SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NCTMBER 52 

Chromosome Numbers in Compositae, 

XII: Heliantheae 

Harold Robinson, 

A. Michael Powell, 

Robert M. King, 

andJames F. Weedin 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1981 
ABSTRACT 

Robinson, Harold, A. Michael Powell, Robert M. King, and James F. Weedin. 

Chromosome Numbers in Compositae, XII: Heliantheae. Smithsonian Contributions to Botany, number 52, 28 pages, 3 tables, 1981.-Chromosome reports 

are provided for 145 populations, including first reports for 33 species and 

three genera, Garcilassa, Riencourtia, and Helianthopsis. Chromosome numbers 

are arranged according to Robinson’s recently broadened concept of the 

Heliantheae, with citations for 212 of the ca. 265 genera and 32 of the 35 

subtribes. Diverse elements, including the Ambrosieae, typical Heliantheae, 

most Helenieae, the Tegeteae, and genera such as Arnica from the Senecioneae, 

are seen to share a specialized cytological history involving polyploid ancestry. 

The authors disagree with one another regarding the point at which such 

polyploidy occurred and on whether subtribes lacking higher numbers, such 

as the Galinsoginae, share the polyploid ancestry. Numerous examples of 

aneuploid decrease, secondary polyploidy, and some secondary aneuploid 

decreases are cited. The Marshalliinae are considered remote from other 

subtribes and close to the Inuleae. Evidence from related tribes favors an 

ultimate base of X = 10 for the Heliantheae and at least the subfamily 

As teroideae. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf clearing from the 

katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Main entry under title: 

Chromosome numbers in Compositae, XII. 

(Smithsonian contributions to botany ; no. 52) 

Bibliography: p. 

Supt. of Docs. no.: SI 1.29:52 

1. Compositae-Genetics. 2. Chromosomes. 3. Plant cytotaxononiy. I. Robinson, Harold 

QKl.S2747 no. 52 [QK495.C74] 581 81-607855 [583’.55] AACR:! 

Ernest, 1932-. 11. Title: Heliantheae. 111. Series. 
Contents 

Introduction ......................................................... 

Acknowledgments ................................................. 

General Considerations ............................................ 

Chromosome History of the Tribe .................................... 

Ultimate Base Number of the Tribe .................................. 

Review of the Subtribes .............................................. 

AMBROSIINAE ............................... 

ESPELETIINAE ...................................................... 

MELAMPODIINAE .................................................. 

POLYMNIINAE ...................................................... 

MILLERIINAE . . ................................................. 

DESMANTHODIINAE ................................................. 

CLIBADIINAE ....................................................... 

GUPlRDIOLINAE ..................................................... 

ENHYDRINAE ...................................................... 

MONTANOINAE ..................................................... 

RUDBECKIINAE .................................................... 

ZALUZANIINAE .................................................... 

ECLIPTINAE ........................................................ 

NEUROLAENINAE .................................. 

HELIANTHINAE .................................................... 

GALINSOGINAE .................................................... 

DIMERESIINAE .................................................... 

COREOPSIDINAE .................................................... 

FITCHIINAE ......................................................... 

COULTERELLINAE ................................................... 

PECTIDINAE ........................................................ 

FLAVERIINAE ...................................................... 

VARILLINAE ....................................................... 

CLAPPIINAE ........................................................ 

MADIINAE ......................................................... 

HYMENOPAPPINAE .................................................. 

PERITYLINAE ...................................................... 

BAERIINAE ......................................................... 

CHAENACTIDINAE ................................................... 

GAILLARDIINAE .................................................... 

MARSHALLIINAE ................................................... 

JALMEINAE ......................................................... 

Literature Cited ....................... 

Page 

1 

5 

5 

7 

9 

10 

10 

10 

17 

18 

18 

18 

18 

18 

18 

19 

19 

19 

19 

20 

20 

21 

21 

22 

22 

22 

22 

23 

23 

23 

23 

23 

23 

23 

24 

24 

24 

24 

25 

... 

111 

Chromosome Numbers in Compositae, 

XII: Heliantheae 

Harold Rob inson, 

A. Michael Powell, 

Robert M. King, 

and James F. Weedin 

Introduction 

The present paper continues a series dealing 

with chromosome numbers of Compositae 

(Raven et al., 1960; Raven and Kyhos, 1961; 

Ornduff et al., 1963, 1967; Payne et al., 1964; 

Solbrig et al., 1964, 1969, 1972; Anderson et al., 

1974; Powell et al., 1974, 1975; King et al., 1976; 

Tomb et al., 1978) and is the first using a revised 

and broadened concept of the Heliantheae (Robinson, 1981). Reports are provided for 145 populations including first reports for 33 species and 

three genera, Garciiassa, Riencourtia, and Helianthopsis (marked by asterisks in Table l), and the 

cytological history of the tribe is discussed. 

The new reports in this paper are based on 

material collected by R. M. King and counted by 

A. M. Powell and J. F. Weedin. The chromosome 

counts have been made from acetocarmine or 

aceto-orceine squashes of microsporocytes in 

meiosis. Voucher specimens of the King collections are in the U.S. National Herbarium (US), 

Harold Robinson and Robert M. King, Department of Botany, 

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. A. Michael Powell and James F. Weedin, 

Department of Biology, Sul Ross State University, Alpine, Texas 

79830. 

with a nearly complete set at the Missouri Botanical Garden (MO). 

The treatment of the tribe by Robinson (1981), 

used as the basis for comparison in this paper, 

recognizes 265 genera containing approximately 

2970 species distributed in 35 subtribes. Included 

are elements that have been considered as separate tribes beginning with the Ambrosiinae followed by the typical Heliantheae in the paleaceous series. The system continues with Dimeresza, 

which is sometimes placed in the Inuleae, the 

Coreopsidinae and Pectidinae, which have recently been treated as tribes, and the epaleaceous 

series, which includes many genera from the traditional Senecioneae and most of the Helenieae. 

The isolated subtribe Marshalliinae is treated 

last. The tribe therefore is considerably altered 

from the traditional versions of Bentham and 

Hooker (1873) and Hoffmann (1890-1894) and 

from the recent revised treatment by Stuessy 

(1977), though the latter has proved to be a useful 

source of data. The tribe is considered somewhat 

specialized in the family by Robinson (1981), in 

contrast to the central phyletic position proposed 

by Cronquist (1955). The morphological, anatomical, and chemical characters that serve as a 

basis for Robinson’s treatment are reviewed in 

that paper (198 1). 

1 
2 SMITHSONIAN COKTRIBUTIOKS TO BOTANY 

TABLE 1 .-Chromosome counts, localities, and vouchers for species of Heliantheae (asterisk 

(*) = first report) 

Species 

Acanthospermum hispidum DC. 

Alloispermum integrlfolium (DC.) 

H. Robins. 

* Alloispermum scabrum (Lag.) H. Robins. 

* Alloisperum scabrum (Lag.) H. Robins. 

Ambrosia artemrsiordes (W’illd.) Meyen & 

Baltimora recta L. 

Bidens andicola H.B.K. 

Bidens andicola H.B.K. 

Bidens andicola H.B.K. 

Bidens andicola H.B.K. 

Bidens andicola H.B.K. 

Bidens andicola H.B.K. 

Bidens andicola H.B.K. 

Bidens chiapensis Brandegee 

Bidens ostruthioides (DC.) Sch.Bip. 

Bidens pilosa L. 

Bidens squarrosa H.B.K. 

Bidens squarrosa H.B.K. 

Walp. 

Bidens sp. 

Bidens sp. 

Clibadium anceps Greenm. 

Clibadium armanii Sch.Bip. ex Baker 

Clibadium armanii Sch.Bip. ex Baker 

Clibadium armanii Sch.Bip. ex Baker 

Clibadium glomeratum Greenm. 

Clibadium glomeratum Greenm. 

* Cltbadium leiocarpum Steetz in Seem. 

* Clibadium leiocarpum Steetz in Seem. 

* Clibadium microcephalum Blake 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Clibadium surinamense L. 

Coreopsis lanceolata L. 

Coreopsis mutica DC. 

Coreopsis mutzca DC. 

Cosmos caudatus H.B.K. 

Cosmos caudatus H.B.K. 

n Localib and collection number 

11 

16 

17-18 

ca. 16 

18 

12- 13+2-4” 

24 

12 

12 

12 

12 

12 

23 

12 

36 

24 

24 

ca. 

24 

34-36 

16 

16 

16 

16 

16 

16 

16 

ca. 17 

16 

16 

16 

16 

16 

16 

16-17 

16 + frag.‘ 

17-20 

16 

13 

28 

28 

ca. 24 

24 

Bolivia: La Paz. 19 km from Chulumani, K7450 

Guatemala: Chimaltenango. 8 km WSW of Chimaltenango, 

Guatemala: Alta Verapaz, 6 km W of San Cristobal \.’eraGuatemala: Quezaltenango, 20 km of Quezaltenango. 

Ecuador: Pichincha, 20 km S of Quito. K6615 

K7196 

paz. K7113 

K7256 

Guatemala: Escuintla, Escuintla, K7017 

Bolivia: La Paz, 24 km from La Paz, K7411 

Bolivia: La Paz. 30 km from La Paz. K7413 

Bolivia: La Paz, 67 km from Chulumani, K7473 

Bolivia: La Paz. 32 km from La Paz, K7498 

Bolivia: Oruro, 32 km from Oruro, K7518 

Ecuador: Azuay, 6 km SE of Cumbe, K6694 

Ecuador: Chimborazo, 1 km W of San Juan, K6597 

Guatemala: Chimaltenango, 17 km N of Tecpan, K7075 

Guatemala: Baja Verapaz, 3 km NE of Granados, K7086 

Ecuador: Azuay, 3 km E of Paute, K665 1 

Guatemala: Sacatepequez. 1 I km N of Escuintla, K7176 

Guatemala: Alta L’erapaz. 15 km W of San Cristobal VeraBolivia: Cochabamba. near Parotani. K7572 

Bolivia: Cochabamba, 26 km from Tolata, K7601 

Costa Rica: Heredia. 2 km SW of Alto Gallito, K6805 

Brasil: Bahia, 12 km from Ilheus, K8008 

Brasil: Distrito Federal, 32 km N of bridge at Asa Norte, 

Brasil: Goias, 21 km N of .41to Paraiso, K8286 

Costa Rica: Cartago, El Munieco. K6766 

Costa Rica: San Jose, 2-3 km N of Nubes, K6778 

Costa Rica: San Jose. 16 km W of Santa .4na, K6746 

Costa Rica: Cartago. 3 km S of Pacayas. K683 1 

Ecuador: Canar, ca. 40 km E of bridge at Guayaquil, K7735 

Bolivia: La Paz. 1 km from Chulumani, K7423 

Bolivia: La Paz, 7 km from Chulumani, K7433 

Bolivia: La Paz, 3 km from Chulumani, K7440 

Bolivia: La Paz. 52 km from Chulumani. K7461 

Bolivia: Cochabamba. 49 km from Colomi, K7701 

Costa Rica: San Jose, 5 km NE of Santiago-Puriscal, K6750 

Ecuador: Caniar, 77 km ESE of Guayaquil, K6871 

Ecuador: Guayas, along road to Naranjal, K7005 

Ecuador: Loja, 24 km S of Catamayo. K795l 

United States: Maryland, Prince Georges Co.. K7389 

Guatemala: Solola, 13 km SE of Panajachel, K7231 

Guatemala: Alta Verapaz, 5 km WNW of San Cristobal 

Bolivia: La Paz, 19 km from Chulumani, K7453 

Ecuador: Guayas, along road to Vinces, K7009 

paz, K7344 

K8205 

\’erapaz. K7338 
NUMBER 52 3 

TABLE 1.-Continued 

Species 

* Cosmos peucedanthifolius Wedd. 

Echinacea tennessensis (Beadle) Small 

Eclipta alba (L.) Hassk. 

Flaveria bidentis Kuntze 

* Flourensia heterolepis Blake 

* Flourensia heterolepis Blake 

* Galinsoga mandonii Sch.Bip. 

* Garcilassa rivularis Poepp. & Endl. 

* Helianthopsis hypargyreus (Blake) H. 

* Helianthopsis hypargyreus (Blake) H. 

* Helianthopsis pseudoverbesinoides (Hieron.) 

* Helianthopsis lehmannii (Hieron.) H. 

Galinsoga quadriradiata Ruiz & Pavon 

Helenium mexicanum H.B.K. 

Robins. 

Robins. 

H. Robins. 

Robins. 

Hymenoxys robusta (Rusby) K. Parker 

Kinginnthus paniculatus (Turcz.) H. 

Kingianthus paniculatus (Turcz.) H. 

Lasianthaea fruticosa (L.) Becker 

Melampodium americanum L. 

Melampodium paniculatum Gardn. 

Melampodium perfoliatum (Cav.) H.B.K. 

Monactis kingii H. Robins. 

Neurolaena lobata (L.) R. Br. 

Neurolaena lobata (L.) R. Br. 

Robins. 

Robins. 

* Monactis holwayae (Blake) H. Robins. 

* Oyadaea boliviana Britton 

* Oyadea boliviana Britton 

* Perymenium gymnolomoides (Less.) DC. 

* Perymenium nicaraguense Blake 

Podachaenium eminens (Lag.) Sch.Bip. 

Podachaenium eminens (Lag.) Sch.Bip. 

Parthenium hysterophorus L. 

Podachaenium eminens (Lag.) Sch.Bip. 

Podachaenium eminens (Lag.) Sch.Bip. 

* Podachaenium skutchiz (Blake) H. Robins. 

* Podachaenium skutchii (Blake) H. Robins. 

Porophyllum ruderale Uacq.) Cass. 

Porophyllum ruderale Uacq.) Cass. 

Schistocarpha eupatorioides (Fenzl) 

Schistocarpha eupatorioides (Fenzl) 

* Riencourtia tenuiflia Gardn. 

Kuntze 

Kuntze 

n Locality and collection number 

12-14 

11 

11 

ca. 18 

ca. 18 

ca. 18 

24 

16 

17+ld 

12-13 

17 

9 

17 

ca. 17 

15 

ca. 30 

32-34 

ca. 8 

10 

18 

11 

32 + ca. 6' 

11 

11 

14 

14 

ca. 18 

140r 15 

15-17 

18 

19 

ca. 19 

19 

19 

19 

34-36 

22-24 

ca. 32' 

ca. 16 

a 

8 

Bolivia: Cochabamba, 46 km from Challa, K7543 

United States: iMaryland, Prince Georges Co., cult., K6849 

Costa Rica: Cartago, El Munieco, K6765 

Ecuador: Loja, 16 km W of Catamayo, K6925 

Bolivia: Cochabamba, 1 1 km from Quillacolla~ Ki559 

Bolivia: Cochabamba, 19 km from Tolata, K7585 

Bolivia: La Paz, 1 km from Chulumani, K7419 

Ecuador: Loja, 7 km S of Saraguro, K7830 

Ecuador: Chimborazo, 35 km NE of El Triunfo, K6954 

Guatemala: Baja Verapaz, 32 km SE of Salama, K7094 

Ecuador: Tungurahua, 25 km SE of Ambato, K6577 

Ecuador: Azuay, 16 km N of Sigsig, K6646 

Ecuador: Tungurahua, 1 km SE of Ambato, K6533 

Ecuador: Chimborazo, 25 km SW of Riobamba, K6970 

Bolivia: Oruro, 32 km from Oruro, K7522 

Ecuador: Pichincha, 3 km SW of Chillogallo, K6509 

Ecuador: Tungurahua, 11 km S of Ambato, K6578 

Costa Rica: Cartago, 8 km W of Tirro, K6843 

Guatemala: Baja Verapaz, 21 km SW of Granados, K7089 

Guatemala: Escuintla, ca. 19 km NW of Escuintla, K7014 

Costa Rica: San Jose, San Jose, K6745 

Ecuador: Azuay, 6 km SE of Cumbe, K6697 

Ecuador: Canar, ca. 25 km N of Canar, K7739 

Ecuador: Canar, 23 km ESE of El Triunfo, K6998 

Guatemala: Sacatepequez, 7 km N of Escuintla, K7174 

Bolivia: La Paz, 23 km from Chulumani, K7455 

Bolivia: La Paz, 52 km from Chulumani, K7464 

Bolivia: Cochabamba, 1 1 km from Quillacolla, K7564 

Guatemala: Solola, 16 km S of Chichicastenango, K7065 

Guatemala: Jalapa, 8 km NE of Jalapa, K7137 

Costa Rica: Cartago, 8 km W of Tirro, K6842 

Guatemala: Alta Verapaz, 12 km W of San Cristobal VeraGuatemala: Suchitepequez, 24 km N of Patubal, K7236 

Mexico: Oaxaca, Cerro San Filipe, K6445 

Guatemala: Quezaltenango, 20 km S of Quezaltenango, 

Guatemala: Totonicapan, 14 km NE of Totonicapan, K7279 

Ecuador: Loja, 16 km W of Catamayo, K6927 

Ecuador: El Oro, 13 km N of Machala, K7976 

Brazil: Goias, 20 km N of Alto Paraiso de Goias, K8808 

Bolivia: La Paz, 8 km from Chulumani, K7436 

paz, K7119 

K7257 

Ecuador: Tungurahua, 11 km E of Banos, K6548 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1.-Continued 

Species 

Schistocarpha eupatoriotdes (Fenzl) 

* Schistocarpha longiligula Rydberg 

* Schistocarpha paniculata Klatt 

Kuntze 

Schkurhia multtflora Hook. & Am. 

Schkuhria multtylora Hook. & Am. 

Schkuhrza pinnata (Lam.) Kuntze 

Sigesbeckia jorullensis H.B.K. 

Simsia dombeyana DC. 

Simsia holwayi Blake 

* Smallanthus fruticosa (Benth.) H. Robins. 

Smallanthus maculatus (Cav.) H. Robins. 

Smallanthus maculatus (Cav.) H. Robins. 

* Smallanthus microcephala (Hieron.) H. 

Robins 

Smallanthus riparius (H.B.K.) H. Robins. 

Spilanthus alba L’Her. 

Spilanthes alba L’Her. 

Spilanthes alba L’Her. 

Spilanthes oppositiyolia (Lam.) D’Arcy 

* Steiractinia mollis Blake 

Tagetes terniyolia H.B.K. 

Tagetes zypaquirensis H. B. K. 

Tagetes zypaquirensis H.B.K. 

Tagetes zjpaquirensis H.B.K. 

Tithonia diuersifolia (Hemsl.) A. Gray 

Tithonia rotundzfolza (Miller) Blake 

Tithonia tubaeformis (Jacq.) Cass. 

Tithonia tubaeformis (Jacq.) Cass. 

Verbesina apleura Blake 

* Verbesina boliiiana Klatt 

* Verbesina bolitiana Klatt 

* Verbesina egsersii Hieron. 

Verbesina encelioides (Cav.) Benth. & 

Verbestna gtgantea Jacq. 

Verbesina glabrata Hook. & Arn. 

Verbesina lindenii (Sch.Bip.) Blake 

Hook. f. 

* Verbesina latisquamata Blake 

* Verbesina minuticeps Blake 

* Verbesina oerstediana Benth. 

* Verbesina oerstediana Benth. 

* Verbesina pentantha Blake 

* Verbesina pentantha Blake 

* Verbesina semidecurrens Kuntze 

* Verbestna semidecurrens Kuntze 

* Verbestna sodiroi Hieron. 

* Verbesina sodiroi Hieron. 

Verbesina turbacensis H.B.K. 

n Locality and collection number 

8 

8 

8 

12 

12 

10 

15 

17 

17 

29+ 1 

ca. 34 

ca. 16 

27-30 

16 

30-35 

ca. 39 

ca. 39 

ca. 26 

14 

ca. 24 

12 

12 

12 

17 

17 

ca. 17 

17 

17 

ca. 17 

17 

17 

17 

17 

ca. 34 

ca. 34 

17 

17 + 1 frag. 

ca. 17 

15-17 

34 

34 

17 

17 

34 

34 

17 

Ecuador: El Oro, 30 km NE of Machala, K6932 

Guatemala: Alta Verapaz, 3 km ENE of Coban, K7326 

Costa Rica: Cartago, 6 km NE of San Rafael de Irazu, 

Bolivia: Cochabamba, 11 krn from Quillacolla, K7566 

Bolivia: Cochabarnba, 19 km from Cochabamba, K7667 

Ecuador: Loja, 16 km W of Catamayo, K6926 

Ecuador: Azuay, 7 km SE of Cumbe, K6707 

Ecuador: Guayas, 17 km E of Palestina, K7008 

Guatemala: Alta Verapaz, 5 km WNW of San Cristobal 

Ecuador: Chimborazo, 25 krn SW of Riobamba, K6971 

Costa Rica: Cartago, slopes of Volcan Irazu, K6810 

Costa Rica: San Jose, 5 km NE of Santiago-Puriscal, K6749 

Ecuador: Azuay, 23 km SW of Giron, K6691 

K6826 

Verapaz, K7337 

Bolivia: La Paz, 1 krn from Chulurnani, K7423 

Guatemala: Jutiapa, N of Asuncion Mita, K7386 

Ecuador: Canar, 67 km W of Caniar, K7794 

Ecuador: El Oro, 5 km W of Pinias, K7970 

Costa Rica: Cartago, El Munieco, K6769 

Ecuador: Chimborazo, 38 km SW of Riobamba, K6973 

Ecuador: Tungurahua, 39 km SSW of Banos, K6990 

Ecuador: Pichincha, 3 krn SW of Chillogallo, K6507 

Ecuador: Pichincha, 1 krn W of Aloag, K6522 

Ecuador: Chirnborazo, 25 km SW of Riobamba, K6969 

Costa Rica: Heredia, 2 km N of San Isidro, K6993 

Guatemala: Suchitepequez, 10 km E of Mazatenango, 

Guatemala: Guatemala, 8 km N of Villa Canales, K7084 

Guatemala: Huehuetenango, 16 km E of Chiantla, K7316 

Guatemala: San Marcos, 20 km NE of San Marcos, K7039 

Bolivia: Cochabamba, 11 km from Quillacolla, K7558 

Bolivia: Cochabamba, 1 1 km from Quillacolla, K7563 

Ecuador: Guayas, vicinity of Guayaquil Airport, K6944 

Bolivia: Santa Cruz, Comarapa, K7620 

K7243 

Guatemala: Escuintla, 16 km SW of Amatitlan, K7168 

Bolivia: Santa Cruz, 2 km from Comarapa, K7636 

Ecuador: Canar, 20 km NW of Azogues, K6884 

Mexico: Hidalgo, Jalapa, K6492 

Ecuador: Chimborazo, 12 km NE of Bucay, K6991 

Costa Rica: San Jose, 8 km N of Nubes, K6779 

Costa Rica: Cartago, 3 km NE of San Rafael de Irazu, 

Ecuador: Loja, 4 km W of Loja, K6923 

Ecuador: Loja, 7 km W of Loja, K7882 

Bolivia: Cochabamba, 11 km from Quillacolla, K7561 

Bolivia: Cochabarnba, 9 km from Parotani, K7575 

Ecuador: Pichincha, 20 km S of Quito, K6514 

Ecuador: Chimborazo, 1 km W of San Juan, K6594 

Costa Rica: San Jose, 22 km S of Cartago, K6753 

K68 19 
NUMBER 52 5 

TABLE 1.-Continued 

Species 

Verbesina turbacensis H.B.K. 

Viguiera cordata (Hook. & Am.) D’Arcy 

Viguzera cordata (Hook & Am.) D’Arcy 

* Viguiera lanceolatu Britton 

* Viguiera parensis Rusby 

* Viguiera parensis Rusby 

* Viguiera procumbens (Pers.) Blake 

* Viguiera procumbens (Pers.) Blake 

* Viguiera procumbens (Pers.) Blake 

* Viguiera rudbeckioides (H.B.K.) H 

* Wedelia grandflora Benth. 

Robins. 

Wedelia helianthoides H.B.K. 

Wedelia helianthoides H.B.K. 

Wedelia helianthoides H.B.K. 

* Wedelia holwavi Blake 

n LocaliQ and collection number 

ca. 17 

ca. 40 

40 

ca. 34 

ca. 34 

ca. 34 

ca. 34 

ca. 34 

17 

17 or 18‘ 

Guatemala: Baja Verapaz, city limits of Purulha, K7100 

Guatemala: Suchitepequez, 3 km W of Cocales, K7239 

Guatemala: Suchitepequez, 30 km \V of Mazatenango, 

Bolivia: La Paz, 3 km from Chulumani, K7427 

Bolivia: La Paz. 69 km from Chulumani. K7477 

Bolivia: Cochabamba, 1 km from Quillacolla, K7552 

Bolivia: Oruro, 20 km from Caracollo, K7526 

Bolivia: Cochabamba, 46 km from Challa. K7542 

Bolivia: Cochabamba, 26 km from Tolata, K7602 

Ecuador. Loja, 14 km S of Loja, K7862 

K7248 

12 

ca. 21 

ca. 22 

20-22 

11 

Ecuador: Loja, 18 km N of Vilcabamba, K7870 

Ecuador: Azuay, 10 km SW of Azogues, K6613 

Ecuador: Caniar. 7 km SM’ of Azogues, K6880 

Ecuador: Guayas, \vicinity of Guayaquil Airport. K6946 

Bolivia: Cochabamba, 5 km from Parotani, K7573 

a 12-13 + 2-4 univalents or supernumery chromosomes. ’ ca. 12 (possibly n=ll). 

‘ 16 + 2(4)6 fragment. 

1711+11 or n=18. 

32 + ca. 6 ghost chromosome or n+3011+11~. 

ca. 32 (30-34). 

17 or 18 (probably 17). 

It should be emphasized that the revised classification has been used as a basis for organizing 

the data in the present study, but it is of critical 

significance only in the discussions of various 

subtribes. The general conclusions are valid in 

comparison to any of the modern systems of 

classification, and one of the major conclusions, 

acceptance of higher base numbers for most of 

the subtribes of the Heliantheae, was anticipated 

by Smith (1975) on the basis of traditional subtribal concepts. 

following grants to Robert M. King: National 

Science Foundation BMS 70-00537, Penrose 

Fund of the American Philosophical Society, and 

the National Geographic Society; and the following grant to A. M. Powell: National Science 

Foundation BMS 73-06851 A01. The help of Dr. 

J. J. Wurdack in furnishing material of Echinacea 

tennesseensis from his garden is appreciated. We 

wish to thank Drs. David Lellinger and Vicki 

Funk for reviewing the manuscript. We credit a 

ACKNOWLEDGMENTS.-Supported in part by the 

previous, unnamed reviewer for providing a chart 

of chromosome number frequencies after which 

Table 2 is patterned. 

General Considerations 

Many genera and subtribes of the Heliantheae 

show an extreme diversity of chromosome numbers (Tables 2 and 3; Strother, 1977; Stuessy, 

1977; Turner and Powell, 1977b). Variation is 

particularly marked among groups having n less 

than 10. Stability, where it occurs, is usually 

found where n=17-19. Much diversity in numbers occurs within groups that are obviously 

closely related, and such diversity of chromosome 

number is not indicative of phyletic diversity. In 

contrast, diverse elements of the tribe tend to 

have uniformity of chromosome numbers at levels 

of n=17-19. 

Examination of the Heliantheae seems to indicate that every possible process of karyotype 

evolution is involved in producing the chromo- 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

soma1 variation. As in other groups of plants, the 

chromosome numbers of the Heliantheae tend to 

correlate with the habit of the plant. The trend 

toward reduced DNA content in weedy, shortlived, herbaceous species (Bennett, 1972; King et 

al., 1976; Solbrig, 1977) is seen in the subtribes 

Galinsoginae, Dimeresiinae, Madiinae, and 

Chaenactidinae, among others. In some groups 

such as the Madiinae, the more herbaceous members of the subtribe in North America have lower 

chromosome numbers, whereas the more woody 

members in Hawaii show higher base numbers. 

The highest base numbers in the tribe are associated with more dendroid elements, such as the 

Espeletiinae, Montanoa of the Montanoinae, and 

Podachaenium of the Ecliptinae. 

Polyploidy is apparently the most common 

evolutionary process in the tribe. Examples include series in the Galinsoginae on a base of n=8, 

Chaenactis on a base of n=6, many Coreopsidinae 

and Pectis of the Pectidinae on a base of n=12, 

Spifanthes (Acmelfa) on base of n=13, members of 

the Ambrosiinae on a base of n=18, and species 

of Arnica on a base of n= 19. Polyploids of Montanoa 

having a base of n=19 reach n=l lO-120, the 

highest count known for the Asteraceae (Funk 

and Raven, 1980). It is clear that polyploid populations and individuals commonly are produced 

in the tribe and that they commonly survive. 

Aneuploid increase, in contrast to polyploidy, 

seems to be rare in the Heliantheae. Examples 

undoubtedly exist, but numbers such as 11 or 12 

are not assumed to represent simple increases 

from n=9 or 10 in this survey. Even the example 

of increase to n= 11 and 12 from a base of n= 10 

in Melampodium (Stuessy, 1979) cannot of considered proven, since that study assumed primitiveness of lower numbers in the tribe (Stuessy, 1977; 

see also the discussion of Melampodiinae below). 

There is no detailed evidence on the cytological 

mechanisms involved, but aneuploid increase 

should not be any more difficult to produce than 

some forms of aneuploid decrease. The primary 

difference must be in the ability of the aneuploid 

increase to survive and stabilize. Any extra chromosome in an out-crossing plant would tend to 

be unpaired and possibly eliminated in a subsequent generation. Most processes that increase 

chromosome number involve production of superfluous genetic material, and in the absence of 

other factors, elimination and simplification normally are favored. 

Aneuploid decrease is common in the Heliantheae, and a number of unquestionably long 

series are evident. Either loss of superfluous chromosomes or reduction of centromeres through 

translocation could easily explain descending series from higher numbers such as n=18 exemplified by Perityfe. Still, examples of reduction in 

other tribes, as in Fleischmannia in the Eupatorieae 

(Baker, 1967) and Crepis in the Lactuceae (Tobgy, 

1943), indicate that extreme reductions correlated 

with annual habit are mostly by rearrangement 

of chromosome material on fewer centromeres 

rather than by loss. Such a mechanism seems to 

occur in the Heliantheae in Lasthenia (Ornduff, 

1966) and Pseudobahia of the Baeriinae, Sabazia in 

the Galinsoginae (Solbrig et al., 1972), Calycadenia 

and Hofocarpha in the Madiinae (Carlquist, 1959), 

and in two species of Chaenactis in the Chaenactidinae (Raven and Kyhos, 1961). Evidence from 

polyhaploidy might be an indication that rearrangement of chromosome material on fewer centromeres has occurred in members of the tribe 

with higher chromosome numbers. 

Polyhaploidy may be a more common process 

than is generally realized. Its discovery depends 

on demonstrating that the ancestor was a polyploid, and this is usually not evident. When 

groups show more than one ploidy level, the 

higher levels ordinarily are considered derivatives 

of the lower ones. An apparent example in the 

Heliantheae seems to be the n=9 cited for Flaueria 

campestris Johnston (Anderson, 1973) in a subtribe 

of three genera, which otherwise show X= 18 consistently. A related process seems to give the X=8 

and n=9 of Heliomerzs and the n=9 in Helianthopsis 

hypargyrea (Blake) H. Robins., both in the subtribe 

Helianthinae having a base of X= 17. If our suppositions on the chromosomal history of the tribe 

are correct, many other lower numbers in the 

tribe may represent products of polyhaploidy. 
NUMBER 52 7 

Apomixis seems to be comparatively rare in the 

Heliantheae. The most obvious examples are the 

generic pair Monactis-Kingianthus, with n = ca. 30- 

34. Monactis dubia H.B.K. has been erroneously 

considered dioicous and has been placed in a 

separate genus, Astemma Less., because of the 

defective anthers of the type specimen (Robinson, 

1976). 

Allopolyploidy is sometimes cited as the basis 

for chromosome variability or the basis for groups 

having obscure origins in the Heliantheae. In a 

recent example, Gardner (1977) cites a hybrid 

between elements of the genus Lipochaeta with 

X=15 and X=26 and suggests derivation of typical Lipochaeta with X=26 from a hybrid between 

elements having X=l 1 and X= 15 (see also Rabakonandriana and Caro, 1981). Such a mechanism could explain many of the otherwise unexplained chromosome numbers in the tribe. Gardner’s suggestion presumes that barriers to hybridization were not complete between plants that 

had achieved various karyotype differences. In 

this way, variation in a group, once present, could 

generate more variation. The process would be 

accentuated by any general reduction in the effectiveness of isolating mechanisms in the tribe. 

Intergeneric hybridization is evident in the 

tribe. The sterile or nearly sterile natural hybrid 

cited by Gardner (1977) in his study of Lipochaeta 

was between elements not considered to be congeneric in this paper. Numerous experimental 

crosses have been attempted by one of the authors 

(Powell, 1972), and some sterile intergeneric hybrids have been produced. Rare but significant 

instances of fertile intergeneric hybrids are suspected on the basis of erratically distributed structural features in the Heliantheae (Robinson, 

1981). Similar evidence is found in other tribes 

that have been examined critically, such as the 

Eupatorieae (Robinson and King, 1977) and the 

Senecioneae (Robinson and Brettell, 1974). 

Chromosome History of the Tribe 

The treatment of the Heliantheae in the recent 

Symposium on the Biology and Chemistry of the 

Compositae (Steussy, 1977), along with an earlier 

paper on the subtribe Melampodiinae by the 

same author (Steussy, 1973), are particularly notable because they most strongly represent the 

recent trend to give the chromosome number 

primacy in resolving relationships. This is most 

evident in the suggested overall phylogeny of the 

tribe in which Stuessy (1977) says, “The first and 

largest line is that centering around the Verbesininae with chromosome numbers based on X= 15, 

16, and 17 (and presumptive aneuploid derivatives). The second line is that centering around 

the Galinsoginae with chromosomal bases of X=8 

and 9. The third and smallest line is that represented by the Coreopsidinae and its offshoot, the 

Fitchiinae, with base numbers of X=12 and 16.” 

The schematic diagram provided by Stuessy 

places the Helianthinae in his first group, while 

the Galinsoginae, which have proved to be closely 

related, on the basis of anatomy and chemistry 

(Robinson, 1981), are in the second group. The 

Verbesininae and Ecliptinae are in Stuessy’s first 

group, but the intimately related Engelmanninae 

are in the second. The Melampodiinae are placed 

in the first group although n= 11 and adjacent 

numbers are common in the subtribe and although n=9, 10, 11, 12, 18, 20, and 23 occur in 

Melampodium itself. The Bahiinae from the Helenieae are included in the second group, although 

the base numbers are mostly X=ll and 12 or 

higher. As such, Stuessy’s proposed phylogeny is 

internally inconsistent and seems to imply considerable chromosome number stability in a tribe 

where extreme variability is a most undeniable 

fact . 

In a study of the chromosome numbers of 

Coreopsis, which has implications for the whole 

tribe, Smith (1975) suggests a generic ancestor 

with X= 14 and separate aneuploid reductions to 

various groups with n=12 and 13. Smith further 

suggests derivation of Coreopsis from plants with 

chromosome numbers one to several aneuploid 

steps higher than Coreopsis. A high base number 

of X= 17- 19 is suggested for the tribe on the basis 

of the observations on the Coreopsidinae and the 

high numbers of other genera such as Helianthus, 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Verbesina, and Rudbeckia. The Smith proposal furnishes a remarkable contrast to the chromosomal 

history proposed by Stuessy (1977) and runs 

counter to all previous suggestions of simple derivations of higher numbers in the tribe from 

lower base numbers. 

In spite of the great variability in the chromosome numbers of the Heliantheae, a pattern can 

be seen when the evidence is fully reviewed. A 

few groups in the tribe, such as Galinsoginae, 

Baeriinae, and the Engelmannia group of the Ecliptinae, show a conventional pattern of polyploidy, 

with potential lower base numbers well represented and with higher numbers that can be 

interpreted directly as polyploids of those numbers. For instance, when n= 18 occurs, it clearly is 

derived from a doubling of X=9 or a tripling of 

X=6 (in Chaenactzs of the Chaenactidinae). In 

most subtribes distributed throughout most of the 

tribe, however, there are few or no lower chromosome numbers, and what stability there is in 

the tribe seems to be concentrated at the higher 

chromosome numbers. Numbers above X= 10 often form patterns easily interpreted as decreasing 

aneuploid series. The pattern usually is confirmed 

when actual relationships of species are known, 

and such decreasing patterns in the Coreopsidinae led Smith (1975) to postulate a higher base 

number for the subtribe and tribe. The subtribes 

with high base numbers predominate throughout 

the paleaceous and epaleaceous groups of the 

Heliantheae, with only the isolated Marsallinae 

unquestionably excluded. Various groups with 

lower base numbers are few and mostly isolated 

in the tribe and are subject to conflicting interpretations by the authors of the present paper 

(see below). 

A more complete picture of the nature of higher 

base numbers can be seen by using the example 

of other tribes that seem to show similar chromosome number patterns at earlier developmental stages. In the Senecioneae, the senecionoid 

series retains mostly X=10 and X=20, with one 

small group having n=23. A decreasing aneuploid 

series with n=9 and 7 is seen in the divergent 

Blennospermatinae, however, and the cacalioid 

series with a base of X=30 has a decreasing 

aneuploid series reaching n=25 (Robins and Brettell, 1973c, 1974: Nordenstam, 1977). In the Vernonieae, paleotropical species placed in Vernonia 

have bases of X= 10 and 20 with some reductions 

to n=9, but the Western Hemisphere species have 

n=16, 17, and 18 as a base (Jones, 1977). In the 

Eupatorieae, the base of X= 10 is widespread with 

one apparent increasing series to n=l 1 and 12 

and two separate decreasing series to n=9. HowTABLE 2.-Frequency of chromosome numbers in the Heliantheae by genus (chromosome 

number on horizontal axis. number of genera on vertical axis) 
NUMBER 52 9 

ever, a large element in the tribe seems to have 

base numbers of 19, 18, or 17 with some reductions to 16 (King et al., 1976). All three of these 

tribes show that the predominant form of increase 

is polyploidy and that aneuploid series are mostly 

decreasing. The latter two tribes show further 

that elements of the tribes with stabilized higher 

numbers often have bases less than twice the 

amount of the original base number. It is not 

difficult to see the majority of the Heliantheae as 

elements like those in the Eupatorieae and Vernonieae, which have attained higher numbers 

through polyploidy and show subsequent aneuploid decrease. 

The case for higher base numbers in most 

subtribes of the Heliantheae seems clear, but the 

extent to which they share their polyploid ancestry is less obvious. Predominance of subtribes 

with higher base numbers leads Robinson and 

King, of the present authors, to consider that all 

the extant Heliantheae except the Marshalliinae 

are derived from a single ancestral stock with 

higher chromosome numbers and that there is a 

subsequent progression from stable higher numbers to increasingly unstable lower numbers, ending finally in either scattered recurrence of polyploidy or extreme reduction associated with the 

annual habit. There are a few examples where 

secondary polyploidy seems to be followed by a 

secondary aneuploid decrease. In contrast, Powell 

considers the higher numbers to be derived from 

a number of separate occurrences of polyploidy, 

with such subtribes as the Galinsoginae and Baeriinae having lacked polyploid ancestory. The 

authors agree that, considered in isolation, the 

internal evidence from the latter two subtribes 

favors an original base number for them of X= 10. 

Ultimate Base Number of the Tribe 

Even with a general acceptance of the higher 

base number of X=17-19 suggested by Smith 

(1975) for the Heliantheae, there is still the question of the ultimate base number from which the 

higher number is derived. In a previous paper of 

this series (Solbrig et al., 1972), a base of X=8-12 

was suggested for the Heliantheae, and in the 

review of the tribe by Stuessy (1977) an original 

X=8 or 9 was suggested. Such proposals did not 

take into account possible higher base numbers 

for most subtribes of the Heliantheae, and these 

suggested lower base numbers are given less credence here. Still, these suggestions to some extent 

were an attempt to bring the Heliantheae into 

conformity with other tribes of the Asteraceae, 

and this does seem to be the only profitable 

approach for a group in which the original base 

number has been so completely obscured by 

polyploidy. 

A review of the entire family indicates a base 

number of X=9 or X=lO. It is notable that n=9 

is the most common number in the family (Solbrig, 1977), and it has been proposed as the base 

number for the family by Raven (1975). However, other evidence for an n=9 base for the family 

is weak. Examples of the number in older tribes 

such as the Eupatorieae are easily recognized as 

reductions from n=10, and tribes such as the 

Astereae and Anthemideae without evidence of a 

higher base number are highly advanced tribes 

with extensive evidence of aneuploid decrease. 

We find that the wide occurrence of n=18 offers 

little support for X=9 in view of the already 

mentioned tendency of higher numbers to stabilize at levels less than double the original base 

number. 

The evidence for X=10 is very different. That 

base in the Vernonieae has been mentioned 

above, but it is more common in the subfamily 

Asteroideae, to which the Heliantheae belong. 

The tribes marked with the greatest stability in 

chromosome number, the E;upatorieae and Senecioneae, have nearly certain base numbers of 

X=lO. The Calenduleae have a probable X=lO 

(Norlindh, 1977), and the Inuleae, which seem 

closest to the Heliantheae, are partly based on 

X=10. There seems to be no evidence that is 

convincing for any alternative number. Further 

supporting evidence from within the Heliantheae 

is the frequent occurrence of the number X= 19, 

which would be more difficult to explain from an 

original base of X less than 10. 
10 SMITHSOKIAN CONTRIBUTIONS TO BOTANY 

There remains perhaps the most divergent view 

on tribal and family base numbers to consider, 

the proposal that genera in various groups with 

low chromosome numbers evolved at a very early 

time, essentially to the morphological level as 

expressed in the genera now, at the basic level of 

X=5 and/or 4 (Turner et al., 1961; Johnston and 

Turner, 1962; Turner, 1977). The concept derives 

from the frequent absence of the n=6, 7, or 8 in 

groups having both n=9 and numbers of n=5 or 

below. In those examples where the phylogeny is 

well known, however, the direction of the change 

seems to be from the higher to the lower numbers. 

The present evidence does not preclude that there 

are only four or five basically different chromosomal elements in the ancestral complement of 

the Heliantheae and the family Asteraceae, which 

would facilitate reduction to lower numbers, but 

it does not seem that the ancestral stock of the 

Asteraceae actually had so few chromosomes. 

Review of the Subtribes 

The following 32 subtribes are treated in the 

order given by Robinson (1981). Three subtribes 

are omitted for lack of chromosome data, the 

Pinillosinae and Heptanthinae of the Greater 

Antilles and the Lycapsinae known only from 

San Felix and San Ambrosio islands off the coast 

of Chile. Chromosome reports are known for 2 12 

of the approximately 265 recognized genera of 

the tribe (Table 3), but no attempt is made in 

the table to indicate the number of species of each 

genus that have been counted. This information 

is given in the text for some genera. 

AMBROSI1NAE.-In the previous paper on the 

Ambrosiinae (Payne et al., 1964), X=18 was regarded as the primitive chromosome number. 

Since that time, two genera have been transferred 

into the subtribe by Stuessy (1973). Seven of the 

eight genera in the subtribe have been reported. 

All have some counts of n=18, and both Zua and 

Ambrosia have multiples of that number. Reports 

of n=27 in Iva and Parthenium might indicate some 

traces of an original X=9, but the latter number 

apparently is not extant in any of the genera. A 

more likely origin for n= 27 in the subtribe would 

be hybridization between different ploidy levels 

of n=18. Lower numbers, including the n=12 

found in two genera, also seem secondarily derived from X= 18. 

Stuessy (1 973) hypothesized an Engelrnannialike ancestor for the Ambrosiinae, having a base 

number of X=9. Engelmannia, which is placed in 

the Ecliptinae in this treatment, has achene complexes reminiscent of some Ambrosiinae. The 

Ambrosiinae, along with the Espeletiinae and the 

Clibadiinae, have an unstriated achene wall that 

may indicate relationship to the Ecliptinae, and 

all four subtribes have at least some species with 

male disk flowers. In determining the base number of the Ambrosiinae, it seems significant that 

the related subtribes Epeletiinae and Clibadiinae 

have bases of X=19 and X=16 respectively, and 

that more primitive elements of the Ecliptinae, 

such as Podachaenzum, have n= 18-19. The genus 

Engelmannza is regarded here as a more specialized 

element in the Ecliptinae. 

ESPELETiih.AE.-This distinctive group of the 

paramo and subparamo of northern South America has recently been elevated to subtribal rank 

(Cuatrecasas, 1976). Seven genera presently are 

recognized, with chromosome reports for five of 

them (Powell and King, 1969; Powell and Cuatrecasas 1970, Powell and Powell, 1978). All reports are n=19, making the Espeletiinae one of 

the few subtribes of the Heliantheae without 

known variation. The recent proliferation of genera is based in part on striking differences in 

habits of the plants, but the consistent chromosome number and the comparatively recent origin 

of most paramo habitats would indicate that 

diversity within the subtribe is of rather recent 

origin. 

Neither structure nor chromosome number 

seems to link the Espeletiinae with the Melampodiinae in which it has traditionally been 

placed. Two other genera with arborescent habits 

and chromosome numbers of n=19 are Montanoa 

of the Montanoinae and Podachaenium of the 

Ecliptinae, but other characters indicate that 
NUMBER 52 11 

TABLE 3.--General distribution, habit, and chromosome number of genera of the Heliantheae 

(A = annual, B = biennial, P = perennial herb, sS = subshrub, S = shrub, L = liana, T = tree) 

Subtribe and genus 

Ambrosiinae Lessing 

Ambrosia 

Dicoria 

Hymenoclea 

Iua 

Parthenice 

Parthenium 

Xanthium 

Espeletiinae Cuatr. 

Carramboa 

Espeletia 

Espeletiopsis 

Libanothamnus 

Ruilopezia 

Acanthosperumum 

Melampodiinae Lessing 

Ichthyothere 

Lecocarpus 

Melampodium 

Smallanthus 

Polymniinae H. Robinson 

Milleriinae Benth.& Hook 

Polymnia 

Guizotia 

Milleria 

Rumfordia 

Sigesbeckth 

Trigonospermum 

Desmanthodiinae H. Robinson 

Desmanthodium 

Clibadiinae H. Robinson 

Cli badi um 

Rtencourtia 

Guardiolinae H. Robinson 

Gua rdi o la 

Enhydrinae H. Robinson 

Enhydra 

Montanoinae 

Montanoa 

General Approx. number Chromosome numbers 

distribution Habit of species (mostly haploid) 

North America, Mexico, Central 

America, Andes, adventive 

sw US, n Mexico 

sw US, n Mexico 

North America 

sw US, n Mexico 

North America, tropical America, 

North America, tropical America, 

adven tive 

adventive 

Venezuela 

n Andes 

Colombia, Venezuela 

Columbia, Venezuela 

Venezuela 

tropical and subtropical America 

Central and South America 

Galapagos 

tropical America 

adventive 

e North America, Mexico, Central 

and South America 

e North America 

tropical Africa, cultivated 

Mexico, Central and n South 

Mexico, Central America 

tropical America, Africa, Asia 

Mexico, Guatemala 

America 

Mexico, Central and n. South 

America 

tropical America 

South .4merica 

Mexico 

pantropical 

Mexico, Central and n South 

America 

30 

4 

3 

19 

1 

16 

2 

4 

51 

24 

13 

21 

6 

25 

3 

36 

20 

2 

6 

2 

6 

12 

3 

10 

40 

8 

10 

10 

33 

12, 17, 18, 36, 54, 62 or 

18 

18 

16, 17, 18, 27, 36 

18 

12, 17, 18, 27, 

2n=3511+ 11 

18 

63, 72 

19 

19 

19 

19 

19 

10, 11, 12, 20 

16. ca. 33 

11 

9, 10, 11, 12, 18, 20, 

23, 25f1, 27, 30, 33 

ca. 15, 16, 17, 18, 27- 

30, 30 

15 

15, ca. 30 

15 

24 

10, 12, 15, 30 

15 

18 

16, 17, 17-20, 24, 32 

ca. 16 

12 

11, 15 

19, 19f1, 57, 76, 110- 

120 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 3.-Conrinued 

Subtribe and genus 

Rudbeckiinae H. Robinson 

Dracopis 

Rati bida 

Rudbeckia 

Zaluzaniinae H. Robinson 

Hybridella 

Zaluzania 

Ecliptinae Lessing 

‘4spilia 

Balsumorrhiza 

Baltitnora 

Berlandiera 

Blaincillea 

Borrichia 

Ca lyp t oca rp us 

Chiysogonum 

Delilia 

Dugesia 

Echinacea 

Eclipta 

Eleutheranthera 

Encelia 

Encellopsis 

Engelmannia 

Flourensia 

Geraea 

Heliunthella 

HeliupJis 

King i an th u~ 

LaAianthaea 

Lindheimera 

Lipochaeia 

,Welanthera 

‘Uonactis 

Otopappus 

Philaclis 

Phoeban thus 

Podachaenium 

Podanihus 

General 

distribution 

Approx. number Chromosome numbers 

Habit of species (mostly haploid) 

s US 

US n Mexico 

North .4merica 

Mexico 

Mexico 

tropical America, ‘Africa 

North America 

tropical America 

sw US 

pantropical 

Gulf of Mexico, West Indies, Peru 

s US, Mexico. West Indies, Cene US 

tropical America 

Mexico 

e US 

pantropical 

tropical America, Madagascar 

w US, Mexico, w South .4merica. 

w us 

sw US 

tropical America 

sw US. Mexico 

w US. Mexico 

w US, Mexico, Central America. 

.Andes 

Ecuador 

Mexico, Central America 

sw US, Mexico 

Hawaii 

pantropical 

Ecuador, Peru 

Mexico. Central America, n South 

America 

Mexico, Central America, Andes 

Mexico, Central America, n South 

tral and e South America 

Argentina 

America 

s Mexico. Guatemala 

s us 

Mexico. Central America 

Chile. Argentina 

A 

P 

P 

P 

p, s 

p, s 

P 

A 

P 

A, p 

S 

A 

P 

A 

P 

P 

A 

A 

S 

P 

P 

S 

.A, P 

P 

P 

S 

S 

A 

S 

P. s, L 

S 

s, L 

p, s 

P, S, L. T 

S 

P 

s, T 

S 

1 

6 

15 

2 

12 

60 

14 

2 

4 

5 

3 

4 

1 

2 

1 

9 

4 

2 

15 

4 

1 

30 

2 

8 

13 

2 

4 

2 

5 

20 

10 

9 

12 

26 

4 

2 

2 

2 

16 

13, 14, 16, 17-19, 2n= 

15, 17, 18, 19, 20, 20- 

27 

24, 27, ca. 28, 36, 38, 

51+ 

16 

17. 18 

14, ca. 17, 34k2 

19, 19+1,20, 100f2 

15 

15 

8, 17, 39 

13, 14 

12, 35-+1, 36 

16 

12 

18 

11,22 

9, 10, 11, 22 

15-16 

17, 18 

17. 18 

9 

18 

18 

15 

14, ca. 24 

ca. 30. 32-34. 16? 

ca. 8, 10, 11, 11-12 

8 

26 

15 

30+B, 32+c.6 

ca. 15. 16 

14, 28 

14-15, 15, 15-17. 16, 

ca. 30, ca. 45, ca. 86 

14; 28, 42 

17. 34 

18, 19 

11 
NUMBER 52 

TABLE 3.-Continued 

Subtribe and genus 

Salmea 

Sanvitalia 

Stlphium 

Spilanthes 

Steiractinia 

Synedrella 

Verbesina 

Wedelia 

Wollastonia 

wuyja 

Wyethia 

Zexmenia 

Zinnia 

Keurolaeninae (Rydb.) 

Steussy, Turner & Powell 

Brasilia 

Ca lea 

Neurolaena 

Tetrachyron 

C'nxia 

Aldama 

Helianthinae Dumort 

Aluordia 

Garcilassa 

HtlianthopJis 

Helianthus 

Heliomeris 

Iostephane 

Lagascen 

PapPobolus 

Rhpsolepis 

Sca lesia 

Sclerocarpus 

Simsia 

Stuessya 

Tithonin 

General 

distribution 

Approx. number Chromosome numbers 

Habit of species (mostll, haploid) 

Mexico, Central America, West 

sw US, Mexico, Central America 

e US 

pantropical 

Indies 

n Andes 

pantropical 

US, tropical America, Africa 

s US, tropical America, Africa 

central and w Pacific 

Panama, West Indies, South 

w US 

sw US, tropical America 

sw US, Mexico, Central America, 

America 

Andes, cultivated 

Brazil 

tropical America 

Mexico, Central America, Andes 

Mexico, Guatemala 

Panama, n South America 

Mexico, Central America, VeneBaja California 

Central America, w South AmerAndes 

North America, cultivated, adventive 

w US. Mexico 

Mexico 

tropical America, adventive 

Peru 

Mexico 

Galapagos 

Mexico, Central America, n South 

s US, tropical America 

Mexico 

Mexico. Central America. cultivated 

zuela 

ica 

America, Africa 

7 

7 

23 

60 

6 

2 

150 

60 

ca. 20 

4 

14 

40 

22 

1 

95 

9 

1 

3 

1 

3 

1 

20 

ca. 50 

1 

2 

9 

4 

3 

14 

8 

35 

3 

10 

18, 32-33 

8, 9, 11. 16 

7 

7?. 12, 13, 16, 26, 39, 

14 

16, 18, 19, 20. 36 

16, 17, 18, ca. 30, 34, 

2n = ca. 88 

11, 12, 14, 20-22, ca. 

21. 22, 23. 25, ca. 26, 

28. 28+ 1, 29k 1, 33, 

ca. 36 

41 

15 

25, 30, 30k1 

19 

10. 11, 14, ca. 17 

ca. 24, 42+ 

10. 1 I, 12. 20, 21, 22. 

19 

16, ca. 17, 18. 19. ca 

11 

16 

16 

32 

17 

15, 30. 60 

17+1 or 18 

9. 17 

17. 34. 5 1 

8. 9. 16 

17, ca 34 

17, 18 

17 

17 

34 

11. 12. 14. 18 

17 

17 

16. 17 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 3.-Continued 

Subtribe and genus 

Viguiera 

Galinsoginae Benth. & Hook 

A lloispermum 

Aphanactis 

Bebbia 

Cuchumatanea 

Cymophora 

Galinsoga 

Jaegeria 

Sabaria 

Schistocarpha 

Tetragonotheca 

Tridax 

Dimeresiinae H. Robinson 

Coreopsidinae Lessing 

Dimeresia 

Bidens 

Chrysanthellum 

Coreocarpus 

Coreops is 

Cosmos 

Dahlia 

Dicranocarpus 

Glossocardia 

Glossogy ne 

Hen rickson ia 

Heterosperma 

Hida lgoa 

Narvalina 

Thelesperma 

Fitchiinae Carlquist 

Coulterellinae H. Robinson 

Pectidinae Lessing 

Fitchia 

Coulterella 

Adenophy llum 

Chrysactinia 

Dy ssodia 

Hydropectis 

General Approx. number Chromosome numbers 

distribution Habit of species (mostb haploid) 

US, tropical America 

Mexico, Central and South AmerCentral and n South America 

sw US, Mexico 

Guatemala 

Mexico 

tropical America, adventive 

tropical America 

Mexico, Central America, n Andes 

Mexico, Central America, Andes 

s US, Mexico 

Mexico, Central and South America 

ica, adventive 

w us 

cosmopolitan 

tropical America, Africa 

sw US, n Mexico 

North and tropical America, Africa, cultivated 

tropical America, cultivated 

Mexico, Central America, cultisw US, Mexico 

India 

se Asia, East Indies, Australia, w 

Mexico 

sw US, tropical America 

Mexico, Central and n South 

America 

West Indies 

w US, Mexico, e South America 

vated 

Pacific 

s Pacific 

Baja California 

sw US, Mexico, Central America 

sw US, Mexico 

sw US: Mexico, Central America 

Mexico 

160 

9 

7 

2 

1 

3-4 

14 

8 

15 

16 

4 

25 

1 

230 

10 

10 

114 

26 

27 

1 

2 

8 

1 

5 

5 

1 

12 

6 

1 

8 

4 

8 

2 

12, 16, 17, 18, 21, ca. 

33, ca. 34, 40k2, 

50k2 

9, 15, 16, 17, or 18, ca. 

19, 24, ca. 27, 32 

8 

9 

8 

8, 9 

8, 9, 16, la?, 24. 32 

9, 18, 36 

4, 8, 9, 16, 24, 36 

8 

.- I/ 

9, 10, 18, 27 

7 

10, 11, 12, 17, 18, 23, 

8 

24, 34, 36, 38, 73+2 

IL 

6, 7, 9, 10, 12, 13, 14, 

24, ca. 26, 28, 32, 39, 

ca. 56 

24, 33, 36 

11, 12, 13, 17, 22, 23, 

16, 17, 18, 32, 36 

10 

12 

12 

18 

9, 11, 13, 24, 25 

15, 16 

60 

8, 9, 10, 11, 12, 20, 22, 

24, 2n=44 

45 

18 

13 

15 

7, 13 

9 
NUMBER 52 15 

TABLE 3.-Continued 

Subtribe and genus 

Hymenatherum 

hicolietia 

PPCllS 

Porophf llum 

Strotheria 

Tagetes 

L7rbinella 

Flaueria 

Flaveriinae Lessing 

Haploesthes 

Surtceellia 

\.arillinae Turner and Powell ex H. Robinson 

Varilla 

Clappiinae H. Robinson 

Clappill 

Pseudoclappia 

Jaumea 

Achyrachaena 

Argyoxiphium 

Blepharipappus 

Blepharironia 

Cabcadenia 

Dubautia 

Hemitonia 

Ho ioca rp ha 

Holoron ia 

Lagophy ila 

Layia 

Madia 

Raillardeila 

Wilkesia 

Ga lea nu 

Hymenopappus 

Loxothysanus 

Vilianoua 

Peritylinae Rydb. 

Amauria 

Eutetras 

Pericoma 

Perityle 

Jaumeinae Benth. Br Hook. 

Madiinae Benth. Br Hook 

Hymenopappinae Rydb 

General 

distribution 

Approx. number Chromosome numbers 

Habit of sbecies ('mostly hubloid) 

sw US. Mexico. Central America. 

West Indies 

sw US, Mexico 

sw US, Mexico, West Indies, 

sw US, tropical America 

South America 

n Mexico 

sw US. Mexico. Central and 

central Mexico 

South America 

A. P, S 17 7. 8, 13. 16. 24. 26, 32, 

2n=52 

A, p 3 10 

A. P ca. 100 12. 2n=36, 48, 72 

A, P. S 30 11. 12. 15. 22. 24. 

2n=36. 72 

ss 1 8 

A, p, s ca 50 11, 12. 24, 2n=36 

A 1 8 

su US, Mexico, Central and A. P 18 18, 9 

SLV US, n Mexico P, s 3 18 

SM US. n iMexico A. P 4 18 

South America, Australia 

s US, Mexico S 2 18 

sw US. n Mexico 

sw US, n Mexico 

S 

S 

1 16 

2 18+1, 180r 19, 19 

w US, s South America P 2 19 

w us 

Hawaii 

w us 

California 

w US. w Mexico 

Hawaii 

w US, w Mexico 

California 

w US 

w us 

California, w Mexico 

w US, w Mexico, Chile 

w US 

Hawaii 

1 

4 

1 

1 

11 

26 

31 

1 

1 

5 

15 

18 

5 

1 

8 

13, 14 

8 

14 

+, 5. 6, 7, 9 

13. 14 

9, 10, 11, 12, 13, 14 

'1, 5? 6 

14 

7 

7, 8, 16 

6, 7, 8, 9, 14, 16, 24 

8, 9, 18 

ca. 12 

Mexico, Central America A 3 9 

US, Mexico B, p 10 17, 34 

Mexico p, s 2 15, 17? 

Mexico, Andes A, P 10 19 

w Mexico P 3 ca. 17, 18 

central Mexico P 1 18 

sw L'S, Mexico P 1 18 

w US, Mexico, Peru, Chile A, p 54 11, 12, 13, 16, 17, 18. 

19. ca. 54 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 3.-Continued 

Subtribe and genus 

Baeriinae Benth. & Hook. 

Amblyopappus 

Antheropeas 

Baeriopsis 

Eatonella 

Eriophy llum 

Lasthmia 

Lembertia 

,Won o lop i a 

Pseudobahia 

Achl ropappus 

.4rnica 

Ox2PaPPus 

Chaenactidinae Rydb 

Bahia 

Barllettia 

Chaenactis 

Chamaechaenactis 

Florestina 

Huhea 

Hymenothrix 

Hjpericophj llum 

Ma 110 topus 

Orochaenactis 

Pahfixia 

I’eucephjllum 

Pla?schkuhria 

Psathyrotopsis 

Schkuhria 

S>ntrichopappus 

I ’en qpes ia 

Ct’hitnqa 

Artinorpermum 

ilmblplepsis 

Bailqya 

Baku ina 

Duga ldia 

Ga 1 /la rdia 

Gaillardiinae Lessing 

Htlenium 

HI menox) s 

Psathi rotes 

General Approx. number Chromosome numbers 

distribution Habit of species (mostb haploid) 

California. nw Mexico. Peru. 

Chile 

California 

Baja California 

w us 

w US. British Columbia 

California 

California 

California 

w Mexico 

California 

Mexico 

north temperate zone 

sw US. Mexico, Guatemala, Chile 

sw US. n Mexico 

sw US: nw Mexico 

w us 

Texas, Mexico: Central America 

\v US. Baja California 

sw US. Mexico 

tropical .Africa 

e Asia 

California 

s and sw US. n Mexico 

sw US. nw Mexico 

w us 

Texas, n Mexico 

sw US. Mexico, Central and w 

South America, .Argentina 

sw cs 

California. Baja California 

California 

s US 

Texas, n Mexico 

s\+ US, Mexico 

s us 

sw US, Mexico, Guatemala 

s and M US. Mexico. s South 

America, culri\ ated 

US. tropical America 

c and M US, n Mexico, s South 

u US, nu Mexico 

.4merica 

A 1 

A 5 

A 1 

A, B 1 

A. P 11 

A 16 

A, P 1 

A 4 

A 2 

A 3 

A 1 

P 30 

,A, P 13 

A 1 

.4. P 40 

P 1 

’4 7 

A. P 8 

A. B 5 

P 7 

P 1 

‘A 1 

A. P 12 

S 1 

P 3 

P 2 

.A. P 10 

A 2 

P 1 

P 1 

A, B 3 

,4 1 

P 4 

B. P 3 

P 3 

A. P 29 

A. P 40 

A. P 25 

8 

4, 5 

8 

19 

4, 5, 6, 7, 8, 15, 16, 24, 

4,5,6,7,8,12,16,24 

10 

10 

10 

3, 4, 6, 8 

32-34 

10 

19, ca. 28, ca. 36, ca. 

38, 2n=54, 56, 57, 

58, 60, ca. 67, 70, ca. 

74, 76, 95, ca. 97 

8, 10, 11, 12, 24, 36 .. I1 

5, 6, 7, 8, 12, 18 

16 

10, 12 

19 

12, 24 

9 

9 

9 

10, 11, 12 

lo> 20 

12, 24. 36 

19 

10, 11> 12, 20, 21k1 

6. 7 

19 

19 

18 

18, 19 

16, 17 

18, 36 

15 

17. 18, 19, 34 

13, 14, 15, 16, 17, 18 

11, 15 

17 
NUMBER 52 17 

TABLE 3.-Continued 

Subtribe and genus 

Psilostrophe 

Tetraneuris 

Trichoptilium 

Marshallia 

Marshalliinae H. Robinson 

General Approx number Chromosome numbers 

distribution Habit of species (most4 haploid) 

sw US, Mexico P 6 16, 17. 18, 32 

c and w US, n Mexico A, p 26 14, 15, 28, 35, ca 45, 

sw US, w Mexico A 1 13 

56 

e and s US P 10 9 

these are not particularly closely related to the 

Espeletiinae or to each other. 

MELAMPODI1NAE.-The subtribe traditionally 

has contained most of the Heliantheae having 

functionally male disk flowers. Stuessy (1973, 

1977) has successively removed various genera 

from the group, but the present view reduces the 

subtribe further to include only five genera. Differences between the remaining genera leave 

some question about the naturalness of the group, 

but the genera themselves seem entirely natural. 

In spite of some variation in chromosome number, a case can be made for a base number of 

X=l 1 in the three herbaceous genera of the subtribe, with the more woody Smallanthus being a 

remotely related element having X= 16-1 7. 

The chromosome number in Melampodium 

shows considerable instability. The variation has 

been discussed by Turner and King (1962) and is 

reviewed by Stuessy (1971, 1972, 1979). The annual and perennial section Melampodium has X= 10 

on the basis of 14 species, the annual section 

Zarabellia has X=9 on the basis of four species, the 

annual section Serratura has X=12 on the basis of 

three species, the rhizomatous perennials of section Rhizomeria have X=ll on the basis of two 

species, and the annuals of section Afoina have 

X= 11 on the basis of three species with a few 

reports of n=12. Stuessy (1971, 1979) suggests 

that section Melampodium with X= 10 is an ancestral type in the genus. 

Because Melampodium represents the most citable example of presumable aneuploid increase in 

the tribe, it is worthy of special reconsideration 

(see page 6). Stuessy’s conclusions in his 1971 

paper were restated in cladistic form in 1979. On 

the basis of his earlier treatments of the genus, 

the characters used include chromosome number, 

habit, distribution, number of outer involucral 

bracts, form of bract margin, and presence or lack 

of a unique type of ovary in the disk flowers. 

Primitive states are determined in part by comparison with the apparently related genera Acanthospermum and Lecocarpus. The chromosome numbers are evaluated on a simple distance basis, 

apparently without considering that aneuploid 

decrease might be much more likely than increase. Careful study of Stuessy’s (197 1, 1979) 

concept shows that it requires three parallel aneuploid increases from 10 to 11 or 12, two in Melampodium and one in the Acanthospermum-Lecocarpus 

line, and it requires two parallel origins of the 

unique undifferentiated ovary of the disk flowers. 

In our own view, a monophyletic origin of the 

unique type of ovary is more likely, even at the 

cost of considering the type of ovary in section 

Melampodzum as a limited revcrsion. Such reversion or de-differentiation is not difficult when it 

represents a return to a form present in the ray 

florets of all the species. Our concept would eliminate the need for parallel aneuploid increase at 

the n=ll level. The n=12 of section Serratura 

possibly did occur by aneuploid increase. 

An anomaly in Melampodzum not yet explained 

is the annual habit of most of the species with 

higher base numbers. The prrennial subshrubs 

seem restricted mostly to two groups apparently 

specialized into less tropical habitats: section Rhztomarza of higher elevations in central and southern Mexico is among those with X= 11, and section Melampodzum series Leucantha of the southwestern United States and northern Mexico is 

among those with X=lO. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Ichthyothere is usually placed adjacent to the 

genus Clibadium, and the two are considered 

closely related by Stuessy (1977). Robinson (198 1) 

separates the two and places Ichthyothere in the 

Melampodiinae nearer Smallanthus on the basis of 

achene wall structure. The reports of n=16 

(Turner et al., 1979) and n = ca. 33 in Ichthyothere 

(Coleman, 1970) could correlate with either suggested relative. 

POLYMNIINAE.-The one genus of the subtribe 

has been placed in the Melampodiinae in previous systems, and Polymnia has included Smallanthus in its synonymy. Achene structure combined 

with nonpapillose limbs of the rays distinguish 

Polymnia from all members of the Melampodiinae. 

The X=15 of Polymnia furnishes a slight but apparently consistent additional difference from 

Smallanthus with X=16-17. The similarity between Polymnia and the Melampodiinae does not 

necessarily indicate close relationship. Unxia, 

which has similar characters, and which has species until recently placed in Melampodium and 

Polymnia, proves to be a relative of Calea in the 

Neurolaeninae (Robinson, 1980). 

MILLERIINAE.-The present concept retains 

only Milleria of the genera traditionally placed in 

the subtribe. The other genera included here have 

previously been placed in the Melampodiinae or 

the Ecliptinae (Verbesininae), depending on the 

extent to which the disk flowers are bisexual. 

One genus, Guizotia, has been placed traditionally in the Coreopsidinae, a position first questioned by Robinson and Brettell (1973a) and 

later by Baagoee (1974). More recently, Stuessy 

(1977) has returned the genus to the Coreopsidinae, and Baagoee (1977) has concurred. Guizotia is 

one of the two members of the Compositae in 

which the cytochrome c sequence has been studied. The eight amino acid differences between 

Guizotia and Helzanthus are at least a partial basis 

for the recent elevation of the Coreopsidinae to 

tribal rank (Turner and Powell, 1977b). In both 

anatomy and cytology, however, Guizotia is out of 

place in the Coreopsidinae. There is much better 

correlation with members of the Milleriinae, but 

the genus falls outside of the closely related group 

formed by the other members of the subtribe. 

The variation of chromosome number in Sigesbeckia is discussed briefly by Keil and Stuessy 

(1975), and the predominance of n=15 and 30 is 

emphasized. There are reports of n=12 (Subramanyan and Kamble, 1967) and 2n=20 (Hsu, 

1967) for S. orientalis L. A base of X=12 has also 

been suggested for the related genus Rumfordia 

(Sanders, 1977). 

is available for one species, Desmanthodium fruticosum Greenm. The n= 18 conforms with the general pattern of high base numbers in the tribe. 

CLIBADIINAE.-ChrOmOSOme counts are known 

from about 10 species of Clibadium, including 

three new reports for C. armanii Sch. Bip. ex Baker 

of Brasil, and X= 16 seems to be the base number 

of the genus. A previous count of n=24 from C. 

armanii (Coleman, 1968) suggests a more basic 8 

chromosome complement, but such a count could 

also result from hybridization between plants 

with n=16 and n=32. The latter interpretation is 

favored by the lack of lower numbers in Clibadium 

and by the derived nature of lower numbers in 

most subtribes of the Heliantheae. 

The closely related Riencourtia is reported for 

the first time in this paper, with n = ca. 16. 

GuARDIoLINAE.-The single genus Guardiola has 

usually been placed in the Melampodiinae because of the male disk flowers. Recently, Stuessy 

(1973, 1977) transferred the genus to the Coreopsidinae, with which it shares a number of characteristics. The chromosome number of X= 12, 

based on counts of eight species (Van Faasen, 

1973), seems to support this latter placement. 

Unfortunately, the genus differs from the Coreopsidinae in three of the most definitive features of 

that subtribe. The fact that X=12 is the base 

number of only specialized elements of the Coreopsidinae, but not the base of that whole subtribe, lessens the value of the number as evidence 

of close relationship. 

ENHYDRINAE.-The subtribe consists of a single 

genus of specialized aquatics and is obviously the 

result of morphological reduction. There is a 

DESMANTHODIINAE.-One chromosome report 
NUMBER 52 19 

superficial resemblance to some members of the 

Ecliptinae where the same chromosome numbers, 

11 and 15, are also common, but details of the 

style and achene indicate there is no close relationship. 

MONTAN0INAE.-The single genus Montanoa has 

been placed in the broad and poorly defined 

subtribe Verbesininae that was considered the 

typical element of the tribe Heliantheae in the 

traditional systems of Bentham and Hooker 

(1873) and Hoffmann (1890-1894). The subtribe 

was interpreted much more narrowly by Stuessy 

(1977), but continued to include Montanoa. Anatomically, the achenes of Montanoa are totally 

distinct from those of the Verbesininae (included 

here in the Ecliptinae), and they are more like 

those of the Melampodiinae and Milleriinae. The 

chromosome number of X=19 is based on data 

from five species and seems stable. Polyploidy in 

the genus reaches n= 1 10- 120 (probably 12x, 

n= 114), the highest number reported for the Asteraceae (Funk and Raven, 1980). 

RUDBECKI1NAE.-The three genera of the subtribe form a closely related group with a probable 

base number of X=18-20. Both Rudbeckia and 

Ratibida have a series of numbers that can be 

interpreted as separate reductions from higher 

base numbers, but the single species of Dracopis 

seems to represent a derived element with n=16. 

Robinson’s (198 1) revised classification removes 

Echinacea from this group to the strictly defined 

Ecliptinae on the basis of anatomy. The transfer 

is supported by the chromosome numbers n= 11 

and 22 in the latter genus. 

ZALUZANI1AE.-There are reports of n= 17 and 

18 for Zaluzania and n=16 for the recently resegregated Hybridella (Olsen, 1977). Emphasis on 

chromosome number has led Olsen (1979) to 

equate the n= 17 of Z. grayana B. L. Robinson and 

Greenm. with the helianthine genus Viguiera, but 

partially fertile rays, a basal flange on the corolla, 

and details in resin ducts clearly mark the species 

as a Zalutania. There is no particularly close relationship to the Helianthinae. 

ECLIPTINAE.-The subtribe is interpreted here 

to include the Verbesininae, the Engelmanniinae, 

and the Zinniinae. In view of patterns in other 

subtribes, a higher original base number is suggested here. The element containing Verbesina, 

Podachaenium, and the closely related series EnceliaEnceliopsis-Flourensia-Geraea reflects this original 

number, lower numbers not being seen in these 

genera. Balsamorrhiza and Wyethia, which also 

have n=19 or 20, represent a temperate element 

in this basically tropical subtribe. The higher 

number in the latter genus seems of passing interest in view of the ancestral aspect credited to 

the genus by Cronquist (1955). 

The typical Ecliptinae, including Calyptocarpus, 

Delilia, Eclipta, Wedelia, and Zexmenia, seem to 

have a base number at or near 11 with some 

redoubling or tripling. The closely related Synedrella with n=16, 18, 19, and 20 may be the result 

of secondary polyploidy and subsequent secondary aneuploid loss. Echinacea has X= 1 1, and both 

Zinnia and Sanvitalia seem to have lower base 

number from 8- 12. 

Numerous genera show an intermediate base 

number, such as Melanthera, Steiractinia, Otopappus, 

Philactis, Helianthella, Heliopsis (Fisher, 1957), Borrichia (Semple, 1977), Perymenium (Fay, 1978), and 

Oyedaea with X= 14- 16. The counts in Philactis 

and Heliopsis tend to confirm the artificial nature 

of the traditional Zinniinae. The X= 15 of Wollastonia (King, 1964) seems to support removal of 

the genus from Wedelia (Fosberg and Sachet, 

1980). Lipochaeta of Hawaii has been interpreted 

to include species related to Wollastonia (Gardner, 

1977), but the former genus is limited here to the 

five species in the typical group having X=26. 

Gardner (1977) suggests two possible origins of 

this number: from a polyploid of X=15 with 

subsequent aneuploid loss from n=30 and from 

polyploidy of a hybrid between plants with n= 15 

and n=l1. 

Spilanthes seems to be on a separate descending 

series. Keil and Stuessy (1975) claim four base 

numbers for the genus, but allowing for miscounts 

and erratics, the counts resolve easily into X= 16 

(Spilanthes) and X= 13 (Acmella) . Eleutheranthera, 

with n=15 and 16, seems closely related to Blainvillea, with 72-8, 17, and 39?, suggesting that the 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

higher number of the former genus might be a 

doubling of 8. On the same basis, the specialized 

temperate group containing Engelmannia might 

represent secondary polyploidy. Stuessy’s (1973) 

suggested chromosomal history of the group 

might be correct, with Engelmannia representing a 

retained basic X=9, Lindheimera representing a 

derived X=8, Chrysogonum with X=16 being a 

polyploidy of the X=8, and Berlandiera with X= 15 

being a polyploid with aneuploid loss. Related to 

this group (Stuessy, 1977) is Silphium, with X=7. 

Of the present authors, Robinson and King consider the X=9 of Engelmannia as derived from 

much higher numbers, traces of the higher numbers still being evident in the closely related Wedelia series. 

The Andean genera Monactis and Kingianthus 

share high numbers of ca. 30-34. The new counts 

agree generally with the previous report of n = 

ca. 30 for Monactis wurdackii H. Robins. (Turner 

et al., 1967, as Monopholis jelskii (Heiron.) Blake) 

and n=32 + 2 frag.. in M. kingii H. Robins. (as 

M. flaverioides H.B.K., Jansen and Stuessy, 1980). 

One species of Monactis has aborted anthers, and 

apomixis seems probable in both genera. The two 

genera form a closely related pair of obscure 

relationship to other members of the subtribe. 

The report by Olsen (1979) of n=18 for Kingianthus paniculata (Turcz.) H. Robins. (as Zalutanza sodiroi Hieron.) conflicts with the present reports for the same species and conflicts with the 

otherwise consistent counts of ca. 30-34 for the 

generic pair. The Olsen count does conform with 

those of the Mexican genus Zalutania of the subtribe Zaluzaniinae, in which Olsen placed the 

species. 

NEcROLAENIIZAE.-The subtribe contains at 

least two distinctive elements represented by 

Neurolaena and Calea. The two totally different 

base numbers, based on numerous counts of both 

genera, reenforce the distinction. Because of 

trends seen elsewhere in the tribe, the lower numbers are regarded as the result of aneuploid loss 

from primitive polyploid ancestors. This seems to 

be confirmed in Calea by the lack of lower numbers and in the closely related Brasilia with X= 19 

(Turner et al., 1979). The X=ll of Neurolaena 

might represent aneuploid gain from X= 10, but 

the genus is not so isolated in the subtribe as to 

suggest such a different history. 

Previous treatments have associated both Neurolaena and Calea more closely with genera now 

placed in the Galinsoginae. Neurolaena has been 

closely linked with Schistocarpha, but basic differences were first suggested by Robinson and Brettell (1973b), and the lack of close relationship has 

been confirmed since by anatomical and chemical 

studies (Robinson et al., 1978; Robinson, 1979). 

The base numbers of the two genera have long 

been recognized as different, but with the present 

overview, the X=8 of Schistocarpha is seen to conform with other Galinsoginae, whereas the X= 1 1 

of Neurolaena is completely foreign to that subtribe. 

Confusion in the case of Calea has resulted from 

the previous inclusion of the superficially similar 

genus Alloispermum. Anatomy and chemistry show 

that the two genera are not closely related (Robinson et al., 1978; Robinson, 1979). In spite of 

some shared numbers, the chromosome complements of the two genera also are fundamentally 

different. In Calea, X= 18 and 19 are basic for the 

genus. In Alloispermum, the base is X=8 or 9, with 

n= 16- 18 derived by polyploidy. 

Traditionally, Unxia has been placed in the 

subtribe Melampodiinae because of the male disk 

flowers and the lack of pappus. The genus proves 

to belong to the Neurolaeninae (Robinson, 1980) 

and shows the generally high chromosome number characteristic of the subtribe. The count of 

n=16 is based on the more widely distributed 

annual species U. camphorata L. f. The two suffruticose or fruticose Amazonian species, which are 

structurally more like Calea, have not been studied 

cytologically. 

HELIANTHIxAE.-The subtribe is considered by 

Robinson (198 1) to be completely natural in the 

present interpretation, and the base number is 

obviously X=17. A few examples of descending 

aneuploid series are seen in Viguiera and Sclerocarpus. Feddema (197 1) used the chromosome differences as a partial basis for the resurrection of 
NUMBER 52 21 

Aldama from the synonymy of Sclerocarpus. Eliasson 

(1974) has correctly interpreted the Galapagan 

genus Scalesia, with X=34, as being polyploid 

from X=17. There is a single report of n=18 in 

Lagascea by Chopde (1965), but all other counts 

for 10 of the 11 taxa are n=17 (Stuessy, 1978). 

There seems to be a polyploid series based on 

X=15 in Alvordia. The n=1711+11 or n=18 in 

Garcilassa represents the first report for the genus 

and tends to confirm the position in the Helianthinae given by Robinson (1981). 

Two occurrences of lower numbers in the subtribe present a problem of interpretation. They 

do not seem to represent an original condition in 

the subtribe, but neither are they connected with 

an evident aneuploid series. Heliomeris, recently 

resegregated from Viguiera by Yates (in herb.), is 

scarcely distinguishable except by details of the 

style and the nearly consistent X=8 based on 

counts of five species. One count of n=9 is cited 

by Keil and Pinkava (1976). In Helianthopsis hypargyrea (Blake) H. Robins, reports of n=9 and 

n= 17 in this paper are accompanied by another 

report of n= 17 for the closely related H. pseudoverbesinoides (Hieron.) H. Robins. The n=9 in Helianthopsis and the X=8 of Heliomeris must be 

interpreted as derived numbers within the subtribe, unless one accepts very unlikely relationships or unlikely parallelisms in chromosome 

number stabilization. Since the reduction in numbers seems so abrupt, we suggest that polyhaploidy or a closely related process is involved (Raven 

and Thompson, 1964; Ornduff, 1970; DeWet, 

1971; Anderson, 1973). It would seem that the 

ancestrally polyploid chromosome complement 

in the Helianthianae is unusually susceptible to 

abrupt reversions to approximations of the diploid number. 

It is notable that Robinson (1981) places the 

Helianthinae close to the Galinsoginae and Dimeresiinae, two subtribes that show no present 

evidence of ancestral polyploid numbers. Two of 

the present authors, Robinson and King, believe 

the three subtribes all share a polyploid ancestry 

and probably also share the susceptibility to 

polyhaploidy or a related process. Powell reserves 

judgment, especially on the Galinsoginae. An 

alternate proposal to that of Robinson and King 

is possible involving nonpolyploid ancestry for 

the Galinsoginae and Dimeresiinae and an independent polyploid development in the Helianthinae. Robinson and King believe such an interpretation places considerable weight on a few, 

mostly isolated subtribes specialized toward annual and short-lived perennial life-forms, and it 

does not adequately explain the preponderance 

of polyploidy in all other elements of the tribe 

Heliantheae. 

GALINSOCINAE.-The subtribe as delimited by 

Robinson (1981) is regarded as natural, and it is 

one of the few in the tribe showing no relicts of 

higher base numbers. All higher numbers in the 

tribe are readily interpreted as polyploids from 

X=8 or X=9. As indicated in the previous paper 

of the series (Solbrig et al., 1972), the count of 

n=4 in Sabazia is from a specialized annual species. 

In Alloispermum, the only base number thus far 

reported is X=9 with the derived n=17-18 and 

ca. 27, but X=8 is well represented in counts of 

n=16, 24, and 32. There is general agreement on 

X=8 for Galinsoga. The n= 18 reported by Nawaschin (1935) seems out of place, and Canne 

(1977a) makes the point that she has not seen the 

voucher. Tridax is credited with bases of X=9 and 

10 by Powell (1965) with polyploids of 9 only. 

The most recent of a series of studies of the 

Mexican genus Cymophora, with n=8, indicates 

that it should be kept separate from Tridax 

(Turner and Powell, 1977a), but the difference is 

based only on one count of one species (Turner et 

al., 1973). In aspect, Tridax dubia Rose, with n=9, 

is closer to Cymophora. An attempt by Powell to 

count the chromosomes of Tridax veneiuelensis Arist. & Cuatr. (Venezuela: Distrito Federal, Steyermark LY Aristeguieta 122, VEN, US) gave only 

approximate results, only three cells with rather 

sticky chromosomes, bivalents appeared heteromorphic, 2n = ca. 18. Canne (1977b) transferred 

the latter species to Cymophora, but the geographic 

disjunction plus other differences mentioned by 

Canne suggest a distinct phyletic line is involved. 

DIMERESI1NAE.-The subtribe consists of a sin- 
22 

gle, minute, annual herb from western North 

America having a chromosome number of n=7. 

Dimeresia seems to follow the pattern of annual 

habit correlating with reduced numbers. 

COREOPSIDINAE.-The subtribe has been expanded recently by Stuessy (1977) and, with a 

few important omissions, the limits are regarded 

here as natural. The most common chromosome 

number in the subtribe is n=12, and Stuessy 

seems to regard this as the base number. In a 

study of the chromosome numbers of Coreopsis, 

which has implications for the whole tribe, Smith 

(1975) suggests a generic ancestor with X= 14 and 

separate aneuploid reductions to various groups 

with n=12 and 13. The reduction series in the 

typical line includes X= 13, 10, 9, 7, and 6. Smith 

suggests ultimate derivation of Coreopsis from 

plants with still higher chromosome numbers one 

to several aneuploid steps higher than Coreopsis. 

A high base number of X= 17- 19 is suggested for 

the tribe on the basis of the observations on the 

Coreopsidinae and the high numbers of other 

genera such as Heiianthus, Verbesina, and Rudbeckia. 

In support of Smith, it is notable that within the 

Coreopsidinae higher numbers are characteristic 

of the distinctive genera Hidalgoa with X=15 and 

16, Dahlia with 16, 17, and 18, and Henricksonia 

with n=18 (Turner, 1978), forming some of the 

most divergent elements in the subtribe. 

The apparent restriction of X=12 to more derived elements of the Coreopsidinae lessens the 

significance of that number in the distinctive 

genus Guardioia as a basis for including it in the 

subtribe. Remarks on another omitted genus, Guizotia, may be found under the Milleriinae. 

FITCHIINAE.-The subtribe contains only the 

Polynesian genus Fitchia. One chromosome count 

of n=45 has been reported for F. speciosa Cheesm. 

(Solbrig et al., 1972). Studies by Carlquist (1957) 

have shown basic similarities between the genus 

and the Coreopsidinae. The high number of chromosomes is correlated with an arborescent habit 

and large pollen. A base number of X= 15 seems 

possible. 

COULTERELL1NAE.-The subtribe contains one 

species, Coulterelia capitata Vasey & Rose, of southSMITHSONIAN CONTRIBUTIONS TO BOTANY 

ern Baja California. A count of n=18 (Powell, 

pers. comm.) conforms to the general pattern of 

high numbers in the tribe. 

PECTID1NAE.-Strother (1 97 7), treating the 

group as a tribe, Tageteae, has provided a schematic diagram of relationships between the genera. The two basic groups are recognized as separate subtribes, Pectidinae with only Pectis, and 

Tagetinae with the remaining genera. In the 

Tagetinae three groups are recognized. The first, 

containing Urbanella, Strotheria, Hymenatherum, and 

Hydropectis, is credited with the greatest concentration of advanced characteristics combined 

with most of the chromosome numbers between 

X=7 and X=9. The genera of the second group 

include Tagetes, Adenopappus, Adenophyllum, Dyssodia, Gymnolaena, and Schizotrichia. These have 

X=11-13, except for Dyssodia with X=7 and 13. 

In the remaining group, the more advanced genera include Porophyllum with X=ll, 12, and 15 

and Nicolletia with X= 10, while Chrysactinia, with 

X= 15, is placed near the base. The scheme shows 

the general pattern seen in many other subtribes 

where lower numbers seem derived. The highest 

number, X=15, might be derived from lower 

numbers, but it is in a group of genera in which 

numbers below X=10 are notably absent. Of the 

present authors, Robinson and King believe the 

X=15 is representative of a more ancestral condition in the subtribe. 

Strother (1969), in his treatment of “Dyssodia,” 

discusses the apparent dibasic condition of the 

genus, where there is an enigmatic lack of intermediate numbers. Adjacent n=8 and n=13 populations of Hymenatherum pentachaeta DC. (as Dyssodia) are cited. Strother indicates the n=13 species in Adenophyllum (as Dyssodia subg. Clomenocoma) 

are more primitive, whereas the n=7 species are 

advanced. Strother comments further on the taxonomic remoteness of “Dyssodia” species with n=7 

and n=8, a fact that makes the X=4 hypothesis 

of Johnston and Turner (1962) difficult to accept. 

Strother seems to favor low base numbers of X=4 

and 5, but all extant populations are regarded as 

complex products of polyploidy and (subsequent?) aneuploidy. Higher numbers such as 
KUMBER 52 23 

n=16 and n=26 are accepted as the result of 

secondary polyploidy. Regarding the lack of intermediate numbers, Strother cites the study of 

Gilia (Grant, 1966), where hybrid lines showing 

a full range of numbers from 2n=36-56, when 

selfed for nine generations, showed predisposition 

to revert to 2n=36, 38, and 50. 

FLAvERIINAE.-The review of structure by 

Robinson (198 1) supports the association of Flaueria and Sartwellia (Gray, 1852; Rydberg, 1915) 

and the addition of Haploesthes from the Senecioneae (Turner and Johnston, 1961; Powell, 1978). 

On the basis of many counts, the three genera 

have a consistent X=18 (Turner, 1971, 1975; 

Powell and Powell, 1978; Powell, 1978). An interesting case of probable polyhaploidy with n=9 

has been cited for Flaueria campestris Johnston 

(Anderson, 1973). 

VARILLINAE.-The single genus Varilla, with 

X=18, conforms to the general concept of high 

basic chromosome numbers in the tribe. Details 

of structure tend to confirm the close relationship 

between Varilla and the Clappiinae that were 

parts of the somewhat amorphous subtribe Varilliinae proposed by Turner and Powell (1977b). 

Other paleaceous genera of the Heliantheae that 

were mentioned by Turner and Powell are of 

remote relationship. The paleae of Varilla are 

considered a secondary development from epaleaceous ancestry by Robinson (1981). 

CLAPPI1NAE.-The two epaleaceous genera 

Clappia and Pseudoclappia seem to belong to a 

larger natural group including the paleaceous 

genus Varilla and the epaleaceous subtribes Flaveriinae and Jaumeinae. The chromosome numbers of all four subtribes are in the general high 

range found throughout most of the tribe. 

Keil and Stuessy (1977) indicate that the 

proper count for Pseudoclappia arenaria Rydb. is 

probably n=19. Powell and Powell (1977) report 

n= 19 for P. watsonii Powell &Turner, but indicate 

the need for verification in both species. 

JAC.MEIKAE.-AS delimited here, the subtribe 

contains only one genus with two species. Both of 

the disjunct species in the genus share X=19 

(Powell et al., 1975). The genera Espejoa and 

Hypericophyllum, once included in the synonymy of 

Jaumea, are placed in the Chaenactidinae by Robinson (1 98 1). 

MADIINAE.-The subtribe has been well characterized in the past, and chromosome data are 

comparatively complete (Carlquist, 1959; Clausen et al., 1945). Solbrig et al. (1972) indicate 

that “the basic number in the Hawaiian Madiinae appears to be n=14, while in the Pacific 

Coast species, the most pervasive number is X=7, 

although n=8 and n=9 are also frequent.” The 

common occurrence of n=7 along with n= 14 suggests the latter is derived by polyploidy from the 

former in the subtribe. This impression is 

heightened by the fact that most occurrences of 

n=14 are in the geologically younger portion of 

the subtribal range. It is notable that the Hawaiian genera are more woody. The tendency for 

more shrubby habits in insular members of herbaceous groups is well known. A lesser example 

of perennial development is seen in a few North 

American Madiinae on the close offshore islands 

of California. Increased chromosome numbers are 

to be expected in such shrubby insular groups. 

In spite of evidence from within the subtribe, 

the general trend in the Heliantheae would indicate derivation of the Madiinae from ancestors 

with higher numbers. Such continental represent at ives as Calycadenia, Madia, and Raillardella 

are interpreted here as aneuploid series from an 

original number of not less than X=9. The continuous series from X= 14 downward in Hemizonia 

also might include relicts of higher ancestral numbers. 

HYMENOPAPP1NAE.-The present concept may 

be artificial, but any artificiality is within narrow 

limits. The revised alignment introduces no new 

problems in interpretation of the chromosome 

data, since the anomalous element, Galeana with 

n=9, has traditionally been placed adjacent to 

Villanova with n= 19. The presence of higher numbers in three of the genera plus the example of 

other subtribes leads Robinson and King to consider the number in Galeana as derived. 

PERITYL1NAE.-The redelimitation of the subtribe by Robinson (1981) eliminates all elements 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

lacking distinct petioles and possessing striations 

on the achene. As such, all genera included by 

Powell and Turner (1974) with chromosome 

numbers lower than X=18 also are eliminated, 

except for the obvious declining aneuploid series 

in Perityle (Powell, 1968, 1969). 

BAERIINAE.-The subtribe as presented seems 

to be a natural group. The distribution of chromosome numbers indicates a common ancestor of 

X=10. Eatonella, with X=19, seems close to Lembertia and apparently is derived from a polyploid 

of X=10. The higher numbers in Lasthenia, an 

obviously specialized genus, are also likely to be 

of polyploid origin (Ornduff, 1966). It is the 

evidence of related subtribes that causes Robinson and King to believe the X=10 is derived, in 

turn, from higher numbers, rather than being a 

direct reflection of the original X=10 of the 

subfamily Asteroideae. 

CHAENACTIDINAE.-The subtribe is interpreted 

broadly to include the Bahiinae, although the 

latter group forms a rather distinctive element 

within the subtribe. The obviously related subseries Bahia, Florestina, Palafoxia, Platyschkuhria, 

and Schkuhria has a rather consistent X=10-12. 

Since aneuploid decrease is more likely, the X= 12 

is presumed to represent the ancestral number of 

the series. The typical Chaenactidinae have a 

mixture of higher and lower numbers with the 

X= 19 of Arnica, Hulsea, Psathyrotopsis, Venegesia, 

and Whitneya representing the primitive condition. The n=16 of Chamaechaenactis also seems a 

relict of originally higher numbers. The series of 

numbers in Chaenactis, however, seems to indicate 

that the n=18 in some specimens of C. douglasii 

(Hook.) Hook. & Arn. is of polyploid origin from 

X=6. 

The n=9 for one species of Hypericophyllum 

(Powell et al., 1975) tends to support its separation from the genus Jaumea, with which it has 

often been placed. 

The genus Arnica, one of a series of Heliantheae 

that has been traditionally placed in the Seneci- 

. 

oneae, is clearly in this subtribe. The structural, 

chemical, and cytological reasons for excluding 

the “Arniceae” from the Senecioneae are summarized by Nordenstam (1977), with a subsequent note of interest by Robins (1977). 

GA1LLARDIINAE.-The subtribe is closely related to the typical element of the Chaenactidinae, especially to such genera as Arnica and Psathyrotopsis. In the present group, however, the 

higher base numbers are more consistently retained. The most obvious reduction series is in 

the closely knit generic group with Helenium, Hymenoxys, and Tetraneuris. Within Helenium, section 

Hecubaea is considered most primitive by Bierner 

(1972). Both species of the section are now known 

to have chromosome numbers of n= 17 (Keil and 

Stuessy, 1977). 

Trichoptilium with X=13 is regarded here as a 

product of a separate reduction series in the 

relationship of Psathyrotes with X= 17. 

MARSHALLIINAE.-There is a single chromosome report for Marshallia graminiflia (Walt.) 

Small with n=9 (Jones, 1970). 

The genus Marshallia traditionally has been 

placed in the Heliantheae, but recently the position has been questioned, and a position in or 

near the Eupatorieae has been suggested (Stuessy, 

1977; Turner and Powell, 1977b). Robinson 

(198 1) rejects a relationship to the Eupatorieae 

and cites characters that place the genus definitely in the Heliantheae-Inuleae relationship. 

The latter position is supported by chemistry 

(Bohlmann et al., 1979). Robinson retains the 

genus as a subtribe in the Heliantheae isolated 

from the rest of the tribe. Improved understanding of the Inuleae may eventually show that 

Marshailia is better accommodated in that tribe. 

Tails of short to medium length are present on 

the anthers of some species. 

At best, Marshallia is isolated from other members of the Heliantheae and probably does not 

share the ancestry of higher base numbers that is 

evident in most other subtribes. 
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