Compositae of Ecuador, II. Diversity and distribution - Funk 1997

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Estud ios sobre ]" ·--··_.wersidad y

c.ologia .· .'_: de : I ntas

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ESTUDIOS SOBRE DIVERSIDAD Y ECOLOGIA DE PLANTAS Memorias del II Congreso Ecuatoriano de Botanica realizado en la Pontificia Universidad Catolica del Ecuador, Quito 16—20 Octubre 1995

editores Renato Valencia & Henrik B aislev

Publicado por PONTIFICIAUNiVERSIDADCATOLICADEL ECUADOR en colaboracidn con UNIVERSIDADDEAARHUS,DINAMARCA (PROYECTO ENRECA) Y PROGRAMADANESDEINVESTIGACIONDELMEDIOAMBIENTE (PROYECTO DIVA)

1997

CONTENIDO

p.

Contenido Introduccion y presentacion Dedicatoria Agradecimientos Asesores cientfficos Florfstica y taxonomia 1. Cornelia Ott- Notes on the systematics and ecology of Menispermaceae of Ecuador 2. Laurence E. Skog & Lars Peter Kvist - The Gesneriaceae of Ecuador 3. Vicky A. Funk- Wemeria s.l. (Compositae: Senecioneae) in Ecuadof 4. Maximilian Weigend -Some aspects of the biogeography. morphology and systematics of Loasoideae in northern ~th~~a

iii v

-vi viii

1

i3 25

r

5. Lynn G. Clark - Diversity and biogeography of Ecuadorean bamboos (Poaceae: Bambusoideae) and their ~~

6. J-:Iarold Robinson & Vicky A. Funk- Compositae of Ecuador I: Key to frequently collected genera 7. Vicky A. Funk - Compositae of Ecuador II: Diversity and distribution 8. Lars W. Chatrou, Paul J. M. Maas, Carola P. Repetur & H. Rainer - Preliminary list of Ecuadorean Annonaceae Descripcion de Ia vegetacion 9. John Littner Clark- Preliminary floristic inventory of the Bilsa Biological Station, Esmeraldas, Ecuador 10. Blanca Leon, Kenneth R. Young & Asuncion Cano - ¡ Fitogeografia y conservaci6n deJa Costa Central del Peru 11. Tamara Nunez- Inventario florfstico y zonificacion de Ia vegetacion en Isla La Plata, Parque Nacional Machalilla 12. Carlos E. Ceron & Consuela Montalvo A. - Composici6n y estructura de una hectarea de bosque en la Amazonia Ecuatoriana - con informacion etnobotanica de los Huaorani

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65 79 97

123 129 143

153

Compositm: of Eciiiimii"

Etn-positae of Ecuador, II. Di'\lersity-and distribution V. A.-Funk U. S. National Herbarium, Department of Botany, Smithsonian ln~titution, MRC 166, Washington, D.C., 20560, USA, tel (+1) 202-357-2560,fax (+1) 202786-2563, emnil nmhbo003@sivm.si.edu Abstract The Compositac are a large and d ive('Sc tamily in Ecuador with 152 genera and over 753 species found over 1000 min elevation. The family is divided into many tribes, 12 of which are found in Ecuador: Anthemideae, Astereae, Barnadesieae, Chichorieae, Eupatodeae, Gnaphalieae, Heliantheae s.L Liabeae, Mill.iSie.ac,_I'.Lu..che.e.a.e,_Senecioncac~anc:lj{ernonieac....The-provinccs with the most taxa are Pichincha, Chimborazo,Az uay, and Loja. The tribes with the largest number of species in Ecuador ¡are the Heliantheae, Eupatorieae, Senecioneae, and Astereae. Urief descriptions arc provided for each tribe. All of the genera known to occur in Ecuador, inclu ding those under 1000 min elevation are listed in Appendix A. Resumen La fam il ia Compositae es mra familia grande y may diliersa En el ccuaoor; se ha cncontrado 152 generos y mas que 753 especies encima de 1000 metros sobre el nivel del rna,. La familia induye un gran numero de tribus, de los cuales 12 estan reprcsen tados e n el Ecuador: Astereae, Barnadesieae, Chichorieae, Eupatorieae, Gnaphalieae, Heliantheae s./., Liabeae, Mutisieae, Pluchccae, Senecioneae y Vernonieae. Las provincias que mas taxa tienen son Pichincha, Chimborazo, Azuay y Loja. Los tribus con el numero mas alto de especies en el Ecuador son Heliantheae, Eupatorieae, Scnccioneae y Astereat>. Este articulo presenta descripciones cortas de cada uno de los tribus. El Appendice A incluye una lista de todos los generos de Compositae que se han registrado en el Ecuador.

Introduction The Compositae (Asteraceae) is the largest family of flowering plants in the world with approximately 25,000 species, 1I 10 of all flowering plants. In the Andes the Compositae range from ephemerals to 20 m tall trees and are found in all habitats from forests to paramo and puna, although they are not well-represe nted in the low elevation rain forests. Except fbr areas of permanent snow and lowland tropical forests, Compositae make up an important part Of the flora and if one wishes to understand the composition and diversity of an area one must understand the Compositae.

The Compositae for the Flora of Ecuador are being treated by tribe; the Liabeae (Robinson, 1978) and the Mutisieae s.l. (Harling 1991) are completed, the Ser}ecioneae is nearing completion (Nordenstam MS), a nd the Heliantheae is in draft form (Robinson MS). There are several books and checklists for Ecuador: the Amazonian Ecuador (Renner et al. 1990), trees and shrubs of the Andes (Ulloa Ulloa & }0rgensen 1993), plants of the high Andes

79

1,-: A. F11•rk

Compv~ita c

(J0rgensen & Ulloa Ulloa 1994), a check list for pa ramos ~Luteyn, MS), and a checklist of Compositae of the Andes found over 1000 m (Funk et a/. 1995; Fu nk iv'!S). Jerg::!nsen is spearheading a project to prcduce a checklist of all plants of Ecuador but the Compos.itae arc not finished. Even the checklists that exist have problems as they mus t I.Je constantly updated because Of nomenclature changes, new records, and newly described genera and species.

Materials ami 111ethods All specimens in the U. S. National Herbarium we.re surveyed (ca. 600,000) and those species th<1t were collected in the Andes over 1000 m in elevation were recorded in the data base. As additional collections of recorded species were found in a new province or at elevations not already covered, these were added to the data base. The collections at US are strong for the northern and central Andes. The total number of species of Compositae in the Andes above 1000 m is ca. 3700 and wi!l.prob2bly reach dose to 4000 by the time the project is completed.

As with any checklist the names are constantly changing as genera and species are described or sunk and generic realignments are made. A number of changes have occurred since the data base was finished including the descriptions of new genera, such as Idiopappus (Robinson & Panero 1994), Xenophyllulm (Funk in press), and the locating of Ecuadorian species of genera not previously known from Ecuador, such as Crossothamnus King & H. Rob. (King & Robinson 1995) and Talnlllnnca/{a H. Rob. & Cuatrec.

(Nordenstam & Pruski 1995). Realignments of taxa include the species commonly referred to Aster which in Ecuador have been placed into .

Symphyotrichum Nees (Nesom 1994), many species of Pentacalia being transfered into Monticalin (Nordenstam 1996), and of all the species of Culcitium in Ecuador that were moved into Senecio, one now appears to be recognized, Culcitium rufescens. Other additions were simply missed, such as the placement of all the Ecuadorian species of Aspilia Thouars into Elaphandra Strother (Robinson 1992). These types of changes are constantly altering the data base.

In order to make Appendix A, the list of genera found in Ecuador, as complete as possible it has been updated to reflect all these types changes. In addition, about 15 genera have been added that are found under 1000 m. The result is the most complete list of taxa for Ecuador that could be compiled at this time, all based on identified material. Unfortunatley, the maps and tables are current only to October 1995.

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80

of Ew111i!;r

Results and discussion Based on these data, Ecuador has approximately 152 genera and over 750 -species of Compositae found over 1000 m in elevation (Table l). For the Andean Countries this is significantly ·exceeded -6ii1YlW Peru (187 genera'/1143 species) and Colombia (171 /993) and it is very close to the fig ures for Bolivia (161 I 802). Given the size of its montane area H has an extremely diverse Compositae flora with approximately 20% of the Compositae species in the Andes having been found in Ecuador.

Table I. Composi.tae, number of genera and s pecies above 1000 m in the And ean countries.

Country I species

#of gen.

Venezuel2 Colom bia Ecuador Peru Bolivia Argentina* Chile*

142/ 535171 / 993 152/755 187/ 1143 161 /802 136/629 57/270

1QOtb30DO m

>30oo-m=

-·------- -13.2t.A6G160/786 150/621 173/816 139/566 127 I 485 47/194

-36/115 62/345 70/3i 7 1()2/519 74/339 44/189 . 27/92

·- · - *US herbarium is net-s~rong fo r these countries

The distribution of the taxa in Ecuador reflects the elevation with the provinces of the Andes having the most followed by those of the slope forests (Fig. 1). The provinces with the most taxa are two fro in the north central area of Ecuador, Pichincha (104 genera /288 species) and Chimborazo (91 /225), and two from the south, Azuay (81 / 218) and Loja (84/ 203) (Table II).

Pichincha and Chimborazo are near the big herbaria in Quito and one might argue that they are better collected than those in the south, so the current work under way in the south by Peter Jergensen, Carmen Ulloa Ulloa, and their collaborators, may bring these two southern provinces closer in number to those near Quito.

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V. A. Funk

Composit11c of ECimdor

Table II. Compositae genera and s pecies above 1000 m in each p rovince of Ecuador.

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

Province

# o f gen.

I s pecies

Pichincha .

Chimborazo Azuay Loja ¡ Napo(+Sucumbios) Carchi lmbabura Tungu rahua Cotopaxi Cafiar J3olivar

104/288 91/225 81/218 84/203 63/163 66/150 62/ 137 66/120 55/113 57/96 44/77

1000- .

3000 Ill 94/213 73/139 68/165 80/190 35/68 54/95 43/62 55/88 37/54 46 /68 32/51

>3000m

art:a in sq km

60/165 47/112 60 / 144 29/54 40/106 42/105 46/110 25/40 33/70 34/56 22/29

12,872 6,522 8,098 11,214.

53,835 3,750 4,459 3,128 6,248 3,184 4_.105

The four tribes with the most species in Ecuador are the Heliantheae, Eupatorieae, Senccioneae, and Astereae, all with over.lOO.species..(Tab-1~--Ill}.~ The Heliantheae (47 genera /149 ~pecies;-Eig 2), and Eupatprieae (32f1:SIJ are the la.rges.t and they have important centers of-distribution in the Andes and are found wholly or primarily in the western hemisphere. Both mprphology and molecular data indicate that these tribes are each others closest relatives and that together they form a well supported monophyletic group (Bremer 1994, Kim eta/. 1992). The monophyletic group formed by these two tribes has a combined count of ca. 80 genera and 300 species. Table III shows that these taxa are found mostly between 1000 m and 3000 m with just over 1/3 above 3000 m. The H eliantheae have only one genus that is found exclusively above 3000 m, Aphanactis.

Remain'ing nine provinces have small numbers of species.

-0-

HEUANTHEAE

Figure 1. Map of Ecuad or showing the distribution of the genera/ species of the Compositae by province (Sucumbios is included in Napo).

82

Figure 2. Map of Ecuador showing the distribution of the genera/ species of the tribe Heliantheae (Compositae) by province (Sucumbios is included in Napo).

83

V. A. Fuuk

Table tribe.

W. Compositae, numbers of genera and species above 1000 m ir. each

Tribe ··-····..._ __

l.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

Cum1'ositae of Ecundor

Heliantheae Eupatorieae Sen-ecioneae Astereae Gnaphalieae Mctisieae Liabeac Vernonieae ·chichorie ae Barnadesieae Anthemideae

·- - - #of gen.

I species

1{)00· 3000 m

47/ 149 32/151 13/126 9 / 112 10/51 9/42 8/42 10/28 2/20 3/13 7/17

46/142 32/136 12/84 8/75 8/27 9/ 37 7/40 10 /27 2/14 3/12 7/12

>3000 Jr.

-

---· -----23/45 14/67 9/97 ---9/90 9/45 6/ 27 8/21 2/5 2/14 3/9 5/13

The next largest tribes are the Senecioneae (13 genera/ 126 species; Fig. 3) ar,d the Astereae (9 I 112) which are well represented in the Andes and have specialized element3 there, but are common in both the eastern and western hemispheres. These two tribes are not sister taxa and therefore are not as closely related to one another as the Heliantheae and Eupatorieae. Table III shows that these two tribes have half or more of their genera above 3000 m. !here appears to be a split between the Helianth eae/Eupa torieae and the Senecioneae I Astereae that follows the lines of their worldwide distributi on.

One cannot help but wonder if there is some connection.

Pichincha is most likely the. best collected province and so it should give us a window into overall diversity. There are over 100 genera and nearly 300 species in this province. The numbers of species within each of the tribes of the Compositae it is rather impressive with the Eupatorieae (19 genera/ 56 species), Heliantheae (32/51), Senecioneae (10 /50), and Astereae (9 / 46) h aving the largest numbers.

The Liabeae in Ecuador (8/42) is the tribe that is most likely of Andean origin with possibly several different dispersal events into Mexico, Central America, and the Caribbea n (Funk et al. 1995). For most of the tribes there are only data for collections over 1000 m in elevation, however, for the Liabeae the data have been collected for all elevations. These additional low elevation data produced an increase of ca. 10 new province records for Ecuador but the 84

new data did not add any additiona l species for the country. The tribe Liabeae has eight of its 15 genera represented in Ecuador and 42 out of approximo:tely 160 species. These taxa are distribute d in 18 prov mces (Sucumbios included in Napo). There are four provinces with relatiYf4Uarge= numbers of Liabeae, Loja (7 /18), Azuay (7 / 13), Pichincha (6/ln and Chimbor azo (6/14) (Fig. 4). Munnozia and Liabum are by far the largest genera in the tribe, each with 12 species in Ecuador, and although they ere widespread in the Andes with some species ·ranging into Central America and the Caribbean, they are centered in Peru and Ecuador.

The tribe Mutisieae is well represen ted for it's size (worldwide: 76 genera, ca. 970 species). The remainin g tribes such as the Plucheeae and Chichorieae (Lactuceae) are limited to a few endemic s and/ or a few widespread. species (Table ITi;.Appendix- A). There were som e genera that one would expect- to find in the mountains of"Ecoad or that were- not-feufld---sueft=as Oyedea" (I-Ielianth eae ).

·0

SENECIONEAE

Figure 3. Map of Ecuador showing the distributio n of the genera/species of the tribe Senecioneae (Compositae) by province (Sucumbfos is included in Napo). ·

85

V. A. Ft111k

C6mpositm• of Ecuador

ally yellow; anthers appendaged at apex, base obtuse without tails; styles without appendages, apex truncate; pappus absent or reduced. (Worldwi de:109 ge1'1era, 1740 species; Ecuador: 7 I 17) 2:-Astere ae: mainly annua s; eaves usually alternate; no milky sap; no cha: racteristic pl;lbescel}ce; phyl lari~equal,..imb.cicatain-Seve~ep=­ tacle naked; ray florets present and in large numbers; disk florets regular; anthers appendag ed at apex, base obtuse; styles with sterile appenda ges, adaxially devoid o£ style hairs, apex rounded; pappus usually setose.

(Worldw ide: 170/3000; Ecuador: 9 / 112) 3. Barnad esieae - perennial herbs or shrubs; leaves alternate, axillary spines frequently pres(JJ:lti no milky sap; no characteristic pu~esceHce, imolucral bracts in several ro vv s, cltartaceo us, freq uently spine-tipped; recepta€1e= naked;__ray florets p resent or abse11t, bilabiate or subligula le; disk florets m ore or less regular, deeply 5-lobed or 5-cleft, with equal or unequal segments, but never truly bilabiate; anthers apical appendag es rounded to obtuse o r acu te, emargina te to bilobed, base with very short or absent appendages; styles shallowly d ivided at the apex; pappus of villose bristles, rarely of scales or absen t. (Worldwide: 9 /92; Ecuador 3/ 13)

Figure 4. Map of Ecuador showing the distributio n of the genera/ species of the tribe Liabeae (Composi tae) by province (Sucumbfos is included in Napo).

A Key to the tribes of the Composi tae in Ecuador and a glossary are found in the following article (Robinso n & Funk, this volume).

Tribe descripti ons (All genera and species numbers are approximate) 1. Anthemideae - mainly annuals some subshrubs or shrubs, often odoriferous; leaves generally alternate usually much divided or toothed; no milky sap; no characteristic pubescence; involucra! bracts imbricate in several rows, wholly scarious or herbaceo us with a scarious margin; receptacle naked or with thin deciduous bracts; ray florets present or absent; disk florets regular; ray florets mostly white or yellow sometime s pinks or blues; disk florets usu86

4. Chichorieae (Lactuceae) - herbs, rarely shrubs or trees; leaves alternate, rarely much divided; milky sap; no characteristic p ubescence; involucr a! bracts..imbricate in several rows· or equal and uniserate; receptacle naked or with thin deciduou s paleae; florets all ligulate, often yellow or blue; anthers apical appendage elongate, base calcarate and caudate; style branches slender, papillose and sometimes flattened; pappus usually with one or more rows of se!ose or plumose bristles, sometime s paleaceo us or awns or absent.

(Worldw ide: 98/ 1550; Ecuador: 2/20).

5. Eupatorieae- commonly herbs or shrubs; leaves opposite; no milky sap; no characteristic pubescen ce; phyllarie s imbricate in several rows; receptacle naked; ray florets absent; d isk florets regular; florets blue, purple, pink, or white, never yellow; anthers apical append age ovate to oblong, base obtuse; style appendages long and flattened with apex club shaped, distinct in that they are· the color of the corolla; pappus usu ally of fine but ridged setae, sometime s plumose or scaly or absent. (Worldwide: ca. 170/2400; Ecuador: 32/ 151) 6. Gnaphalieae- Perennial or annual herbs or shrubs; leaves alternate; no

87

V. A Fu11k

Compt!sitnc of Ecuador

milky sap; leaves often with woolly hairs; phyllaries imbricate in several rows, papery and often showy; receptacle usually naked; ray florets present or more Lo;nmonly abseni; marginal female florets when present filiform, rarely radiate; disk florets regular, often functionally male; florets often yei!ow; anthers apical appendage flat cr less frequently concave, base with short tails; style arms various without terminal appen<!ages; pappus setose, barbeilate or plumose, or of scales or Clbsent. (Worldwide: 130/2000; Ecuador: 10/51)

gular, often functionally male; florets often yellow; anther apices flat, base usually tailed; style arms various without terminal appendages; pappus of barbellate capillary bristles or of scale-like bristles. (Worldwide: 28/220; Ecuador: 2/2)

7. Heliantheae s.l. - Perennial-~erbs or shrubs; leaves usually opposite; no milky sap; no characteristic pubescence; involuceral bracts usually biserate sometimes imbricate in several rows; receptacle usually paleaceous; ray florets usually present, ligu 1atc; disk florets reguiar; ray florets usually-white-or yellow; disk florets usually yellow but both also orange-red; anthers appendaged at apex, base not tailed but sometimes with sharp points, often black; style branches various, with or without appendages, apex various; pappus 2-3 ridged awns or scales or paleaceous or absent. (Worldwide: 189/250J;Ecuacto;:47/149) 8. Liabeae - Perennial herbs or shrubs; leaves opposite or congested in rosette; often with milky sap;. often with dense white hairs on under surface of leaves; involucre usually subimbricate with bracts in many graduated series; receptacle naked or nearly so; ray florets usually present, ligulate; disk florets regular but deeply lobed; ray and disk florets nearly always yellow; anthers appendaged, base sagittate, rounded or fringed; style branches with little or no appendage; pappus various. (Worldwide: 15 I 160; Ecuador: 8/42).

11. Senecioneae - Perennial herbs, shrubs or trees; leaves¡ alternate although sometimes alternate; no milky sap; cottony hairs are common; involucra! bracts more or less equal and generally in one series often subtended ¡by a row of shortened bracts (calyculus); receptacle usually naked; ray florets when present, ligulate; disk florets regular or disciform; ray florets yellow, white, or purple; disk florets yellow; anther apically appendaged, base sagittate; style branches various, with or without¡ appendages but often with -ap~runcate and fringed; pappus setose, with numerons-fine-wft;- simple bristle-S~(Worldwide: 120 /3000; Ecuador: 13 /126) 12. Vernonieae - Perennial. herbs or shrubs, less frequently vines and large trees; leaves alternate; no milky sap; no characteristic pubescence; iltvolucral bracts imbricate in several rows; receptacle naked; ray florets absent; disk florets regular; florets usually blue or purple; anthers with apical appendage elongate, or obtuse, base sagitate, filamenrsinserted nign above base; styles deeply divided into 2 long semi-cylindrical pointed limbs, acute at ape~; pappus of many barbellate bristles. (Worldwide: 93/ 1300; 10 /28)

10. Plucheeae - Perennial herbs or shrubs; leaves alternate; no milky sap; leaves often with woolly hairs; phyllaries imbricate in several rows, cartilaginous, sclerenchymatous portion undivided; receptacle usually naked; ray florets rare; marginal female florets usually present, filiform; disk florets re~

Genera There are 152 genera of Compositae recorded from Ecuador (Appendix A has 180 listed but only 152 are in maps and tables) with approximately 753 species. The genera with the largesf number of species are Pentacalia (33 species), Gynoxys (32), and Senecio (24) in the Senecioneae; Baccharis (47) and Diplostephium (25) in the Astereae; and Mikania (38) and Ageratina (23) in the Eupatorieae. Illus trations of some of these genera can be found in Robinson and Funk (this volume). Interestingly enough while these three. tribes all have large numbers of species in Ecuador, the tribe that has the largest number of specie$ in the country, the Heliantheae (47/154), has no genera with more than 18 species, the largest being Verbesina (18) and Clibadium (14; Fig. 4a, Robinson & Funk, this volume), and the remainder all have 10 or fewer species. Comparing the number of species to the number of genera in the four larg'Tst tribes, the Heliantheae have a ratio of 3:1, the Eupatorieae 5:1, the Senecion~ae 10:1, and Astereae 10:1. The Heliantheae, and to a lesser extent the Eupatorieae, have fewer species per genus than the other major tribes.

This may be a result of the fact that the Eupatorieae and Heliantheae are

88

89

9. Mutisieae - Perennial herbs or more frequently a shrub; leaves alternate, rarely much dissected or prickly; no mil~y sap; no characteristic pubescence; involucra! bracts imbricate in several rows; receptacle naked; florets, especially the marginal ones, bilabiate; anthers appendaged at apex, base sagittate with long basal sometimes laciniate, appendages; style branches usually short or erect, connivent, obtuse or truncate at the tips without appendages; pappus of one or more series of bristles, rough or plumose, seldom formed of paleae or absent. (Worldwide: 76/970; Ecuador: 9/42)

V. A. Ftlltk

almost exclusively new world tribes, have been worked on extensively, and segrega ted at the generic level quite a bit while the Senecioneae and Astereae are larger and mo re global in their distribution and still contain large core genera that probabl y will be broken up in the future. Indeed the Astereae are being revised now by Nesom .(1994).

The 22 genera w ith 10 or more species are as follows: ASTEREAE: Baccharis (47), Conyzn (13), Diplostephium (25), Erigeron (10) CICHORIEAE: Hieracium (10}, Hypochoeris (10) EUPATORIEAE: Ageratina (23}, Mikania (38}, Stevia (13) GNAPHALEAE: Gnaphalium (15) HELIANTHEAE: BideHs (11), Clibadium (14}, Verbesina-{18} LIABEAE: Liabum (12}, Munnozin (12) MUTISIEAE: Mutisia (12), Jungia (11) SENECIONEAE: Gynoxys (32}, Pentacalia (33}, Senecio (24), La_?iocephalus {13) VERNONIEAE: Critoniopsis (12) In addition , there are a 35 genera that have less than 10 species in Ecuador but have at least one species that is very frequently collected. They are as fo11ows: ASTEREAE: Oritrophium BARNADESIEAE: BamadeS'ia, Chuquiragn EUPATORIEAE: Adenostemma, Ageratum, Aristeguietia, Austroeupatorium ,

Badilloa, Chromo/aena, Fleischmannia, Heterocondylus, Polyanthina GNAPHALIEAE: AcT1yrocline, Gamochaeta, falcophila, Loricaria HELIANTHEAE: Acme/la, Aph11nactis, Galinsoga, Pappobolus, Smallanthus, Tridax, Wulffia LIABEAE: Erato, Philoglossa MUTISIEAE: Perezia PLUCHEEAE: Tessaria SENECIONEAE: Culcitium (all but one species of this genus belong m Senecio), Pseudogynoxys, Werneria, Xenophyllum VERNONIEAE: Cyrtocymura, Lepidaploa, Piptocarpha, Vernonanthura There are a total 57 genera in the two groups listed above. A key to these genera (includi ng a glossary) is provide d in Robinso n and Funk, this volume.

For the Compos itae the species of the very high elevatio n areas are probably better known than other areas and this is reflected in the fact that most of

90

Composilae of Ecuador

the recently describe d new species are collected elsewhere, primari ly below 3000 m on the eastern slopes of the Andes. Recently Robinso n and Cuatrecasas (1993) describe d over 20 new species of Pentacalia from Eettado r and Peru, all from the eastern slope forests. fn addition, Sagastegui-Alva-and Dillon (1994); Zermog lio and Funk (in press) and others have found:new taxa on the eastern slopes of Ecuador and Peru.

Conclusion Ecuador has a ri.ch. and diverse Compos itae flora. with a concent ration of species in the provinc es of the Andes followe d by those of the eastern slope forests of the Andes. This paper examine s those species with specime ns in the U.S. Nationa l Herbari um with label information that indicate they were Wll~tecl af1000- m or a ove 111 elevatio n (152 genera, 753 species).

There are few species at low elevatio ns in either the west or the east and most of those are either weedy or belong to a few genera that frequent the lowland s such as Mikania and Clibndium. Many of these species are found above-1000 m as well as below. Howeve1~ there are a numbgr-e~ne.ra-that::are:;f{lttnd primarily below 1000 m. An effort has been made to identify these genera and all that were found were added to Append ix -A to bring the total number of genera for Ecuado r to 180. Most of these additions to the generic list are represented by one species and so the total number of species is probabl y close to 785. No doubt it will climb higher as new records and taxa are found.

Acknow ledgeme nts I thank G. McKee for helping maintain the data base and H. Robinson and two reviewers for their critical commen ts.

Literature cited BREMER, K. 1994. Astemceae: Cladistics and Classificatioll. Timber Press, Portland, Oregon.

FUNK, V. A., ROBINSON, H., MCKEE, G. 5. & PRUSKI, J. F. 1995. Neotropical montane Compositae with an emphasis on the Andes, pp. 451-471 in Churchill, S., Balslev, H., Forero, E. & Luteyn, J. (eds.). Biodiversity and Conservation of Neotropical Monta11e Forests.

New York Botanical Garden, New York.

KIM, K.-J., JANSEN, .R. K., WALLACE, R. 5., MICHAELS, H. ]. & PALMER, J. D. 1992.

Phylogenetic implications of rbcL sequence variation in the Asteraceae. Amrals of tire Missouri Bola11ical Garden 79: 428-445.

J0RGENSEN, P. & ULLOA ULLOA, C. 1994. Seed plants of the high Andes of Ecuador -A checklist. AAU Reports 34.

NESOM, G.¡L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. Phytologia77: 141-297.

NORDENSTAM, B. & PRUSKI, J. F. 1995. Additions to Dorobaea and Talamanca/ia (Composi tae: Senecioneae). Compositae Newsletter 27: 31-42.

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V. A F:rnk

Compvsitae of Ecuador

29: 47·50.

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• SAGASTEGUl-ALVA, A. & DILLON, M. 0. 1991. Estudio£ en Ia tribu Liabcac (Aster?.:::eae) ·~n Peru: III. Una nueva cspecie de Cllrysactinium de ~:ortc del ?cru. Amaldon 2(2): 31-35.

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AAU Reports 30. r .

ZERMOGLIO, M. F. & FUNK, V. A.. In press. A new species of Cllrysact:. . nwm {'-ompos 1ta~: Liabeae) from Ch~chapoyas, Peru. BioL/anin.

Appendix A. Compositae genera in Ecuador. * recently added to this list, not indude d on maps and tables; **Only Culcitium rujcscens which W<lS counted in the tables and on maps as..::Senecio; + commo nly collected naturahred gene-ra ; ++ recently describ ed segregate genus- species include d in anothe r genus in the tables and maps ASTEREAE Bacclmris L. Conyza Less.

Diplostephium S. F. Blake Egletes• Cass.

Erigeron L. Laestarlia Kunth ex Less. !.lerasia Triana Noticastmm DC. Oritroplrium (H.B.K.) Cuatrec.

Plagioclreilus Arnott ex DC

Bar"l/ctlilla R. M. King & H. Rob. Brickcllia Elliott Clrromolawa DC. Co,dylirl ium R. M. King & H. Rob. Conclylopodium R. M. King & H. Rob. Critouia P. Br. Critouiella rt M. !<:ng& H:-Rob...; Cronquistiautlrus R. M. King & H. Rob. Crossollramlllts• R. M. King & H. Rob. Fleischmmutia Sch. Bip. Guevnria R. M. King & H. Rob. Grosveuoria R. M. King & H. RTIO. Hebec/iuiwn DC. Heteroconrlylus R. M. King & H. Rob. Kaunin R. M. King&: H. Rob. Koauophyllou Arruda Mikmrin Willd.

Neomirnudca R. M. King & H. Rob. Oplrryosporus Meyen Plm!ncraen DC. Piquerin Cav. Polynuthina R. M. King & H. Rob. Scindoceplw/n Maltf.

StevinCav.

ANTHEMIDEAE Achillea-b.Artemisia L. Cotula L. Leucnuthemum Mill.

Matricaria L. So/iva Ruiz & Pav. Tmracetum L. BARNADESIEAE Banrnrlesia Mutis Clwquiraga Juss.

Dasyphyllmu H.B.K.

GNAPHALIEAE Achyrocline (Less.) DC. Antwnnria• Gaertn.

Be/loa Remy Chevreulin Cass.

Cuntrecnsasiclln H. Rob. Face/is* Cass.

Gamochaeta Wedd.

Gnaphalium L. Jnlcophiln M. 0 . Dillon & Sagast.

Loricaria Wedd.

Lucilia Cass.

Stuckertiella Beauverd

CiCHORIEAE Hieraceum L. Hypoclroeris L. Sonclzus+* L. Taraxacum+• Weber in F. H. Wigg EUPATORIEAE Ageratina Spach AgeratumL. Adenostemnra J. R. Forst. & G. Forst.

Aristeguietill R. M. King & H. Rob. Asplundia nfhus R. M. King & H. Rob. Austroeupaiorium R. M. King & H. Rob. Ayapana Spach.

Ayapanopsis R. M. King & H. Rob. Badilloa R. M. King & H. Rob.

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HELIANTHEAE Acanthospermum• Schrank Acmella Rich.

93

Compositnc of Ec~tador

V. A. Funk Slciractinin S. f. Blake S yncdrei/a Gaert~<.

Tngetcs L. Tridax L. Vcr/Jesina L.

Alloispemwm Willd.

Ambrosia• (,..

Aphnnactis Wed d.

Bnltimora• L. Bidcns L. Calen L. Clibndium L. Coreopsis L. Cosmos+ Cav. Cyathomone• S. F. Blake Deli/in Spreng.

Ecliptn L. Elnphnndrn Strother [Aspilia pro parte) Eleut!Jerant!Jera• Poit. ex Bose.

Enydrp• Lour.

Ericentrodea S. F. Blake & Slmff Espeletia Mutis ex H. & B. Flnverin Juss.

Gnlinsoga Ruiz & Pavon Gnrcilnssa Poepp. & End!.

Heliopsis Pers.

Heterospermn Cav. Hidnlgoa La Llave & Lex. Ichthyuthere Mart. ldiopnppus• H. Robins. & Cuatrec. fnegeria H.B.K. Kingianthus H. Rob. Lngnscea• Cav. Melanthera• Rohr Milleria• L. Monactis H.B.K. Montanan Cerv.

Neuroleana R. Br. Oblivia• Strother Pappobolus S. F. Blake Pnrthenium L. Peel is• L. Perymmium Schrad.

Porop!Jyllum Guett.

Sabnzia Cass.

Sal mea DC. Schistocarp!Ja Less.

Schizoptera• Turcz.

Sc!Jkuhria Roth.

Sigesbeckia L. Simsia Pers.

Smallanthus Mack.

Sphagneticola 0. Hoffm.

Spilanthese" ]acq. · Stachycephalum• Sch. Bip.

Viguiera H.B.I<.

Villanova Lag. Wede/in Jacq.

Wulffin Neck. ex.Cam. (Ti/esin) Xantlrium L. LlABEAE Cacosmia H.B.K: Clrrysactinirrnr (H.B.K.) Wed~.

Erato DC. Ferreyrantlws H. P..a~. &',~rcl_tell Liabum Adans.

Mrmnozia Ruiz & Pav. Oligactis (H.B.K.) C<;f.s.

Philoglossa DC.

I'seudogynoxys (Greenm.) Cabrera Senecio L. Talnmancalin• H. Rob. & Cuatrec.

Wemcrin !-I.B.K. Xenupllyllrf!~~Z+ V. A. Funk VERNONIEAE,...

Critoniopsis Sch. Bip. Cuntrecasantlrus H. Rob. Cyrtocymrrra H. Rob. Eirmoceplrnla H. Rob. Eleplrnntopus L. ·Josemrthus H. Rob. Lepidaplon (Cass.) Cass.

Pacouri11a• Aubl.

Piplocarp!Ja• R. Br. Piplocoma Cass. [incl. Pollnlesla] Pseudelephn11/opus Rohr Venronmrlhurn H. Rob.

MUTISIEAE Clraptalin Vent.

Gcrbcra L. /lmgia L.f.

Mutisia L.f.

011oseris Willd.

Perezia Lag. Stmopndus S. F. Blake Triclrocline Cass:Trixis P. Browne PLUCHEEAE Pseudoconyza Cuatrec. (Blumea DC.) Pluc!Jen• Cass.

Tessaria Ruiz & Pav. SENECIONEAE Aequatorium B. Nord.

Culcitirmr•• Bonpl.

Dendroplrorbium (Cuatrec.) C. Jeffrey Dorobnen Cass.

Emilia• Cass.

Erechtites Raf. Garcibarrigoa Cuatrec.

Gynoxys Cass.

Lnsiocephalus Willd. ex Schltdl.

Monticalia• C. Jeffrey Pentacalia Cass. [include Monticalia] Paragt;noxys (Cuatrec.) Cuatrec.

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