Eiszeitalter
und
Gegenwart
Band 1 4
Seite
96-103
Öhringen/Württ.,
1. September
1963
N o t e s on s o m e P l e i s t o c e n e mammal migrations from the Palaearctic t o the N e a r c t i c V o n BJÖRN K U R T E N , Helsingfors
Mit 2 Abbildungen im Text S u m m a r y . The following dates for mammalian migrations from the Palaearctic to the Nearctic are suggested: Smilodontine sabre-tooths, Elster (Kansan); black bears, Elster (Kansan); brown and grizzly bears, Würm (Wisconsin); wolverine, Saale (Illinoian). Correlation between the mammalian faunas in the Nearctic and the Palaearctic should be based on compilation of many additional migration items. Z u s a m m e n f a s s u n g . Die Einwanderungen einiger paläarktischer Säugetiere ins Nearktikum werden folgendermaßen datiert: Smilodontine Säbelzahnkatzen Elster (Kansan). Schwarz bären Elster (Kansan). Braunbären, bzw. Grizzlybären Würm (Wisconsin). Vielfraß Saale (Illi noian). Für eine befriedigende Korrelation zwischen den eiszeitlichen Säugetierfaunen des nearktischen und paläarktischen Raumes müßten noch eine Reihe von Einwanderungsbeispie'.en analysiert werden. T h e correlation between the Pleistocene mammalian faunas of North A m e r i c a and E u r o p e is at present a topic of much informal discussion a n d controversy, but on which relatively little has been published. T h e knowledge of Pleistocene faunal evolution in both areas is still incomplete, more so in the N e a r c t i c , but a fairly coherent succession has recently been worked out b y H I B B A R D a n d his associates ( H I B B A R D , 1958; T A Y L O R & H I B B A R D , 1959) and related to the standard glacial-interglacial sequence. Perhaps it may still be said that knowledge is too incomplete to permit a stage-by-stage correlation on faunal evidence. Nevertheless the topic is of such interest that contributions to it should be welcomed. It appears to the present writer that the most hopeful method to attack the problem is the detailed study of case histories of intercontinental migration b y means of taxonomic and evolutionary analysis. H e n c e I have chosen to offer some notes on a number of C a r n i v o r a , summarized from a comparative study in progress. T h e y are intended to supply some preliminary correlation items of the desired kind. It m a y per haps be hoped that specialists on this and other groups w o u l d analyse other instances of intercontinental migration in the Pleistocene, of which m a n y w o u l d be available. It may be assumed that migration between the O l d and the N e w W o r l d occurred exclusively across the Bering Strait, as far as the m a m m a l s a r e concerned, a n d that it took place only when there w a s a land bridge; that is to s a y , during a glacial phase. Immigrants are therefore assumed to have migrated in the preceding cold phase, if their first occurrence is in an interglacial fauna. First occurrence in a cold fauna is taken to signify an immigration during the same cold phase. In either case, however, it is a prere quisite that a probable ancestor should be k n o w n to have existed, at the given time, in the area of assumed origin. Errors are bound to arise sometimes because our knowledge is incomplete a n d later discoveries m a y reveal that the immigrant w a s actually present a t an earlier d a t e . A more detailed analysis of the evolving populations m a y help us to a v o i d errors of this kind. If it can be shown that the assumed ancestor a n d the immigrant form an essential mor phological continuum, this m a y be taken as additional indication that the correct time of migration has been found. A morphological g a p between the two will suggest that part of the evolutionary sequence is missing and may have taken p l a c e in either a r e a . I have endeavoured to p a y full attention to this factor in the examples to follow.
Pleistocene mammal magrations from the Palaearctic to the Nearctic A.
Sabre-tooths
of
the
genera
Megantereon
T h e close relationship between the European Megantereon
and
97
Smilodon
and the American
Smilodon
was recognized by S C H A U B ( 1 9 2 5 ) . Unfortunately, however, M A T T H E W ( 1 9 2 9 ) made
the
error of synonymizing the European Homotherium with Smilodon. With the discovery of excellent material of the sabre-tooth Dinobastis in the late Pleistocene of T e x a s ( M E A D E , 1 9 6 1 ) it has become clear that this form, not Smilodon, is the American ally of the European Homotherium, whereas Megantereon is allied to Smilodon. T h e y form two quite distinct groups of sabre-toothed cats, which m a y be termed the tribes Homotheriini and Smilodontini. SCHAUB ( 1 9 2 5 ) pointed to the detailed similarity in the postcranial skeletons of Megantereon and Smilodon. T h e neck, for instance, is much elongated, and the distal segments of the heavy limbs are shortened. In the Homotheriini the neck is less elongated a n d the limbs are not shortened distally, indeed the forearm is extremely long. T h e dentitions and skulls (fig. 1 ) are also quite distinctly constructed in the two groups. T h e Smilodontini have dirk-like, very long upper canines, which were evidently used for stabbing exclusively; they are relatively and absolutely larger in Smilodon, but this is only a specialized character and does not obscure the essential similarity. T h e lower canines were reduced and form flanking elements in the transverse incisor row. T h e cheek teeth are not much modified from the normal feline type, except for a progressive reduction of the anterior elements and of the protocone (internal cusp) in the upper carnassial. T h e skull profile tends to be triangular, with a high occiput, only slightly overhanging the occipital condyles. T h e post-orbital processes a r e well set off with a marked con足 striction behind, separating them from the small but globular braincase. T h e development of the mastoid processes is correlated with that o f the head depressors and is more pronounced in the a d v a n c e d Smilodon with its enormous sabres. In contrast, the Homotheriini have relatively short, flattened sabres with crenulated, sharp cutting edges all along the front and back; w e a r facets show that they were used in biting, as well as stabbing and slashing. T h e incisors form a semicircle, unlike the
Fig. 1. Skulls of Homotheriini (left) and Smilodontini (right) in side view. Upper left, Homotherium crenatidens, Perrier, Villafranchian. Lower left, Dinobastis serus, Friesenhahn (Texas), Wisconsin. Upper right, Smilodon neogaeus, Arroyo Pergamino (Argentina), Pampean. Lower right, Megan足 tereon megantereon, Seneze, Villafranchian. Not to scale. 7
Eiszeit und Gegenwart
98
Björn Kurten
smilodont transverse row. T h e cheek teeth are highly modified, extremely thin slicing blades. T h e skull is long and low with an arched profile, the frontal region is broad without distinctly set off processes, and the braincase is separated from the occipital plane by a m a r k e d constriction. T h e two tribes probably arose independently from orthodox feline cats by divergent evolution along quite different a d a p t i v e paths. T h e first stage in the development of sabre-like upper canines is seen in the present-day Felis nebulosa. T h e idea of an iterated evolution of sabre-toothed cats gains in credibility from the fact that independent evolu tion of sabre-toothed carnivores has been demonstrated in the M a r s u p i a l i a and C r e o d o n t a . T h e history of both groups is mainly or entirely contained in the Pleistocene (inclu ding the Villafranchian), and most or all of the Eurasian a n d American Pleistocene sabre-tooths may be referred to one group or the other. T h e two main types of homotheres, Homotherium and Dinobastis, occur in both hemispheres. T h e former are only early Pleistocene (Villafranchian) in Europe, but may occur later in America (Ischyrosmilusf). T h e latter are middle and late Pleistocene (Dinobastis serus, N o r t h A m e r i c a ; Dinobastis latidens, E u r o p e ; Dinobastis ultimus, China). T h e smilodont cats show a more definite evolutionary trend and indicate a n inter continental migration at a rather narrowly defined point in time. All the Eurasian forms are referred to the genus Megantereon. T h e earliest forms, a p a r t from some possible Indian ancestors in the Pliocene, occur in the earliest Villa franchian in E u r o p e (Villafranca d'Asti, Etouaires). They are relatively small, of perhaps p u m a size. T h e r e is evidence of a gradual size increase, and the late Villafranchian forms (Seneze, V a l d'Arno, N i h o w a n ) are somewhat larger. All of the E u r o p e a n Villafranchian
Fig. 2. Evolution of lower jaw and teeth in the Smilodontini. Bottom, Megantereon megantereon, Seneze, Villafranchian, with P 3 and large jaw flange. Centre, Smilodon gracilis, Port Kennedy Cave, Yarmouth, retaining Pg (alveolus), flange somewhat reduced. Top, Smilodon neogaeus, Lapa Escrivania, Brazil, late Pampean, size greatly increased, Ps lost, flange reduced. All to the same scale.
Pleistocene mammal magrations from the Palaearctic to the Nearctic
99
populations may, however, be referred to a single evolving species, M. megantereon. A t the close of the Villafranchian, the line became extinct in Europe, but in Asia (and Africa) it survived into the middle Pleistocene. T h e last representative of this line in Asia, known at present, is Megantereon inexpectatus from Locality 1 (the Peking M a n site) of C h o u koutien. T h e date of this form is late Elster or early H o x n i a n . M. inexpectatus is larger than M. megantereon, showing that the phyletic growth continued, a n d in fact it is about the same size as the earliest American Smilodons. O u r next glimpse of this evolutionary line comes from the N e w W o r l d with the ear足 liest known members of the genus Smilodon. T h e y come from Port K e n n e d y C a v e out of deposits that appear to be Y a r m o u t h in age ( H I B B A R D , 1 9 5 8 ) . As fig. 2 shows, these forms resemble advanced Megantereon more than advanced Smilodon in m a n y characters. T h e y still retain a well developed Ps (alveolus in the figured specimen) a n d have a distinct inner cusp (protocone) in the upper carnassial. On the other hand, the reduction of the dependent flange on the lower jaw has already got under w a y in the early Smilodon. T h e trend in this character, within Smilodon, shows well enough that the genus evolved from a large-flanged ancestor like Megantereon. A p p a r e n t l y the sabre still bit inside the lower lip in Megantereon, a n d h a d to have a sheath supported by the j a w b o n e . In Smilodon the sabre bit outside of the lower lip, a n d the sheath could be reduced. If this character is m a d e the key character of the two genera, the Port K e n n e d y C a v e form should go into Smilodon in spite of numerous resemblances to Megantereon. T h e development of the j a w flange in the Choukoutien form is unfortunately unknown ( T E I L H A R D , 1 9 4 5 ) . In Smilodon of Illinoian age, from the C o n a r d Fissure (BROWN, 1 9 0 8 ) , further p r o 足 gress in size and dental characters have occurred, but not until S a n g a m o n and Wisconsin time the full-fledged Smilodon known from Rancho L a Brea and other asphalt deposits is met with. It appears highly probable that the migration occurred at a point in time roughly coinciding with the age of the last known Megantereon. Thus the migration would be likely to be of Elster date. In this w a y it m a y be concluded that the characteristic Elster faunas of the O l d W o r l d antedate the Yarmouth faunas in N o r t h A m e r i c a , and that the Y a r m o u t h is a correlative of the E u r o p e a n Holstein. Smilodon also entered South America, where the earliest form, a relatively small a n d primitive one, occurs in the C h a p a d m a l a l ( K R A G L I E V I C H , 1 9 4 8 ) . T h i s appears to give a m a x i m u m age for the C h a p a d m a l a l : it can hardly antedate the Y a r m o u t h and Holstein. T h e outlines of the evolution and migration of the Smilodontini are indicated in the d i a g r a m (table 1 ) . B.
Bears
of
the
genus
Urs
us
T h e bear family, U r s i d a e , is one of the smallest a n d most tecent carnivore families. T h r e e subfamilies are recognised at present ( T H E N I U S , 1 9 5 8 , 1 9 5 9 ) , but one, the A g r i o theriinae, became extinct at the close of the Pliocene, a n d does not concern us here. A n 足 other, the Tremarctinae, is exclusively American in distribution, a n d evolved from immi足 grants dating back to the Pliocene (Plionarctos). T h e presence of large tremarctine bears (Arctodus) may h a v e been a factor in the relatively late spread of Ursus into the N e w World. T h e third subfamily, the Ursinae, has a H o l a r c t i c and Indian distribution. With a single exception (Ursus americanus) all of its species are present in the O l d World, a n d the fossil record shows that it originated in Eurasia. In the Nearctic, ursine bears are not known until the middle Pleistocene, with a single uncertain exception (JOHNSTON & S A V A G E , 1 9 5 5 ) , a specimen from the Blancan Cita C a n y o n ; it is fragmentary and m a y , or m a y not, be ursine. 7 *
Björn Kurten
100
Table 1 Evolution and migration of the Smilodontini
Smilodon neogaeus
Wisconsin
Smilodon " californicus" (Rancho La Brea etc.)
Würm
Sangamon
S. trinitiensis Texas etc.
Eem
Illinoian
S. troglodytes Conard Fissure
Riß-Saale
Yarmouth
S. gracilis Port Kennedy Cave
Holstein
Smilodon ensenadensis S. riggii Chapadmalal
Eurasia
North America
South America
t
t 1
Megantereon inexpectatus Choukoutien
MindelElster
Megantereon
Cromer & Villa franchian Table 2 Intercontinental migrations of Ursus,
excluding U.
North America
M.
spp.
megantereon
maritimus Eurasia
Recent
U. americanus
U. arctos
Wisconsin
U. americanus Many Iocs.
U. arctos <~ Alaska
Recent
U. arctos
U.
thibetanus
U. arctos Many Iocs.
U. thibetanus China
Eem
U. arctos Many Iocs.
U. thibetanus China
Riß-Saale
U. arctos U. thibetanus Tornewton Cave Europe, China etc.
Holstein
U. arctos Grays etc.
MindelElster
U. arctos China
Würm Sangamon
Illinoian
Yarmouth
U. americanus Trinity River U. americanus Cumberland Cave, Conard Fissure U. americanus Port Kennedy Cave t
1 1
'
U. thibetanus China
U. thibetanus China Cromer Villa franchian
U. thibetanus China, Europe U. etruscus Europe
U. ^thibetanus China
Pleistocene mammal magrations from the Palaearctic to the Nearctic
101
T h e first certain ursine species to a p p e a r in N o r t h America is Ursus americanus, the black bear. T h e earliest record appears to be Port K e n n e d y C a v e , as in the case of the smilodonts, and the bears evidently date from the Y a r m o u t h . These early black bears are of moderate size and h a v e several primitive characters, e.g. the l a r g e size of the upper carnassial and the small size of the last molar. In these characters, a n d also in morphology, the P o r t Kennedy C a v e black bear closely resembles the middle Pleistocene Ursus thibe tanus kokeni (Asiatic black bear) known from Chinese deposits, e. g. Choukoutien. It seems probable that the American form descended directly from the closely allied Asiatic species, and that the migration occurred at the same time as that of the Smilodontini, i. e. during the Elster Glaciation. Illinoian black bears in N o r t h America, from C u m b e r l a n d C a v e a n d C o n a r d Fissure ( G I D L E Y & GAZIN, 1 9 3 8 ) , show definite advance over the stage seen a t Port Kennedy. L a t e Pleistocene forms reached great size; in the Postglacial, the size was secondarily reduced ( K U R T E N , in the press). T w o other species of Ursus are at present found in N o r t h America, the polar bears (Ursus maritimus) and the brown and grizzly bears (Ursus arctos). B o t h are conspecific with O l d World species, and the indication is that their migration is o f more recent date. Unfortunately the fossil history of the polar bear is practically unknown. As regards Ursus arctos, however, the fossil record bears out the contention. M o s t or all fossils of this species from N o r t h America appear to date from Postglacial times or possibly the latest Wisconsin ( K U R T E N , 1 9 6 0 ) . It is possible that the species was present in Alaska a t a somewhat earlier date, well back in the Wisconsin. H o w e v e r this m a y be, the intercon tinental migration evidently took place during the W ü r m = Wisconsin. As a matter of fact, the morphology of the A l a s k a n and Eastern Siberian Ursus arctos indicates t w o independent migrations: a northern, with narrow-skulled animals from northern Siberia, a n d a southern, with broad-skulled animals from the K a m c h a t k a population. C.
The
glutton,
Gulo
gulo
T h e glutton or wolverine evolved in the O l d W o r l d . T h e ancestral form is the Cromerian Gulo schlössen, known from various localities in Central E u r o p e , and chiefly distinguished by its much smaller size. T h e first true Gulo gulo appear in the Elster ( M o s bach; see T O B I E N , 1 9 5 7 ) of Europe, and also in China (Choukoutien; see P E I , 1 9 3 4 ) . In N o r t h America, the earliest record of wolverine appears to come from Cumberland C a v e ( G I D L E Y & G A Z I N , 1 9 3 8 ) , where it is part of the cold elements that appear to date from the Illinoian ( H I B B A R D , 1 9 5 8 ) . T h e Illinoian is thus the minimum age of the m i g r a tion, but there is also a possibility that the migration dates back to the Elster. It m a y be noted, however, that the N e w World population is only very moderately differentiated from that in the O l d ( K U R T E N & R A U S C H , 1 9 5 9 ) . a n d only ranks as a distinct subspecies. T h i s would support a relativelv late date, and a t present a Saale-Illinoian migration appears the most likely alternative. Conclusions T h e three migration items discussed here support the correlation of glaciations a n d interglacials in N o r t h America and E u r o p e as far back as the Holstein in Europe a n d Y a r m o u t h in N o r t h America, as shown in tables 1 a n d 2 . T h e e x a c t European corre latives of the K a n s a n , the Aftonian, and the N e b r a s k a n faunas remain doubtful. Perhaps the definitive clearing up of the migration of the elephants will settle p a r t of the problem. T h e earliest true elephants in N o r t h America appear in faunas assigned to the K a n s a n , such as the H o l l o m a n and C u d a h v ( T A Y L O R & H I B B A R D . 1 9 5 9 ) . This early form is Elephas haroldcooki. Detailed analysis of a large material will be necessary to show whether
Björn Kurten
102
this species m a y be derived from the primitive mammoths of the Elster, or from advanced Elephas tneridionalis types of the late Villafranchian. A correlation of the K a n s a n with the Elster would make the Aftonian a correlative of the Cromerian in Europe. T h e latter is a complex stage with two distinct temperate oscillations ( W E S T , 1 9 6 1 ) . On the other hand, m a n y American specialists seem to favour a correlation of the Aftonian with the Villafranchian in E u r o p e . Some implications of such a correlation are somewhat disturbing. Correlation of the Aftonian or p a r t of it with the Villafranchian, and of the Y a r m o u t h or part of it with the Holstein, would suggest one of the schemes ( 1 ) — ( 3 ) below, in contrast with the usual arrangement ( 4 ) . (1)
1 \ )
Kansan Aftonian Nebraskan Rexroadian
Holstein (Elster) Cromer Günz Villafranchian Donau Astian
(2) Yarmouth
Holstein
Yarmouth
Kansan Aftonian Nebraskan Rexroadian
1 > J
Elster (Cromer) Günz Villafranchian Donau Astian
(3) Yarmouth Kansan
Nebraskan Rexroadian
Holstein Elster Cromer (Günz) Villafranchian Donau Astian
(4) Yarmouth Kansan Aftonian Nebraskan Rexroadian
Holstein Elster Cromer Günz Villafranchian
Aftonian
1 \ )
It would obviously be premature to attempt an evaluation of these and other possible combinations at present, when even the intra-continental correlations (e. g., the dating of the American faunal sequence in terms of the American glacial-interglacial sequence) are somewhat uncertain. One possible source of bias in intercontinental correlation should, however, be mentioned. Strictly homotaxial relations for migrating groups can only be expected at the time of the actual migration, i. e., at the time of a glaciation, as a rule. A p p a r e n t homotaxial relations between interglacial faunas m a y be deceptive. F o r a hypo thetical example, suppose that a typical Holsteinian Old W o r l d form migrates to N o r t h America in the Saale (Illinoian), and becomes extinct in E u r a s i a , or perhaps evolves into a more advanced Eemian type. If it is further supposed that the American immigrant happens to be known to us only in strata of Sangamon age, this will be a case of closer homotaxial relationships between the S a n g a m o n and the Holstein, than between the S a n g a m o n and the Eem. T h i s may possibly h a v e some bearing on the apparent relation ship between the Aftonian a n d the Villafranchian. N a t u r a l l y , migrants in the other direction, from the N e w W o r l d to the O l d , will have the opposite effect; but as these migrations have been much less common, their effect is likely to be overshadowed. Numerous migration items await detailed analysis. L y n x e s among felids; wolf and red fox among canids; least weasel, otter and marten among mustelids are obvious cases in the C a r n i v o r a , and the only known American hyaenid m a y be a d d e d : Chasmaporthetes, which appears in the early Pleistocene C i t a Canyon fauna a n d has a close relationship with the European Euryboas, the „ S p r i n t e r hyena". The importance of the Proboscidea has already been stressed, a n d numerous examples will be found among the rodents. Horses and camels exemplify migration from the N e w W o r l d to the O l d , perhaps a subsequent remigration of modernized Equus into N o r t h America ( M C G B E W , 1 9 4 4 ) . Elk
Pleistocene mammal magrations from the Palaearctic to the Nearctic
103
(Alces), reindeer (Rangifer), y a k (Bos), bison (Bison), musk ox (Ovibos) a n d sheep (Ovis) are important artiodactyl migrants from the O l d W o r l d to the N e w , most of them in the late or latest Pleistocene.
References BROWN, Barnum: The Conard Fissure, a Pleistocene bone deposit in northern Arkansas: With description of two new genera and twenty new species of mammals. - Mem. American Mus. Nat. Hist. 9 ( 4 ) , 1 5 7 - 2 0 8 , N e w York 1 9 0 8 . GIDLEY, J . W., & GAZIN, C . L . : The Pleistocene vertebrate fauna from Cumberland Cave, Mary land. - Bull. U . S . Nat. Mus. 1 7 1 , 1-99, Washington 1 9 3 8 . HIBBARD, Claude W.: Summary of North American Pleistocene mammalian local faunas. - Papers Michigan Acad. Sei. 4 3 , 3 - 3 2 , Ann Arbor 1 9 5 8 . JOHNSTON, C. S., & SAVAGE, D . E . : A survey of various late Cenozoic vertebrate faunas of the Panhandle of Texas, part I: Introduction, description of localities, preliminary faunal lists. - Univ. California Publ. Geol. Sei. 3 1 , 2 7 - 5 0 , Berkeley 1 9 5 5 . fvRAGLiEviCH,Lucas Jorge: Smilodontidion riggii, n. gen. n. sp. Un nuevo y pequeno esmilodonte en la fauna pliocena de Chapadmalal. - Rev. Mus. Argentino Ci. N a t . 1, 1-44, Buenos Aires 1 9 4 8 . KURTEN, Björn: A skull of the grizzly bear (Ursus arctos L.) from Pit 1 0 , Rancho La Brea. - Los Angeles County Mus. Contr. Sei. 3 9 , 1-7, Los Angeles 1960. KURTEN, Björn & RAUSCH, Robert: Biometrie comparisons between North American and European mammals. - Acta Arctica 1 1 , 1 - 4 4 , Copenhagen 1 9 5 9 . M G R E W , Paul O.: An early Pleistocene (Blancan) fauna from Nebraska. - Geol. Ser. Field Mus. Nat. Hist. 9 , 3 3 - 6 6 , Chicago 1 9 4 4 . MATTHEW, William D . : Critical observations upon Siwalik mammals. - Bull. American Mus. N a t . Hist. 5 6 ( 7 ) , 4 3 7 - 5 0 0 , New York 1 9 2 9 . M E A D E , Grayson E . : The saber-toothed cat Dinobastis serus. - Bull. Texas Memor. Mus. 2 , 2 5 - 6 0 , Austin 1 9 6 1 . P E I , Wen-chung: On the Carnivora from Locality 1 of Choukoutien. - Palaeont. Sinica, ser. C , 8, C
1-216, Peking 1 9 3 4 .
SCHAUB, Samuel: Über die Osteologie von Machaerodus
cultridens
Cuvier. - Eclog. geol. Helv. 1 9 ,
2 5 5 - 2 6 6 , Basel 1 9 2 5 .
TAYLOR, Dwight D . & HIBBARD, Claude W.: Summary of Cenozoic geology and paleontology of Meade County area, southwestern Kansas and northwestern Oklahoma. - Pamphlet, 1 - 1 5 7 , Ann Arbor 1 9 5 9 .
TEILHARD DE CHARDIN, Pierre: Les formes fossiles. - Les felides de Chine, 5 - 3 5 , THENIUS, Erich: Über einen Kleinbären aus dem Pleistozän von Slovenien nebst Phylogenese der plio-pleistozänen Kleinbären. - Razprave Slov. 6 3 3 - 6 4 6 , Ljubljana 1958. - - Ursidenphylogenese und Biostratigraphie. tierk. 2 4 , 7 8 - 8 4 , Berlin 1 9 5 9 . TOBIEN, Heinz: Cuon
H O D G . und Gulo
Peking 1 9 4 5 . Bemerkungen zur Akad. Znan. 4 , - Zeitschr. Säuge-
FRISCH (Carnivora, Mammalia) aus den altpleistozänen
Sanden von Mosbach bei Wiesbaden. - Acta Zool. Cracoviensia 2 , 4 3 3 - 4 5 2 , Krakow 1 9 5 8 . W E S T , R. G.: The glacial and interglacial deposits of Norfolk. - Trans. Norfolk & Norwich N a t . Soc. 1 9 , 3 6 5 - 3 7 5 , Norwich 1 9 6 1 .
Manuskr. eingeg. 3. 1. 1 9 6 3 . Anschrift des Verf.: Dr. B. KURTEN, Zoologisches Museum der Universität, N . Järnvägsgatan 1 3 , Helsingfors, Finnland.