper generic name for species herein listed under Pachura Selys-Long Champs (1839) to be Suncus Ehrenberg (Symb. Phys. dec. 2. K. 1832). While I have not verified his findings, I presume it to be correct in which case the proper designations will be Suncus occultidens (Hollister), Suncus luzoniensis (Peters), Suncus palawanensis (Taylor) .

INDEX [Names of genera and of new species and new combinations are printed in boldface.]

A Ababa, 57. abramus, Pipistrellus, 295, 296, 298, 299. Vespertilio, 298. Vesperugo, 298. Acanthion, 490. pumila, 491. Acerodon, 58, Ill, 114, 134, 135, 146; distribution of, 136. aurinuchalis, Pteropus, 139. jubatus, 23, 26, 186, 138. jubatus juba:t.us, 23, 26, 136, 137-140, 142. jubatus mindanensis, 136, 138, 142, 144, 146, 168. jubatus, Pteropus, 142. lucifer, 136, 136, 140. lucifer, Pteropus, 136. Acno, 67. Acromys, 477. admirabilis, Vespertilio, 92. adustua, Tryphomus, 27, 473, 476, 477. Aeluroidea, 344. Aeorestus. 278. lEosciurus affinis, 62. requalis. Rhinolophus anderseni. 26. 212. 214., 219, 220. affinis, lEosciurus, 62. Ahmbough. 57. Ahm-mo, 67. ahamobarbus, Sus, 496, 602-604. Sus barbatus, 602. Alalacsin. 67. Alamid. 67. albo:ryensis, Phlreomys, 62, 391. 893. albicauda. Sciurus, 376, 377, 379. albigularis. 419. Epimys, 423. Mus. 27. 423. Ftattus, 27. 68, 416, 417, 419, 420. 423, 424. albipes, Semnopithecus. 61, 336. alecto alecto, Emballonura. 183, 196, 198, 200. Emballonura. 23. 24, 196, 200. Emballonura alecto, 183. 196, 198, 200. N ycticejus. 303. palawanensis, EmbaUonura, 21. 183, 196, 197, 198. Vespertilio (Nycticeus). 23, 196. alexandrinus, Ftattus, 401, 415, 440. alfredi, Cervus, 609, 616. melanaxis, 616. Rusa, 508. 618. 616, 618. 625.

Amac. 57. Ambao, 57. Ambo. 67. ambrosianus, Ussa, 610. Arno. 67. arnplexicaudatum, Pachysoma, 172. Cynonycteris, 172. Pteropus, 172. Pteropus (Cynonycteris), 172. Roussettus, 23. 118, 172, 173. 175. 176. - XlI'Iltharpia, 172. Anathana, 98. anderseni requalis, Rhinolophus. 25, 212, 214. 219. 220. anderseni. Rhinolophus, 212, 214, 219. 220. Hipposideros diadem a, 22, 232, 241-243. 246. Rhinolophus, 213, 219. Rhinolophus anderseni. 212, 214, 219, 220. angolensis, Roussettus (Lyssonycteris), 172. Anoa depressicornis, 629. mindorensis. 628. Anthropoidea, 323, 324, 336. antricola. Hipposideros. 24, 232. 241, 248. Phyllorhina. 24. 241. Aonyx. 69, 345, 352. cinerea, 68, 316, 349, 352, 358. aper montaus. SUB. 57. apicis, Epimys vulcani, 454. Mus vulcani, 27, 454. Rattus vulcani. 466. apoensis. Cynomolgus rnindanensis, 27. 340. Pithecus rnindanensis, 27, 340-342. Tarsomys, 27. 481. 488, 489. Apomys. 27. 68. 401, 408. 459. 463, 469. 489; distribution of, 392. barduB, 27/ 463. hylocretes. 27, 392, 460-462. insignis, 27, 392, 461 , 462, 465, 469. ins ignis bardus, 392, 460, 462-464. insignis insignis, 460, 462, 463. major, 27, 392, 461, 464, 466, 467. microdon, 28, 29, 392, 461. 469. musculus, 27, 392. 461, 468, 469. petrreus, 27, 392, 461, 465, 466. Aquias, 211. archipelagus, Cynopterus, 21, 182, 183, 184, 187, 188. !lrctictis, 68, 59, 346, 354, 360. binturong, 62, 356, 356. whitei, 356-357.

531

532

INDEX

AretoeebuB, 332. Aretoidea, 344. arcuatus arcuatus, Rhinolophus, 24. 212, 213, 224-227. exiguus, Rhinolophus, 213, 225, 227. Rhinolophus, 221, 224, 227-229. Rhinolophus arcuatus, 24, 212, 213, 224-227. typica. RhinolophuB, 224. arietinus, Sus, 495, 496. Arixenia, 323. Artiodactyla, 65, 66, 493, 494. ater? Poephagomys, 62. atheneensis, Ussa, 512. Atherurinm, 490. Augbo, 57. aurinuchalis, Pteropus, 139, 141. Pteropus (Aeerodon), 139. australis, Baimna, 65. Carponyeteris, 122. Maeroglossus, 122. Miniopterus, 254, 266, 268 , 269. Ayong, 57.

B Babuy, 57. Baeulao, 57. bagobus, Bullimus, 27, 411, 413, 473. Baktin, 57. balabacensis, Sus barbatus, 496, 504. Balae-ba1ae, 57. Balmna australis, 65. mystieetus, 65. Balmnoptera mindol'ensis', 65. rostrata, 65. swinhoei, 65. Balatna matinie, 57. Balecon, 57. Bulintong, 57. Balionyeteris, 114, 116. balutus, Pteropus, 28, 149, 150, 168. Bandieota gigantea, 62. barandanus, Rusa, 508, 616. Ussa, 616. barbatus, ahamobarbus, Sus, 502. balabacensis, Sus, 496, 504. palavensis, Sus, 496, 602, 504. Sus, 495, 502-504. vaT. ealamianensis, Sus, 498. Barbourula, 61. ba,rdus, Apomya, 27, 463. Apomys insignia, 392, 460, 462-464. baryeeros, Ussa, 513. Basilan Cervus, 525. basilanensis, Melanaxis, 518. Rattus, 121, 446. Rusa, 508. 618. Sus, 496. basilanus, Epimys, 28, 455. Rattus, 28, 415, 456. Batak, 490. Batomys, 401, 402, 474, 480, 481, 488; distribution of, 481. dentatus, 27, 480, 481, 483, 484. granti, 26. grantii, 480-483.

Batop, 58. bats, Oriental, distribution of the families and the subfamilies of, 110. Bayong, 57. beatus, Croeidura, 27, 79, 83-86. belangeri, Vespertilio, 303, 306. benguetensis, Epimys, 28, 443. Rattus, 28, 415, 441, 443, 445. Bibliography, annotated, 30. bieolor, Hipposideros, 25, 231, 235-237, 239. Hipposiderus, 236. Phyllorhina, 236, 241. Rhinolophus, 236. bilineata, Epeira, 297. Binatang, 57. binturong, Artictis, 62, 355, 356. Viveriu, 355. Bising, 57. bivittatus, Sciurus, 382. blepotis, Miniepterus, 268, 272, 276. Boneia, 114, 116. Booey, tali-rah bali, 76. Bo-ot, 57. Booot, 57, 326. borneensis, Rhinolophus, 217, 218. Bos, 526. Bovidm, 493, 494, 526. braehyeeros, Ussa, 509. Brachyotis, 278. bvaehyotis brachyotis, Cynopterus, 23, 24, 183-185, 187. Cynopterus, 182-184. Cynopterus brachyotis, 23, 24, 183-185, 187. Pachysoma, 184. Bl'aehyul'a, 194. brachyurus, Herpestes, 365. Mungos, 365, 366. Bradypodidm, 67. brevieaudatum, Pa:ehysoma, 185. brevieeps, Melanaxis, 513. browni, Myotis, 22, 243, 279, 288, 291, 292. bubalis, Bubalus, 526, 529. BubaIus, 67, 59, 61, 526. bubalis, 526, 529. bubalus, 527. keraban ferus, 529. mainitensis, 526, 527. mindorensis, 25, 58, 526-529. moll<mdorfii, 526-528. Bubuit, 57. Bucaiao, 57. Buffelus mindorensis, 528. mollendorfii, 527. Bulilit, 57. Bullimus, 27, 58, 401, 411, 419; distribution of, 473. bagobus, 27, 411, 413, 473. luzonieus, 411, 412, 422, 473. bunkeri, Rhinolophus, 22, 213, 222, 225, 228. Bu-ot, 82. Buta-buta, 57. Bu-uc, 57.

INDEX Bu-ut, 67, 326. Buyatat, 67.

o Csabuit, 67, 96. Cabag, 67. vel Panicqui, 14l. Oabao, 57. Cabat-cabat, 57. Cabayo, 57. Cabig, 57. Cabug-cabug, 57. Cmlophyllus, 21l. cllerulea edwa'rdsiana, Pacbyura, 90. caerulesceus, Crocidura, 62. cagayanus, Cynomolgus, 27, 342. Pithecus, 27, 337, 342, 343. Pteropus, 27, 159. Pteropus hypomelanus, 25, 27, 148-150, 158-163, 166. Cagning, 57. Cago, 57, 96. Cagsi, 57. cagsi, SCiUTUS, 367, 38l. Caguan, 96. Caguang, 57. Calabidang, 57. Calaogo, 96. Calambay?, 57. Calamby, 96. calamianensis, Hyelaphus, 519. Insu]mmus, 22, 470, 471, 473. Rusa, 508, 519-52l. Sus, 495, 496, 498, 502-504. Sus barbatus var., 498. Calapnit, 57. calcis, Epimys, 28, 452, 458. Rattus, 28, 121, 415, 446, 458, 459. Calibaga:, 57. Callinycteris, 114. callitrichus, Rattus, 444. Calumbirong, 57. camellii, Cato-Simius volans, 92. Canidm, 344, 348. Canis, 58, 344, 348. familiaris, 348. familiaris sumatran us, 348. pollokensis, 62, 348. capaecini, Vespertilio, 62. Capaccinius, 2'11l. ca'J)istratus, Pteropus, 171. capito, Taphonycteris, 28, 29, 183, 204, 205. Capromys, 396. elegans, 396. Carabao, 57. carbonarius, TaTsius, 23, 325, 327, 33l. Carnivora, 65, 66, 343, 344; distribution of, 345. Carpomys, 401, 471, 472, 474, 475, 480, 483; distribution of, 473. melanurus, 26, 472-475, 483. phmurus, 26, 472-475. Carponycterinm, 119. Carpyonycteris, 120. australis, 122. castaneus, ]dus, 24, 478-480.

533

Catodon m:rcrocephalus, 65. Cate-Simius volans camellii, 92. catus, Felis, 345. caudatus, Dysopes, 32l. cebifrons, Sus, 495, 496, 500. Celmnomys, 401, 402, 406; distribution of, 377. silaeeus, 25, 402, 403. celebensis, Harpyionycteris, 118. philippensis, Sus, 496, 497. Sus, 497. Centetidm, 71, 73. Cephalotes, 176. peronii, 177. cephalotes, Pteropus, 303. Cercopithecus luzonis minimus, 326. Cercopterus, 17l. Cerf des Philippines, 513. Cerivoula, 253. Cervidre, 493, 494, 505 , 507. Cervus, 58, 59, 61, 508, 519, 526. alfredi, 509, 515. Basilan, 525. culionensis, 519. marian nus, 514, 522. nigrican~ 509, 511, 522. philippinensis, 514. philippinus, 514. steerei, 522, 524. steerii, 518. Ceyx melanura, 409. Chrerephon, 57, 112, 115, 310, 311, 314, 316. luzonus , 24, 28, 311, 313-316. plicatus, 311, 314. Cheiromeles, 107, 112, l15, 310, 320. jacobsoni, 320. pru:videns, 320. torquatus, 315, 320-323. cheiropus, Dysopes, 32l. Chevrotain, 494. Chilophylla, 27, 112, l15, 230, 247: distribution of, 243. hirsuta, 27, 243, 248, 249. chinensis, Pteropus, 23, 151-153. Steno, 65. Vespertilio, 63. Chiromeles, 320. torquatus, 32l. Chiromys, 323. Chironax, l16. Chiropeles torquatus, 321. Chiroptera, 65, 66, 72, 107, 108, Ill, 116, 225, 268, 303, 314; distribution of the families of. 108; distribution of Oriental genera of, 114. Chiroptes, 320. Chongo, 57. Chrotomys, 401, 402, 404, 406, 408; distribution of, 377. whiteheadi, 25, 377, 406-408. Chrysochloridm, 71, 73. chrysocomus, ]dus, 444. Chrysonycteris, 230. chrysotrichos, Ussa, 612.

534

INDEX

Cimmaron, 67, 627. cinerea, Aonyx, 68, 315, 349, 862, 368. Lutra, 363. cinereus, Ussa, 613. Cladobates, 99. Cocane, 57, 334. Coele, 67. Crelops, 115. Colago, 67. Colalapnit, 57. coloratus, Epimys, 28, 434. Rattus, 28, 415, 428, 430, 434-436. Colug idre, 91. Colugo, 91, 92. philippinensis, 93. philippinensis, Galeopithecus, 93. Comastes, 278. commissarius, Mus, 27, 478, 479, 481. conchyvorous, Sus, 496, 497. concinnus, Nannosciurus, 368-374. Sciurus, 369. concolor, Rattus, 454. concolor-ephippium, Rattus, 415, 445, 450, 457, 458. Conoryctidre, 67. coronata, Hipposideros, 234. Phyllorhina, 24, 234. coronatus, Hipposideros, 24, 231, 234-236. corteanus, Ussa, 512. crassicornis, Ussa, 513. Crateromys, 26, 391, 397, 399, 400; distribution of, 392. schadenbergi, 57, 392, 397-400. schadenbergii, 326. crinitus, Sciuropterus, 29, 371, 384, 389, 390. crista:us, Macacus, 338. Crocidura, 57, 58, 73, 74, 76, 77, 82, 83, 86. beatus, 27, 79, 83-86. caerulesceus, 62. doria, 91. edwardsiana, 77, 83, 90, 9l. fretida, 91. grandis, 27, 79, 83, 84, 88. grayi, 23, 79, 83, 85-87. halconus, 27, 79, 83, 85-87. horsfieldi, 86. indicus, Sorex, 62. luzoniensis, 80. mindorus, 27, 79, 83, 87, 88. murina, 62. pala wanensis, 21, 79, 83, 85, 88, 89. parvacauda, 21, 79, 83, 84. (Pachyura) luzoniensis, 24, 80. crossi, Paradoxurus, 62. crossii, Paradoxurus, 360 . CrUnOmY8, 401, 402, 4017, 408; distribution of, 377. fallax, 26, 377, 408-410. melanius, 377, 408-410. Cryptoproctinre, 354. Cuayapnit, 57. Cufanit, 57. culionensis, Cervus, 519. PhoJidotus, 68. Rusa, 520.

cumingi, Mus (Phlreomys), 24, 392. Phlreomys, 24, 391-394. Val'. pallidus, Phlreomys, 394. cumingii, Cynopterus marginatus, 185. Cynopterus marginatus var., 23. Macacus cynomolgus, 389. Macacus cynomolgus var., 24. Cuti, 57. Cuyayapnit, 57. cuyonis, Tupaia, 28, 29, 100- 103, 105. Cyclorina, 230. cyclotis, Harpiocephalus, 252. Murina, 252-254. cylindrura, Urogale, 27, 106, 107. Cynamolgos, 336. palpebrosus, 338. philippinensis, 338. Cynocephalus, 59, 61, 92: distribution of, 100. volans, 23, 25, 67, 58, 92, 95, 100. Cynomolgus cagayanus, 27, 342. cynomolgus cumingii, Macacus, 339. Macacus, 338. mindanensis, 27, 340, 342. mindanensis apoensis, 27, 340. philippinensis, 338. suluensis, 27, 342. var. cumingii, Macacus, 24. Cynonycteris, 172. amplexicaudatus, 172. amplexicaudatus, Pteropus, 172. Cynopithecus othiops, 61, 336. Cynopterus, 57, 111, 114, 120, 134, 172, 177179, 181, 182, 187-190; distribution of, 183. archipelagus, 21, 182, 183, 184, 187, 188. brachyotis, 182-184. brachyotis brachyotis, 23, 24, 183-185, 187. harp ax, 184. horsfieldi, 184. jagori, 178. latidens, 189. luzoniensis, 185. marginatus, 62, 185. marginatus cumingii, 186. mm-ginatus nigrescens, 189. marginatus philippensis, 185. marginatus val'. cumingii, 23. marginatus var. philippensis, 23. melanocephalus, 62. princeps, 184. Ptenochirus jagorii, 178. sphinx luzoniensis, 185. Thoopterus nigrescens, 189.

D Daga, 58. dahlmanni, Pholidotus, 68. dailliardianus, Ussa, 51l. Dasypodidre, 67. Datal, 443. datre, Epimys, 444. Mus, 444. Rattus, 415, 441, 443-445. decumanus, Mus, 426.

INDEX Delphinus longirostrls, 65. Dendrogale, 98. dentatus, Batomys, 27, 480, 481, 483, 484. depressicornis, Anoa, 629. Dermoptera, 66, 66, 71, 72, 91. Dermopterus, 92. Desmalopex, 147, 148, 160. leucopterus, 151. Desmodondita!, 108. diadema anderseni, Hipposideros, 22, 232, 241-243, 246. griseus, Hipposideros, 25, 232, 234, 242, 243, 246. Hipposideros, 241, 242, 244. Dingalong, 58. Dinguin, 58. discolor, Emballonura, 24, 196. Distribution and faunal rehrtions, 69. Dobsonia, Ill, 116, 134, 176; distribution of, 173. exoleta, 177. palliata, 177. peronii, 173, 177. domestica, Felis, 346, Dorcatherium, 506. dore, Ombliodon, 36l. dorire, Crocidura, 91. Doryrhina, 230. Drymomys, 477. Dugian, 58. Dugiant, 58. Dugong dugon, 66. Durian, 68. Duriant, 68. Dyacopterus, 116. Dysopes caudatus, 321. cheiropus, 321. tenuis, 24.

E ecaudatus, Megrerops, 191-193. Edentata, 67. eduJis, Pteropus, 62, 163. edwardsiana, Crocidura, 77, 79, 83, 90, 91. Pachyura, 90. Paehyura caerulea, 90. effrenus, Sus, 495-497. elassodontus, Sciuropterus nigripes, 371, 384, 388, 389. elegans, Capromys, 396. Melanaxis, 613. Phlceomys, 391, 396. Elephas, 58. Eleutherura, 171. philippinensis, 23, 173. elorzanus, Us sa, 510, 612. Emballonura, 111, 114, 195. alecto, 23, 24, 200. alecto alecto, 183, 196, 198, 200. aIecto palawanensis, 21, 183, 196, 197, 198. discolor, 24, 196. monticola, 62. 196. Emballonuridre, 108-111, 194, 309; distribution of, 18B.

535

Emballonurina, 195. Emballonurinre, 109, 110, 194, 195. Eonycteris, 111, 114, 119. 126, 127, 131. 176; distribution of, 121. dobson. 127. Iongicauda, 21, 121, 127, 131, 133. major, 131. robusta, 121. 127, 128, 131, 132. robustus, 29. spelrea, 131. Eosciurus, 367. Epeira bilineata, 297. heraldica, 297. ephippium, Epimys, 462. negrinus, Mus, 26, 446. Rattus. 446. Epimys, 414, 477. albigularis, 423. basilanus, 28, 456. benguetensis. 28, 443. ca\Cis, 28, 462, 458. coloratus. 28, 434. datre, 444. ephippium, 452. everetti, 420, 421. gala, 27, 421. kelleri, 440. leucophretus, 28, 456. luteiventris, 452. luzonicus, 411. magnirostris, 430. mayonicus, 28, 457. mindanensis, 437. mindorensis, 433. negrinus, 466, 452. norvegicus, 426. ornatuJus, 28, 457. pantarensis, 453. querceti, 28, 450. rattus, 431, 477. robiginosus, 28, 435. taguJayensis. 522. todayensis, 448. tyrannus, 28. 427. vigoratus, 28. 449. vulcani, 454. vulcani 8'picis, 464. vulcani vulcani, 454. zamboangre, 442. Eptesicus, 115. Equus, 67. Erinaceidre, 71, 73. 74; distribution of, 100. Erinaceus, 99. eschscholtzi, Miniopterus, 266, 268. 272, 274. 276. Vespertilio, 272. eschscholtzii, Miniopterus, 24, 254. Vespertilio, 24. Euhys, 496. Eunycterys, 147. Euplerinre, 354. Euryalus, 211. Eusus, 495. Eut-eut, 58.

536

INDEX

everetti, Epimys, 420, 421. Mus, 420. Rattus, 25, 401, 412, 415, 417, 419--422. Tupaia, 106. Urogale, 58, 100, 106, 107. exiguus, Rhinolophus arcuatus, 213, 225. 227. exilis. Nannosciurus, 62. Sciurus. 367. Exochurus, 278. exoleta, Dobsonia. 177.

F fallax, Crunomys. 26. 377, 408-410. fslvigula henricii, Mustela, 349. familiaris, Canis, 348. sumatran us, Canis, 348. Faunal relations, distribution and, 59. Felidre. 344. 345; distribution of, 315. Felis. 57. 58, 345. catus, 345. domestica, 345, 346. javanensis, 62. javanica, 346. minuta, 25, 315, 346, 347. volans ternatea, 92. ferruginea palawanensis, Tupaia, 103. Tupaia, 62, 100. ferus, Bubalus keraban, 529. flaveolus, Nycticejus, 306. fcetida, Croeidura, 91. formosus, Myotis, 280. Vespenilio, 280. francianus, Rusa, 508, 517, 518. Ussa, 517. fraterculus, Tal'sius, 23, 28, 29, 325, 327. frenatus, Bus. 496, 497. Fructivorre, 118. fructivorus, Macroglossus, 21, 121, 125. Frugivora, 113. fulvidus, Scotophilus, 304. fulviventer, Tragulus, 62, 506. Funambulus vittatus, 62. funel'eus, Hylobates, 324. Pteropus, 153. funerus, Hylobates, 61. fur, Macacus, 338. Furipteridre, 108. fusca, Kerivoula, 258. fuscomanus, Tarsius, 326.

G Gadia, 58. Gagua, 58. Gaguang, 96. gala', Epimya, 27, 421. Rattus, 27, 415, 417, 420, 421. Galeopithecidm, 91. Galeopithecus, 91, 92, 334. marmoratus, 92. philippinensis, 23, 92, 93. rufus, 92. variegatus, 62, 92. .volans, 92. (Colugo) philippinensis, 93. Galeopteridre, 91.

Galeopterus, 91. Galeopus, 92. Galepithecus rufus, 62. garcianus, Ussa, 510. Garia, 58. geminorum, Pteropus hypomelanus, 166. gigantea, Bandicota, 62. giganteus, Sciurus, 367. Gigua, 58, 96. Glipora, 99. Glirisorex, 99. Glischropus, 112, 115, 277, 292. tylopus, 293, 294. tylopus, Vesperugo, 293. Glisorex, 99. Glisosorex, 99. Gloionyteris, 230. Glyptodontidre, 67. gonzalianus, Ussa, 511. gorrichanus, Ussa, 509, 512. grand is, Croci dura, 27, 79, 83, 84, 88. granti, Batomys, 26. grantii, Batomys, 480-483. grayi, Croeidura, 23, 79, 83, 85-87. griseus, Hipposideros diadema, 25, 232, 234. 242, 243, 246, 247. Rhinolophus. 242. grisiventre, 440. g evaranus. Ussa, 512. gll'idoteanus, Ussa, 510. Gymnura, 74, 75. Gymnuridre, 309. Gymnurinre, 73.

H halconus, Croci dura, 27, 79, 83, 85-87. Halintong, 58. hardwieki, Kerivoula, 255. hardwickii, Kerivoula, 254, 256-258, 260, 261. Vespertilio, 258. Vespertilio (Kerivoula) , 258. harpax, Cynopterus, 184. Harpiocephalus, 115, 251. cyclotis, 252. Harpyionycterinre, 109-111, 116. Harpyionycteris, 111, 114, 117; distribution of, 121. celebensis, 118. whiteheadi, 26, 118. 121. heathii, Nycticejus, 303. henricii. Martes. 349. Mustela fiavigula, 349. heraldica, Epeira. 297. herdoni, Paradoxurus, 361. Herpestes, 364. brachyurus, 365. parvus. 365. Herpestinre, 354, 364. herrei, Myotis, 22, 243, 279, 288, 289, 290. 291, 292. Hesperoptenus, 115 . Heterosciurus lateralis. 62. hipolitianus, Ussa, 512.

537

INDEX Hipposideridre, 108-110, 229, 230. Hipposideros. 67, 114. 230', 231, 241. 243, 248; distribution of, 236, 243. antricola, 24, 232, 241, 243. bieolor, 25, 231, 235-237, 239. coronata, 234. coronatus, 24, 231, 234-236. diadema, 241, 242, 244. diadema anderseni, 22, 232, 241-248, 246. diadema griseus, 25, 232, 234, 242, 243, 246. obscura, 24, 232. obscurus, 25, 231-233, 235, 240. pygm"",. 239. pygmreus, 23, 232, 233, 236, 239. wrighti, 22, 231, 236, 237, 238. Hipposiderus, 230. bicolor, 236. hirsuta, Chilophylla, 27, 243, 248, 249. hirsutus, Rbinolopbus, 212-216. Homo, 323, 324, 336. horsfieldi, Croeidura, 86. Cynopterus. 184. Hyaenidre, 344. Hyelaphus calamianensis, 519, Hydromyinre, 402, 410. Hydromys, 406. Hylidre, 61. Hylobates funereus, 324. funerus, 61. byloca!tes, Apomys, 27, 392, 460-462. Hylogaie, 99. Hylogalea, 99. Hylomys, 74, 75. Hylopetes, 384. Hyopsodonta, 72. Hypoderma, 176. Hypodermis, 107, 114, 176. peronii, 177. hypomelanus, Pteropus, 25, 158, 160, 162, 164, 166. cagayanus, Pteropus, 25, 27, 148-150, 158-163, 166. geminorum, Pteropus, 166. hypomelanus, Pteropus, 169. Pteropus hypomelanus, 159. Pteropus (Spectrum), 169. tomesi, Pteropus, 149, 160, 168, 160, 166. Hypsugo, 294. Hystricidm, 366, 489. Hystricinre, 490. Hystrix, 490. pumila, 490.

I la, 116. IIo, 58. imbricatus, Pipistrellus, 25, 295, 296, 301. Vespertilio, 300. Vesperugo, 300. imperator, Rattus, 427.

300,

inconstans. Sus, 495, 496, 601. in dicus, Sorex (Crocidura). 62. inops, Rhinolophus, 28, 213, 226, 228. inornatus, Pteromys, 63. Insectivora, 65, 66, 71-73, 91, 97; distribution of, 73. insignis, 464. Apomys, 27, 392, 461, 462, 465, 469. Apomys insignis, 460, 462, 463. bardus, Apomys, 392, 460, 462-464. insignis, Apomys, 460, 462, 463. Insu]mmns, 69, 61, 401, 469; distribution of. 473. calamianensis, 22, 470, 471, 473. Iro, 58, 348. irretitus, Pipistrellus, 24, 295-297. VespertiUo, 296. Isotus, 278. Iyam, 58.

J jacobsoni, Cbeiromeles, 320. jagori, Cynopterus, 178. Ptenochirus, 173. jagorii, Cynopterus, (Ptenochirusl, 178_ Kerivouhr, 24, 254, 255, 257, 258. Pachysoma (Ptenochirusl, 24, 178. Ptenochirus, 24, 178, 180, Vespertilio, 257. Vespertilio (Kerivoulal, 24, 257. jalaensis, Sus, 496. javanensis, Felis, 62. javanica, Felis, 346. Manis, 68, 70. Tupaya, 62. javanicus, Manis, 58. Pholidotus, 68. Tragu]us, 62, 506. jeannei. Myotis. 22. 243, 279, 284, 285. jeanni, Myotis. 292. joloensis. Sus, 497. jubatus, Acerodon, 23, 25. 135, 138. Acerodon jubatus. 23, 25. 136-140, 142_ jubatus, Acerodon, 23, 25. 136-140, 142. mindanensis, Acerodon, 136, 138. 142. 144. 146. 158. Pteropus, 23, 138, 142, 163. Pteropus (Acerodonl. 142. juvencus, Sciurus, 375, 377-379.

K Kahboog. 58. kelleri, Epimys. 440. Mus, 27, 440. Rattus, 27, 416, 440. 441. keraban ferus. Bubalus, 529. keraudrenii, Pteropus. 62. Keri vola. 115. Kerivoula. 112. 253, 266. 267, 263; distribution of, 254. fusca. 258. hardwickii. 264-258. 260, 261. hardwickii, Vespertilio, 258.

538

INDEX

Kerivoula-Continued jagorii, 24, 254, 255, 257, 258. jagorii, Vespertilio, 24, 257. pellucida, 24, 254, 255, 259, 261, 263, 264. rufopicta, 280. whiteheadi, 26, 254--257, 260, 261. Kerivoulinre, 109, 110, 112, 250, 253. Kiodotinre, 119. Kiodotus, 120. lagochilus, 122. Koh-lagh-chewey, 58. Kohs' -set, 228. Kukam, 334. Kukang, 334. Kuring, 58.

L lagochilus, Kiodotus, 122. lagochilus, Macroglossus, 122, 124--126. Macroglossus, 121, 122, 125, 126. Macroglossus lagochilus, 122, 124--126. Lamiran, 58. Lampog, 68. lanei, Philippinopterus, 22, 314, 315, 318, 323. lanensis, Pteropus, 27, 154. Pteropus vampyrus. 25, 27, 146, 147, 149, 150, 153, 154, 156. Langam, 58. lanosus, Rousettus (Stenonycteris), 172. larvata, Phyllorhina, 62. lateralis, Heterosciurus, 62. latidens, Cynopterus, 189. Leggada, 477. Lemur menagensis, 333. volans, 92. Lemuroidea, 97, 323, 324. Lenomys, 398. leocomus, Sciurus, 367. Lepedota, 65-67. leschenaulti, Rousettus, 176. Leuconoe, 277. leuconus, Sciurus, 367. leucophretus. Epimys, 28, 456. Rattus, 28, 29, , 415, 446, 456. leucopterum, Spectrum, 151. leucopterus, Desmalopex, 151. Pteropus, 23, 148, 150, 162, 153. Pteropus Spectrum, 151. Limnomys, 27, 401, 402, 484.; distribution of, 481. mearnsi, 28. 481, 485-487. picinus. 28, 481, 485, 486. sibuanus, 27, 481, 484-486. Lipotyphla. 71, 72. Lissonycteris angolensis, Rousettus, 172 . longicauda, Eonycteris, 21, 121, 127, 131, 133. longicuspis, Ussa, 509. longirostris, Delphin)lS, 66. Sus, 495. Loo Toong, 58. Loricata, 67.

Loris, 332. lucifer, Acerodon, 135, 136, 140. Pteropus, 136. Pteropus (Acerodon). 136. luctus, Rhinolophus, 62, 213. rufa. Rhinolophus, 23, 213, 223. luncefordi, Nannosciurus, 22, 369, 371, 373, 374. Lunipao, 58. luteiventris, Epimys, 452. Mus, 452. ILattus. 415. 445, 447, 452, 453, 457. luteus, Nycticejus, 306. Lutra, 362. cinerea, 363. sumatrana, 62, 353. Lutrirue, 349, 352. luzonicus, Bullimus, 411, 412, 422, 473. Epimys, 411. Mus, 26, 411. Rattus, 26. luzoniense, Pachysoma, 24, 185. luzoniensis, Crocidura, 80. Crocidura (Pachyura), 24, 80. Cynopterus, 185. Cynopterus sphinx, 185. Pachyura, 24, 25, 77-79, 80-82, 89. luzonis minimis, Cercopithecus, 326. luzonus. Chrerophon, 24, 28, 311, 313-316.

M Macaca, 336. Macacus cristatus, 338. cynomolgus, 338. cynomolgus cumingii, 339. cynomolgus var. cumingii, 24. fur, 338. nemestrinus, 62. nemestri:hus var. philipp ina, 336. palpebrosus, 338. philippinensis, 338, 340. philippinensis, Pithecus, 338. speciosus, 63, 336. macarianus, Ussa, 510. Maching, 58. macrocephalus, Catodon, 65. Macroglossa, 120. Macroglosssinre, 108, 110, 111, 116, 119. Macroglossus, 111, 114, 119, 121, 124, 126; distribution of, 121. australis, 122. fructivorus, 21, 121, 125. lagochilus, 121, 122. 125, 126. lagochilus lagochilus, 122, 124-126. minimus, 62, 122. Macronycteris, 230. Macroscelididre, 71. 97. macrotarsus, Myotis, 24, 243, 279, 281, 283, 286. Vespertilio, 24, 281. macrotis. Rhinolophus. 213, 215, 216. VespertiIio, 300. Magatilokok, 58, 326.

INDEX Magau, 58, 326. magnirostris, Epimys, 430. Mus, 27, 430. Rattus, 27, 415, 425, 428-431, 434, 435. Mago, 58, 326. Magu, 326. Maimon, 336. mainitensis, Bubalus, 526, 527. Sus, 496. Majo, 58. major, Apomys, 27, 392. 461. 464. 466. 467. Eonycteris. 131. malacensis. Viverricula. 62. 358. Malmag. 326. 330. Malmay. 58. Mammalia, 66 . Mammals, local names for, 56. Mananir. 58. Manidre. 67. 68 . Manis, 57. 59, 68. javanica. 68-70. javanicus, 58. Man turon. 58. Manx. 345. maraisianus. Ussa, 511. marchei. Mydaus. 316. 349--362. Sus, 496. marginatus curningii, Cynop'tel'us, 185. Cynopterus. 62. 185. nigrescens, Cynopterus, ' 189. philippensis. Cynopterus. 185. var. curningii, Cynopterus. 23. val'. philippensis. Cynopterus. 23 . marianna. Rusa. 525. mariannus, Cervus, 514. 522. marrnoratus. Galeopithecus. 92 . Martes, 349. henricii. 349. rnarzianus. Ussa, 611. masbatensis. Melanaxis, 513. maurus. Vesperugo. 63. mayonicus. Epimys. 28. 457. Rattus. 28. 121 415. 446, 457. mearnsi. Limnomys. 28. 481, 485--487. Pteropus. 28. 149, 150, 162. 163. Megachiroptera, 108. 111. 113. Megaderma, 112. 114, 207, 209; distribution of, 214. philippinensis, 23, 208, 209. spasrna. 68. 208. 209. spasm a spasma, 23. 208. 210. 214. spasma trifolium. 209 . Megaderrnata. 206. Megadermatidre. 206. Megadermidre, 108-110. 112. 206. Megaderminre. 206. Megrerops, 114. 116, 135. 190, 191; distribution of. 183. ecaudatuB. 191-193. wetmorei, 21. 183. 191, 193. megalodontus, Sus, 495. 496. Megalonychidre. 67. Megatheriidre. 67 .

539

Megera. 190. Melanaxis alfredi, 515. basilanensis. 518. breviceps. 513. elegans. 513. masbatensis. 513. melanius. Crunomys. 377. 408-410. rnelanocephalus. Cynopterus. 62. melanopogon. Taphozous. 62. 200 . melanotis. Sciuropterus. 62. melanura. Ceyx. 409. melanurus. Carpomys. 26. 472-475. 483. meliceps. Mydaus. 62. 360-352. Melinre. 349. menagensis. Lemur. 333. Nycticebus. 57. 332-334. Menotyphla. 61. 66. 71. 72, 97. Meyeni. Tylonycteris. 304. Vespertilio. 24. 296. 297. meyeri. Odontonycteris. 122. 124. michaelinus. Ussa, 612. Microchiroptera. 108. 109, 111. microdon. Apomys. 28. 29, 392. 461. 469. microdontus. Ussa. 611. microtis, Sus, 496. militaris. Sus. 496. Milo. 68. 348. Mimmal. 58. 96. Mimus. Pteropus. 149. 150. 163. 164. 166. 167. m\nax, Paradox urus. 356. 360. 362-364. rnindanensis. Acerodon jubatus, 136, 138. 142. 144. 146. 158. apoensis. Cynomolgus. 27. 240. apoensis. Pithecus. 27. 340. 342. Cynomolgus. 27. 340. 342. Epimys. 437. mindanensis. Pithecus, 27. 340. 342. mindanensis. Rattus. 416. 430. 437. Mus. 27. 414, 437. Pithecus. 337-343. Pithecus apoensis, 341. Pithecus miIldanensis. 27, 340. 342. Rattus, 27. 428. 430. 433, 434, 437. 439. 440. Rattus mindanensis, 415, 430. 437. Sciurus. 25. 367. 376. 377. 381. 383. Sus. 496. 501. Sus verrucosus. 501. tabla'Si, Ra~tus. 22. 415. 428. 430, 439. mindorensis. Anoa. 528. Balrenoptera. 65. Bubalus, 26. 68, 526-629. Buffelus, 528. Epimys, 433. Mus. 26. 433. Probubalus. 25, 628. Rattus. 26, 415. 428. 430, 433, 434. mindorus, Crocidura, 27, 79, 83. 87, 88, Pithecus, 28. 337, 338, 340. minim us, Cercopithecus luzon is. 326. Macroglossus. 62. 122. Miniopterre. 264. Miniopteri, 264.

540

INDEX

Miniopterinre. 109. 110, 113, 251, 264. Miniopteris. 264. Miniopteru8. 113, 115, 264. 266. 269, 278; distribution of, 264. australis, 264. 266. 268, 269. blepotis, 262, 276. eschscholtzi, 266, 268, 272, 274, 275. eschscholtzii. 24, 254. paululus, 25, 28, 254, 266, 268, 269, 271. pusillus. 24, 254. 266, 268, 271. schreibersii, 272. 8chribersii, 62. tibialis, 254, 266. 272. tristis, 24, 254, 266. 275. 277. Minneopterus. 264. minuta, Felis. 25. 315, 346, 347. minutus, Nesosus, 499. 8us. 495, 496, 499, 500. Minyopterus, 264. Miro, 58. mollendorfii, Bubalus, 526-528. Buffelus, 527. 8ciurus. 375, 377. 380. mollendorfii, Tupaia. 100, 101, 103-105. Molossi, 309. Molossidre, 108-110, 112, 323; distribution of, 315. Molossus, 320. monticola, Emballonura, Mormopterus. 314. Mundo, 58. Mungao, 58. Mungos, 59, 345, 364. brachyurus. 365, 366. palawanus, 356, 864. 365. parvus, 356, 364-366. muricula, VespeJ;'tilio, 62. Muridre, 366. 390. Murina, 113, 115, 251; distribution of, 254. cyclotis, 252-254. murina, Crocidura, 62. Murinre, 390, 400, 401. Murininre, 109. 110, 112, 250, 251. murinus, 8ciurus, 367. Mus, 401, 411. 477, 478, 480, 488; distribution of. 481. albigularis, 27, 423. castaneus, 24, 478-480. chrysocomus, 444. commissar ius, 27, 478, 479, 481. datre, 444. decumanus, 425. ephippium negrinus, 26. 446. everetti, 420. kelleri, 27. 440. luteiventris, 452. luzonicus, 26. 411. magnirostris, 27, 430. mindanensis, 27, 414, 437. mindorensis. 26. 433. musculus. 478-480. norvegicus. 425. pantarensis. 27, 453.

Mus-Continued Rattus, 411. rattus. 431. tagulayensis, 27, 422. todayensis. 21, 448. volcani. 27, 454. vulcani apicis, 27, 454. zamboangre. 27. 442. (Phlreomys) cumingi. 24. 392. Musang, 58. Musculus, 477. Apomys. 27, 392, 461. 468. 469. Mus. 478-480. Mustela, 59, 349. flavigula: henricii, 349. nudipes. 349. Mustelidre, 344. 348, 349; distribution of, 315. Mustelinae, 349. Mydau8, 58. 59, 344, 345. 349, 350. 443. marchei, 215, 349-352. meliceps, 62, 350-352. schadenbergii, 315, 349, 350, 352. Mynonycteris, 172. Myotis. 112, 115, 251, 252, 255, 277. 279. 295. 296; distribution of, 243. browni, 22, 243, 279, 288, 291, 292. formosus, 280. herrei. 22, 243. 279. 288, 289, 290, 291. 292. jeannei, 22, 243, 279, 284, 285, 292. macrotarsus, 24, 243, 279, 281, 283, 386. patricire, 22, 243, 279, 286. 288, 292. rufopictus, 24, 25, 243. 279-281, 284. Myrmecophagidre, 67. Mystacopidre, 108. mysticetus, Balrena, 65. Myzopodidre, 108.

N N adius, 184. Naidius, 114, 181. Nannomys, 477. Nannosciurinre, 367, 368. Nannosciurus, 59. 61, 368; distribution of. 371. concinnus, 368-374. exilis, 62. luncefordi, 22, 369, 371. 373. 374. samaricus. 26. 369. 371-373. surrutilus, 28, 369-373. Nannugo. 295. napu, Tragulus, 506, 507. N atalidre, 108. Natalus, 249. natunre, Pteropus vampyrus. 155. neglectus, Rattus. 435, 488. negrinus, Epimys, 446, 452. Mus ephippium, 26, 446. Rattus, 26, 415. 445-447. nemestrinus, Macacus. 62. var. philippina, Macacus. 386. nereis. Rhinolophus, 219.

541

INDEX Nesosus, 495, 499. minutus, 499. nigellus, Rusa, 28, 508, 518, 519. niger <> philippinensis, Pteropus, 62. Paradoxurus, 362. nigre.seens, Cynopterus, marginatus, 189. Cynopterus, Thoopterus, 189. Thoopterus, 189. nigrieans, Gervus, 509, 511, 522. Rusa, 508, 515, 518, 519, 522, 624, 525. Tragulus, 606. Ussa, 5ll, 622. nigricus, TraguJus, 607. nigrifrons, Paradoxura, 62. Paradoxurus, 360. nigripes, 384. elassodontus, Sciuropterus, 371, 388, 389. elassodontus, Sciurus, 384. nigripes, SciuropteruB, 371, 384, 386388. Sciuropterus, 384, 388. Seiuropterus nigripes, 338, 371, 384, 386, 387. Noetilionidm, 108, 194. noctula, Vesperugo, 63. N octulinia, 80l. noctulinus, Vespertilio, 306. norvegicus, Mus, 426. Epimys, 426. Rattus, 401, 416, 417, 425-432. notatus, Sciurus, 367. nubIan us, Ussa, 509. nudipes, Muste1a, 349. Putorius, 349. Nyctalus, 301, 303. alecto, 303. stenopteruB, 302. verrucosus, 303. Nycteridm, 108, 110, 206. Nycteris. 114. Nyeticebidm, 324, 332. Nycticebu8, 59, 832. menagensis, 67, 332, 333. philippinus. 324, 332, 333. tardigradus. 62, 333. NycticejuB, 305. alecto, 303. tla veolus, 306. heathii, 303. luteus, 306. temminckii, 306. Nycticeus alecto, Ve.spertilio, 23, 196. Nyctimene. 114, ll6. Nyctimeninm, 110. Nyctinomus, ll6, 310, 311. 314. plicatus, 62. Nyctophilax, 253. Nyctophilinm, llO. Nyctophilus. 116, Nystactes, 277.

o obscura. Hipposideros, 24. 232. Phyllorhina. 24, 232.

obseurus, Hipposideros, 25, 231-235, 240. occultidens. PachyUra, 28, 29, 77. 78. Ocypates, 251. Odontonycteris, 119, 120, 124. meyeri, 122, 124. Ombliodon dore, 361. O~m-bough' kahwee-lee, 424 . ornatulus, Epimys. 28. 457. Rattus, 28, 29, 121. 415, 446, 457. 468. Orophodontidm, 67. Orycteropodidm, 67. Oryzomys , 484. othiops. Cynopithecus, 61, 336.

P Pachyotus, 305. pachypus , Tylonycteris. 295, 304, 306. Vespertilio. 304. Vesperugo, 304. Vesperus, 304. Pachysoma, 181. amplexicaudatum, 172. brachyotis, 184. brevicaudatum, 185. luzoniense. 24. 185. titthmcheilum, 185. (Ptenochirus ) jagorii, 24, 178. Pachyura, 57-59, 73, 74, 76, 77. 82, 90. caerulea edwardsiana, 90. edwardsiana. 90. luzoniensis, 24, 25, 77-82, 89. luzoniensis, Crocidura, 80. occultidens, 28, 29, 77, 78. palawanensis, 21 , 77, 78, 79, 80. Pagil, 58. Paguma, 362. palavensis, Sus barbatus. 496, 502, 504. Palawan pangolin, 69, 7l. palawanensis, Crocidura, 21, 79, 83, 85, 88, 89. Emballonura alecto, 21, 183, 196, 197, 198. Pachyura, 21, 77, 78, 79, 80. Rattus, 22, 415. 416, 417, 418, 438. Tupaia, 97. 100, 101, 103-105. Tupaia ferruginea, 103. palawanus, Mungos, 356, 364, 365. paliata, Dobsonia, 177. paoJiatus, Pteropus, 177. pallida, Pteromys petaurista var., 62. pallidus, Phlreomys, 24, 391, 392, 394, 396. Phlreomys cumingi, 394. palmarum, Sciurus, 382. palpebrosus, Cynamolgos, 338. Macacus, 338. pangolin, Palawan, 69-71. Panicqui, Cabag vel, 141. Panique, 58. pantarensiB, Epimys, 453. Mus, 27, 463. Rattus, 27, 415, 446, 447, 453, 454, 457, 468. Pantoc, 58. Pantot, 58, 352.

542

INDEX

Panugo, 30l. papuensis, Sus, 62. Paradoxura, 345. nigrifrons, 62. Paradoxure, 362. Parad~xurus, 57, 58, 354, 360, 361, 362. crossi, 62. crossii, 360. jerdoni, 361. min ax, 356, 360, 362-364. niger, 362. nigrifrons, 360. philippinensis, 25, 356, 360-362, 364. saban us, 362. torvus, 356, 360, 361, 363. zelanicus, 362. zeylanicus, 363. Parrett, 58. parvacauda, Crocidura, 21, 79, 83, 84. par vi dens, Gheiromeles, 320. parvus, Herpestes, 365. Mungos, 356, 364-366. Pasimundot, 58. patricire, Myotis, 22, 243, 279, 286, 288, 292. paululus, Miniopterus, 25, 28, 264, 266, 268, 269, 271. pellucida, Kerivoula, 24, 254, 255, 269, 261, 263, 264. pellucidus, Vespertilio, 24, 261. Penthetor, 116. Perodicticus, 332. peroni, Cephalotes, 177. peronii, Cephalotes, 177. Dobsonia, 173, 177. Hypodermis, 177. personatus, Pteropus, 171. petaUl;ista, Pteromys, 62. var. pallida, Pteromys, 62. Petauristidre, 366, 367, 383. Petinomys, 389. petrreus, Apomys, 27, 392, 461, 466, 466. phreurus, Carpomys, 26, 472-475. pharopus, Pteropus, 303. philippensis, Cynopterus marginatuB, 23, 185 . Phyllotis, 214. Sus, 496, 497. Sus celebensis, 496. Tarsius, 23, 325-327, 331. philipp ina, Macacus nemestrinus var., 336. philippinensis, Gervus, 514 . Colugo, 93. Cynamolgos, 338. Elcutherura, 23, 173. Galeopithecus, 23, 92, 93. Galeopithecus (Colugo), 93. Ma'cacus, 338, 340. Megaderma, 23, 208, 209. Paradoxurus, 25, 356, 360-362, 364. Pithecus, 24, 25, 337-342. Pithecus MacacuB, 338. Pteropus, 173. Pteropus niger 6, 62. Rhinolophus, 23, 212-215.

philippinensis- Continued Rousettus, 172, 173. Rusa, 515. Sciurus, 24, 25, 367, 376, 377, 382. Sus, 496. Sus celebensis, 497 . TaphozouB, 23, 25, 183, 200, 202. Tarsius, 326, 334. Philippines, Cerf des, 513. Philippinopterus, 112, 115, 310, 311, 314. lanei, 22, 314, 315, 318, 323. philippinus, Cervus, 514. N'ycticebus, 324, 332, 333. Rusa, 508-510, 513-515, 518. Sus, 495 . Phlreomyinre, 390, 391. Phlreomys, 26, 58, 391, 396-398; distribution of, 392. albayensis, 62, 391, 393. cumingi, 24, 391-394. cumingi, Mus, 24, 392. cumingi var. pallidus, 394. elegans, 391, 396. pallidus, 24, 391, 392, 394. 395. schadenbergi, 398. Pholidota, 67. Pholidotus, 68. culionensis, 68. dahlmanni, 68. javanicus, 68. Phoniscus, 115. Phyllorhina, 230. antricola, 24, 241. bicolor, 236, 341. coronata, 24, 234. larvata, 62. obscura, 24, 232. pygmrea, 239. Phyllostomid"" 108. Phyllotis, 211. philippensis, 214. picinus, Limnomys, 28, 481, 485, 486. Pilandok, 58. Pilosa, 67. Pipistrellus, 112, 115, 277, 293, 294, 296, 301; distribution of, 295. abramus, 295, 296, 298, 299 . imbricatus, 25, 295, 296, 300, SOl. irretitus, 24, 295-297. pipistrellus, 297. tenuis, 295-298. pipistrellus, Pipistrellus, 297. Pithecus, 57, 58, 324, 336, 337. cagayanus, 27, 337, 342, 343. Maeacus philippinensis, 3S8. mindanensis, 337-343. mindanensis apoensis, 27, 340-342 . mindanensis mindanensis, 27, 340, 342. mindorus, 28, 337, 338, 340. philippinensis, 24, 25, 337-342. suluensis, 27, 337, 342, 343. syrichta, 339, 340. Pithex, 336.

INDEX Plecotinm, 277. Pleuropterus, 92. plicatus, Cbmropbon. 811, 314. Nyctinomus. 62. pluto. Pteropus. 168. Taphonycteris. 183, 204--206. Tapbozous. 27. 206. Podogyrnnura. 27, 58, 59, 73. 74. 75. truei. 27. 76. 76, 100. Poepbagomys aters, 62. Polatuca. 68. pollokensis. Canis, 62, 348. Potomogale, 72. Potomogalidm. 71, 73. prevosti, Sciurus. 367. Primates, 66, 66, 97, 323, 324, 336. princeps, Cynopterus, 184. Probubalus mindorensis, 25, 528. Procynonidre, 344. Proglires, 72. Protelidre, 344. Pselapbon, 147. pselaphon, Pteropus, 150. Pseudoconomys, 477. Pteinomys, 384. Ptenochirus, Ill, 114, 134, 178, 190-192; distribution of, 178. jagori, 173. jagorii, 24, 178, 180. jagorii, Cynopterus, 178. jagorii, Pachysoma, 24, 178. Pternopterus, 278. Pteromys inornatua. 68. petaurista, 62. petaurista var. pallida, 62. Pteropidre, 108, 110, 111, 113, 116, 119. Pteropinm, 108, 110. 111, 116, 134. Pteropodidre, 134, 303. Pteropodinm, 184. PteroPu8, 58, Ill, l14, 117, 134, 135, 137, 147, 148, 150, 151, 153, 176, 177; distribution of, 149. amplexicaudatus, 172. aurinuchalia. 139, 141. ba'lutus, 28, 149, 150, 168. cagayanus, 27, 159. capistratus, 171. celmno, 154. cephalotes, 303. chinensia, 23, 151-153. edulis, 62, 163. funereua, 153. hypomelanus, 25, 158, 160, 162, 164, 166. hypomelanus cagayanus, 26, 27, 148-160, 168-163, 166. hypomelanus geminorum, 166. hypomelanus hypomelanus, 169. hypomehnus tomesi, 149, 150, 158, 160, 166. hypomelas, 160. jubatus, 23, 138, 142, 153. kerll'lldrenii, 62. lanensis, 27, 154.

543

Pteropus-Continued leucopterus, 23, 148, 160, 162, 163. lucifer, 136. mearnsi. 28, 149, 160. 162, 168. mimus, 149, 160, 163, 164, 166, 167. niger 6 philippinensis, 62. paliatus, 177. personatus, 171. pharopua, 303. philippinensis, 173. pluto, 153. pselaphon, 150. pumilis, 149. pumilus, 27, 160, 163, 167, 168. pyrrhocephalus, 139, 141. speciosus, 149, 150, 162-164, 166-168. tablasi, 21, 149, 150, 169, 170, 171. temmincki, 169, 171. vampyrus, 147, 153. vampyrus lanensis, 25, 27, 146, 147, 149, 160, 163, 164, 166. vampyrus natunae, 165. vampyrus vampyrus, 104. (Acerodon) aurinucha-lis, 139. (Acerodon) jubatus, 142. (Acerodon) lucifer, 186. (Cynonycteris) amplexicaudatus, 172. (Spectrum) hypomelanus, 169. (Spectrum) leucopterus, 161. (ap.), 168. Pterygistes. l12, l16, 277, 301; distribution of, 295. stenopterus, 295, 302, 808. Ptilocercinre, 97, 98. Ptilocercus, 98. Ptychorhina, 230. pygmrea, Hipposideros, 239. Pbyllorhina, 239. pygmreus, Hipposideros, 23, 232, 283, 289. Rbinolopbus, 23, 239. Tragulus, 62, 506. pyrrhocepbalus, Pteropus, 139, 141. pumila, Acanthion, 491. Hystrix, 490. Pteropus, 149. pumilus, Pteropus, 27, 160, 163, 167, 168. Thecurus, 26, 490, 492, 493. pusi11us, Miniopterus, 24, 264, 266, 268, 271. Putorius nudipes, 349.

Q querceti, Epimys, 28, 460. Rattus, 28, 416, 445, 447, 460, 461, 466. 457, 459. Quingua, 68.

R ramosianus, Ussa, 610. Rattus, 57, 68, 401, 402, 410, 414, 416, 417, 419, 429, 430, 459, 460, 467, 469, 472, 476, 477, 480, 484, 489; distribution of, 121, 417, 441, 447. albigularis, 27, 58, 415, 417, 419, 420,428, 424.

544

INDEX

Rattus-Oontinued a'l exandrinus, 415. basilanensis, 121, 446. basilanus, 28, 415, 455. benguetensis, 28, 415, 441, 443, 445. calcis, 28, 121, 415, 446, 458, 459. callitrichus, 444. coloratus, 28, 415, 428, 430, 434-436. concolor, 454. datre, 415, 441, 443-445. ephippium, 446. everetti, 25, 401, 412, 415, 417, 419-422. gala, 27, 415, 417, 420, 421. imperator, 427. kelleri, 27, 401, 415, 440, 441. leucophretus, 28, 29, 415, 446, 456. luteiventris, 415, 445. 447. 452, 453, 457. luzonicus, 26. magnirostris, 27, 415, 425, 428-431, 434, 435. mayonicus, 28. 121, 415, 446, 457. mindanensis., 27, 428. 430, 433, 434, 437, 439, 440. mindanensis mindanensis, 415, 430, 437. mindanensis tablasi, 22, 415, 428, 430, 439. mindorensis, 26, 415, 428, 430, 433, 434. neglectus, 436, 438. negrinus, 26, 416, 445-441l norvegicus. 401, 415, 417, 4'25-432. ornatulus, 28, 29, 121, 416, 446, 457, 458. palawanensis, 22, 415, 416, 417. 418. 438. pantarensis, 27, 415, 446, 447. 463,. 454. 457, 458. querceti, 28, 415, 445, 447, 460, 451, 456, 457, 459. rattus, 90, 406, 406, 411, 415, 428-433, 440; distribution of, 428. robiginosus, 28, 29, 415, 428, 480, 435, 436. surdus, 449. tagulayensis, 27, 58, 415. 417. 420, 422. todayensis, 27, 415, 445, 447, 448, 450, 454-456. tyrannus, 28, 29, 415, 417, 425, 427, 429. vigoratus, 28, 415, 445, 447, 449, 450. vulcani, 27, 121, 415, 446, 41:)4, 455. vulcani apicis, 465. xanthurus, 422, 424. zamboangre, 21, 415, 440-442. rattus, Epimys. 431. Mus, 431. Rattus, 90, 405, 406, 411, 415, 428-431, 433, 440; distribution of, 428. Reeah, 58. Reptilia, 67. Rhesus, 336. Rhinolopha, 194. Rhinolophi, 210. Rhinolophidre, 108. 109, 112, 206. 210, 229, 230. Rhinolophides, 210. Rhinoiophina, 206, 210.

Rhinolophus, 57, 112, 114. 211, 212. 216, 229. 231. 233. 235; distribution of, 214. 225. anderseni. 213, 219. anderseni requa路lis. 25. 212. 214. 219. 220. anderseni anderseni, 212. 214. 219, 220. arcuatus, 213, 221. 224. 227-229. arcuatus arcuatus. 24, 212. 213. 224-227. arcuatus exiguus. 213, 225. 227. arcuatus typica. 224. bicoior, 236. borneensis, 217, 218. bunkeri, 22, 213, 222, 225. 228. griseus, 242. hirsutus, 212-216. inops, 28, 213, 225, 228. luctus, 62, 213. luctus rufa, 23, 213, 223. macrotis, 213. 215, 216. nereis, 219. philippinensis, 23, 212-215. pygmreus. 23, 239. rufus. 23, 24, 62, 213, 221, 223. simplex, 213, 218, 219. subrufus, 24, 212, 213, 221-223, 225, 228.' 229. virgo, 29, 212-214, 216, 218. Rhinophylla, 230. Rhinopoma:. 114. Rhinopomidre, 108. 110. Rl)inosus, 495. Rhyllorhina pygmrea, 239. Rhynchocyon, 120. RhynchomY8, 401, 4()4, 406; distribution of, 377. soricoides, 26, 377 .. 403, 405. robiginosus, EJpimys, 28, 435. Rattus. 28, 29, 415, 428, 430, 435, 486. robusta. Eonycteris, 121, 127, 128, 131, 132. robustus, Eonycteris, 29. Rodentia, 65, 66, 366. Romicia, 294. Romicius. 294. rosarianus, Ussa, 510. rosenbergi. SciuruB, 367. rostrata, Balrenoptera, 65. RouBettus, 111, 114, 119, 126, 127, 184, 171; distribution of, 173. amplexicaudatuB, 23, 118, 172. 178, 175, 176. lesehenaulti, 176. philippinensis, 172, 173. (Lissonycteris) angolensis, 172. (Stenonycteris) lanosus. 172. RoussB'. 493. roxasianus, Ussa, 511. rubiginosus, Ussa, 612. rubriventer. Sciurus, 367. rufa, Rhinolophus luctus, 23, 218, 228. rufopicta, Kerivoula, 280. rufopictus. Myotis, 24, 25, 243, 279-281, 284. Vespertilio, 24, 279. rufus, Galeopithecus, 62, 92. Rhinolophus, 23, 24, 62, 213, 221, 223.

INDEX Rusa, 69, 508. alfredi, 608, 613, 516, 518, 525. barandanus. 608, 616. basilanensis, 608, 618. ealamianensis. 608, 619-621. eulionensis. 520. francianus, 608, 617. 618. maria'Ilna. 526. nigeUus, 28, 608. 618, 519. nigricans, 608. 616, 518. 619. 522. 524, 626. philippinensis, 515. philippinus, 608-610, 613-615, 618. steeri, 622. steerii, 525.

s saban us, Paradoxurus, 362. Saccolaimus, 200. saccolaimus. Taphozous, 206. Saladung, 58. samarensis, Sciurus, 25, 376, 377. 382, 383. samaricus, Nannosciurus, 26, 369, 371-373. Sambo. 58. Sangio, 58. schadenbergi. Crateromys, 67, 392. 397-400. Phlceomys, 398. schadenbergii, Crateromys, 326 ~ Mydaus, 315, 349, 360, 352. schreibersii, Miniopterus, 272. schribersii. Miniopterus, 62. Sciuridae, 366-368. Sciurinre, 367. 368, 375. Scinropterus. 58. 69, 383, 384, 389; distribu路 tion of. 371. crinitus. 29, 371, 384. 389. 390. melanotis, 62. nigripes, 384. 388, 389. nigripes elassodontus. 371, 388, 389. nigripes nigripes, 371. 384. 386-388. Sciurus. 67, 59, 99, 375; distribution of, 377. albicauda, 375, 377, 379. bivittatus, 382. cagsi, 367, 381. concinnus. 369. exilis. 367. giganteus. 367. juvencus. 375, 377-379. leucomus, 367. mindanensis, 25, 367, 376, 377, 381, 383. mollendorffi, 875. 377, 380. murinus. 367. nigripes elassodontus, 384. notatus, 367. palmarum, 382. philippinensis, 24, 25. 367, 376. 377, 382. prevosti, 367. rosenbergi, 367. rubriventer, 367. samarensis, 25. 376, 377, 382, 383. steerei, 25, 379. steeri, 376. 259453--36

545

Sciurus-Continued steerii, 367. 376, 377. 378, 381, 382. tenuis, 367. Tupaia, 58. weberi, 367. Scotophila, 306. Scotophilu8, 112. 116, 277. 295, 305; distri路 bution of, 295. fulvidus. 304. temminckii. 295, 306, 308. Selysius, 278. Semnopithecus albipes, 61 , 336. Senonycteris, 172. Sericonycteris, 14 7. sibuanus. Limnomys, 27 . 481, 484- 486. Sideroderma, 230. Sikelaphus soloensis, 613. silaceus. Celaenomys, 25 , 377, 402, 403. Xeromys, 402. SileD'us, 336. Simia syrichta, 23. simplex. Rhinolophus , 213, 216. 21 8. 219. Singaiong, 58, 348. Solenodontidre, 71. 73. soloensis, Sikelaphus, 513. Sorex. 89, 99 . (Crocidura) indicus. 62. Sorex-glis. 99. Sor icidre, 71, 73, 74, 76; distribution of, 79. soricoides, Rhynchomys . 26. 377, 403 , 405. spasms:, 207 . Megaderma, 68, 208, 209. Megaderma spasma, 23. 208, 210 , 2i4. spasma, Megaderma, 23 , 208, 210, 214. trifolium, Megaderma. 209. Vespertilio, 208. spatharius, Ussa. 509. Species erroneously' credited to the Philippines. 61. Specimens, collection and preservation of, 52. speciosus. Macacus, 62, 336. Pteropus, 149. 160, 162- 164, 166- 168. Spectrum, 147. hypomelanus. Pteropus, 159. leucopterum, 151. leucopterus, Pteropus, 15l. Tarsius, 62, 326, 327, 334, 335. spelrea. Eonyc1leris, 131. Speorifera, 230. sphinx luzoniensis, Cynopterus, 185. Squamata, 67. stanleyanus. Tragulus. 507. steerei, Cervus, 522. 524. Sciurus, 25. 379. steeri, Sciurus. 376. Rusa. 522. steerii, Cervus, 518. Rusa, 525. Sciurus. 367, 375. 377, 378, 381, 382. Steno chinensis, 65.

546

INDEX

Stenonycteris lanosus, Rousettus, 172. stenopterus, Nyctalus, 302. Pterygistes, 295. 302. 303. Vesperugo, 302. Stylinodontidre, 67. Styloctenium, 114, 116. subrufus, Rhinolophus. 24. 212, 213, 221-223, 225 , 228. 229. Suidre. 493, 494. Suin re, 494. suluensis, Cynomolgus, 27. 342. Pithecus, 27, 337, 342, 343. sumatrana, Lutra, 62, 353. sumatran us, Canis familiaris, 348. surdus, Rattus, 449. surrutilus, Nannosciurus, 28, 369-373. Sus, 57, 58, (9(. 500. aha!nobarbus, 496, 502-504. arietinus, 4,95, 496. barbatus, 495, 502-504. bar-batus ahamobarbus, 502. barbatus balabacensis, 496, 504. baTbatus palavensis, 496, 502, 504. barbatus var. calamianensis, 498. basilanensis, 496. calamianensis, 495, 496, 498, 502-504. cebifrons, 495, 496, 500. celebensis, 497. celebensis philippensis. 496. celebensis philippinensis, 497. conchyvorous, 496. conchyvorus, 497. effrenus, 495-497. frenatus, 496, 497. inconstans, 495, 496, 50l. jalaensis, 496. joloensis, 497. longirostris, 495. . mainitensis, 496. marchei, 496. megalodontus, 495, 496. microtis, 496. militaris, 496. mindanensis, 496, 50l. minutus, 495, 496, 498. 500. papuensis, 62. philippensis, 496, 497. philippinensis, 496. philippinus, 495. verrucosus mindanensis, 50l. vittatus. 62. (aper montanus). 57. swinhoei, Balrenoptera, 65. Syconycteris, 125. 126. Syncerus, 526 . Syndesmotis, 230. Synetherinre, 490. syrichta, Pithecus, S39, 340. simia. 23.

T tablasi, Pteropus, 21, 149, 150, 169, 170, 171. Rattus mindanensis. 22, 415, 428. 430, 439. Treniodonta, 67. Tagbanuas, 58. Tagnan, 58. tagulayensis, Epimys, 422. Mus, 27, 422. Rattus. 27, 58. 415, 417, 420, 422. Tahdeeahn, 58. Talibata. 58. Talibatab, 58. Tali路 rah bali booey, 76. Talonon, 58. Talpidre, 71, 73. Tamaral, 348. Tamaran, 58 . Tamarao. 58. Tana, 98. tangalunga, Viverra. 58. 356-359. Taphonycteris, 111, 114, 195, 204. capito, 28, 29, 183, 204, 205. pluto, 183, 204-206. Taphozous, Ill, 114, 195, 200, 204. melanopogon. 62, 200. philippinensis, 23, 25, 183. 200, 202. pluto, 27, 205. saccolaimus, 205 . 'l7apilac, 58. tardigradus, Nycticebus, 62, 333. Tarra oahbooey, 58. Tarsiidre. 324, 325. Tarsius, 22, 57-59, 61, 66, 325, 340. carbonarius, 23, 325, 327, 331. fraterculus, 23, 28, 29, 325, 327, 329. fuscomanus, 326 . philippensis, 23, 325-327. 331. philippinensis, 326, 334. spectruTn, 62, 326, 327, 334, 335. Tarsomys, 27, 401. 402, 487; distribution of, 481. apoensis, 27, 481, 488. 489.. telesforianus, Ussa, 512. temTnincki, Pteropus. 150, 169. 17l. temminckii, Nycticejus, 306. Scotophilus, 295, 306, 308. Vespertilio, 303, 306. teT)uis, Dysopes, 24. Pipistrellus, 295-298. Sciurus, 367. Vesperugo, 297. ternatea, Felis volans, 92. Thecurus, 57-59, 490. pUTnilus, 25, 490, 492, 493. Thoopterus, Ill, 114. 116, 135, 188. nigrescens, 189. nigrescenS, Cynopterus, 189.

547

INDEX Thyreorhina. 230. Thyroptera. 249, 293. Thyropteridre, 108. tibialis. Miniopterus. 254. 266. 272. Tinucay, 58. titthrecheilum, Pachysoma, 185. todayensis. Epimys, 448. Mus, 27, 448. Rattus. 27. 415. 445, 447. 448, 450. 454456 . tomesi, Pteropus hypomelanus, 149. 150. 158, 160. 166. Tomopeatinre, 250. Tonger vergonzoso, 335. torquatus, Cheiromeles. 315, 320- 323. Chiromeles, 32l. Chiropeles. 321. torvus. Paradoxurus, 356. 360. 361. 363. TraguJidre, 494, 506. Tragulus, 58, 59, 506., fulviventer, 62. 506. javanicus. 62, 506. napu, 506, 50'7. nigricans. 506. nigricus, 507. pygmreus, 62, 506. stanleyanus, 507 . trifolium. Megaderma spasma, 209. Trilatitus. 278. tristis. Miniopterus, 24, 254, Vespertilio, 24, 275. . truei. Podogymnura. 27. 75 , Tryphomus adustus. 27. Tryphomys. 27, 401, 476; distribution oI. 473. adustus. 473, 476, 477. tuasoninus. Ussa. 509. Tubulidentata. 67. Tupai, 99. Tupaia. 59. 97- 99. 101, 379; distribution of. 100. cuyonis, 28. 29. 100-103. 105. everetti. 106. ferruginea', 62, 100. ferruginea palawanensis. 103. javanica, 62. miillendorfii, 100, 101, 103-105. palawanensis, 97, 100, 101, 103-105. Sciurus. 58. Tupaiidre. 71, 97; distribution of, 98. Tupaiime. 97. 98. Tylonycteris, 112. 115. 277, 293, 303; distribution of. 296. meyeni, 304. pachypus, 295. 304. 305. tylopus. G1ischropus, 293, 294. Vesperugo, 293. Vesperugo (Glischropusl. 293. typica, Rhinolophus areuatus. 224.

tyrannus, Epimys, 28, 427. Rattus. 28. 29, 416, 417, 425, 427, 429.

U Umhao kahwee-Iee. 58. ombooisay, 58. takobo. 68. talikoobung, 58. Ungulata, 493. Urogale, 26, 58, 59. 97, 98, 106. cylindrura, 27, 106. 107. everetti, 58, 100. 106. 107. U rsidre, 344. Usa. 58. Ussa l1Imbrosianus, 510. atheneensiB. 512. barandanus, 516. baryceros, 513. brachyceros, 609 . chrysotrichos. 512. cinereus. 513. corteanus. 512. crassicornis, 513. dailliardianus, 511. elorzanus, 510, 612. francianus. 517. garcianus. 510. gon zalinus, 61l. gorrichanus, 509, 512. guevaranus. 512. guidoteanus. 510. hipolitianus, 512. longicuspis, 609. macarianus, 510. maraisianus, 51t. marzianus, 51l. michaelinus, 512. microdontus, 511. nigricans, 511 , 522. nublanus. 509. ramosianus. 510. rosarianus, 510. roxasian us, 511. rubiginosus, 512. spatharius, 509. telesforianus, 512. tuasoninus, 509. verzosanus, 510. 512. vidalious. 513 . villcmerial1us, 513.

V vampyrus, Pteropus, 147, 160, 163. lanensis, Pteropus. 25, 27. 146, 147. 149. 150, 153, 154, 156. natunre. Pteropus. 155. Pteropus vampyrus, 154. vampyrus, Pteropus, 154. Vespertilio, 153. variegatus, Galeopithecus, 62. 92.

548

INDEX

verrucosus mindanensis, SUB, 50l. Nyctalus, 303. Verrusus, 495. verzosanus, Ussa, 510. 612. Vespertilio, 115, 278, 303. abram us, 298. admirabilis. 92. belangeri, 303, 806. capaccini, 62. chinensis. 63, eschscholtzi, 272. eschscholtzii. 24. formosus. 280, hardwickH, 258. imbricatus, 300. irretitus. 296. jagorii, 257. macrotarsus. 24. 281. macrotis. 300. meyeni, 24, 296, 297. muricula, 62. noctulinus. 306. pacbypus, 304. pellucidus, 24, 261. rufopictus, 24, 279. spasma, 208. temminckii. 303, 306. tristis, ' 24, 275. vampyrus. 153. (Kerivoula) haTdwickii, 25'8. (Kerivoula) jagorii, 24, 257. (Nycticeus) alecto, 23, 196. Vespertiliones, 194. 277. Vespertilionidre. 108-110, 112, 194. 206, 249, 250, 264, 303, 309. Vespertilioninre, 109, 110, 112, 251, 253. 255, 277. Vesperugo, 292, 294, 301, 303. abram us, 298. Glischropus tylopus, 293. imbricatu6, 300. maurus, 63. noctula, 63. pachypus, 304. stenopteru8, 302. tenuis, 297. tylopus, 293. Vesper us. 303. paehypus, 304. vidalinus, Ussa, 513. vigoratus, Epimys, 28, 449. Rattus, 28, 415, 445, 447, 449. 450. ViHe, 58.

villemerianus, Ussa., 513. virgo. Rhinolophuf>, 29, 212-214, 216. 218. Visin, 58. vittatus, Funambulus, 62. Sus. 62. Viveria binturong. 355. Viverra, 58, 345, 354, 357. tangalunga, 58, 356-359. â&#x20AC;˘ zibetha, 62, 358. Viverricula mala:censis, 62, 358. Viverridre. 344. 353; distribution of. 356. Viverrinre. 354. volans camellii, Cato-Simius, 92. Cynocephalus, 23, 25, 57, 58, 92, 95, 100. Galeopitheeus, 92. 95. Lemur, 92. ternatea. Felis, 92. vulcani apicis. Epimys, 454. apieis. Mus, 27. 454. apicis, Rattus, 455. Epimys. 454. Epimys vulcani, 454. Mus. 27, 45Âˇ1Rattus, 27, 121. 415, 446, 454, 455. vulcani Epimys. 454.

w weberi, Sciurus, 367. wetmorei, Megrerops, 21. 183, 191, 193. whiteheadi, Chrotomys. 25. 377. 406-408. Harpyionyeteris, 26, 118. 12l. Kerivoula. 26, 254-257, 260. 261. whitei. Aretietis. 355-357. wrighti. Hipposideros. 22, 231. 235, 237.

x Xa'lltharpia amplexicaudatus. 172. Xantharpyia. 171. Xanthurus, Rattus. 422. 424. Xenarthara. 67. Xeromys. 402. 406. silaceus, 402.

y yut-yut. 395.

z zamboangre. Epimys. 442. Mus. 27. 442. Rattus. 27. 415. 440-442. zelanicus. Paradoxurus, 362. zeylanieus, Paradoxurus. 863. zibetha', Viverra. 62. 358.

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