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2Department of Botany, University of Hawaii at Manoa, Honolulu, Hawaii 96822; 3Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; and 4Department of Biology, Sul Ross State University, Alpine, Texas 79832
Received for publication June 12, 1998. Accepted for publication December 1, 1998.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONLITERATURE CITED |
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14II), Feldstonia (2n = 11II), Gochnatia (2n 23II), and Pseudoconyza (n = 10). In addition, new chromosome numbers are established at the generic level in Acourtia, Calea, Craspedia, Gnaphalium, Helipterum, Liabum, Leucheria, Smallanthus, Trixis, and Viguiera and at the specific level in 13 additional species.
Key Words: Asteraceae • chromosome numbers • Compositae • cytotaxonomy • cytogenetics
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONLITERATURE CITED |
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and more fully referenced by Robinson et al. (1997)
. In conjunction with data from other approaches, chromosome information continues to be useful in assessing generic and tribal relationships in this large and complicated family. Publication of broad surveys of chromosome numbers also makes it possible for subsequent workers to identify situations where further study may be rewarded with increased understanding of relationships on a finer scale. Thus, we present 316 new chromosome number determinations for species in tribes Anthemideae (8), Arctotideae (1), Astereae (70), Calenduleae (2), Cardueae (2), Heliantheae (126), Inuleae (47), Lactuceae (11), Liabeae (12), Mutisieae (12), Senecioneae (1), Tageteae (5), and Vernonieae (19). First reports are provided for 66 species and for the genera Cassinia, Feldstonia, Gochnatia, and Pseudoconyza. New chromosome numbers differing from those on record for ten genera and 13 additional species are also reported. Most of our samples came from Mexico to South America and Australia; additional collections were from Africa and elsewhere. |
MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONLITERATURE CITED |
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for further clarification) from material collected by R. M. King. The counts have been made from aceto-carmine or aceto-orcein squashes of microsporocytes in meiosis. Voucher specimens are in US and MO. Indexes consulted for previous chromosome number determinations include: (Ornduff, 1968
; Moore, 1973,
1974,
1977
; Fedorov, 1974
; Goldblatt, 1981,
1984,
1985,
1988
; Goldblatt and Johnson, 1990,
1991,
1994,
1996
). We have usually cited only these in order to save space and still provide a means to access the primary literature. In several instances, however, the complexity of the situation or omissions or errors in the indexes have mandated the citation of primary sources.
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RESULTS AND DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONLITERATURE CITED |
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) but agrees with a report of 2n = 10II for Athanasia bremeri by Strother, Watson, and Panero (1996)
. Other genera of Anthemideae represented in Africa and known to have species with 2n = 10II include Cotula, Inezia, Lasiospermum, and Matricaria (Heywood and Humphries, 1977
; Källersjö, 1991
).
Arctotidae
The report here of 2n = 9II for Arctotis cuprae is the first for this species. Norlindh (1977)
and others have reported the same number for other species of Arctotis and also other genera of the tribe.
Astereae
First reports are provided for species of Baccharis (5), Conyza (1), Diplostephium (3), Grindelia (1), Hysterionica (1), Olearia (2), and Plagiocheilus (1). From chromosome indexes we have concluded that no new numbers have been established in any of these genera as a result of these new species determinations. With the exception of Grindelia, the newly reported species determinations in these genera reflect x = 9, which is also pervasive at the tribal level.
Although most of our determinations for Brachycome mirror earlier reports for the same species, the report here of 2n = 18II for B. cardiocarpa and 2n = 12II for B. aculeata disagree with earlier counts of n = 9 and n = 9, 18, 27, 36, and 45 for these species, respectively (Smith-White, Carter, and Stace, 1970
). For B. cardiocarpa, our count likely reflects previously unreported infraspecific polyploidy; this phenomenon is well documented in other species of the genus. Our report of 2n = 12II for B. aculeata, which is otherwise known to have only numbers based on x = 9, is problematic. In a genus that has proven to be as cytogenetically diverse and taxonomically difficult as Brachycome, the explanation could relate to either cytogenetics or taxonomic interpretation. There are six other species of Brachycome for which n = 12 has been reported (Smith-White, Carter, and Stace, 1970
; Watanabe and Short, 1996
). Watanabe et al. (1996)
suggested that species of Brachycome with chromosome numbers of n = 11 to n = 15 have arisen by amphidiploidy.
Our count of n = 4 for Nevada material of Machaeranthera rigida var. aristata differs from the previous report of n = 5 for Arizona material of M. rigida by Solbrig et al. (1964)
. Additional collections and determinations will be required to assess the possible significance of the different numbers reported for this species.
Calenduleae
Our determination of 2n = 7II for Calendula officinalis differs from previous reports of n = 14 and 16 for this species; 2n = 7II has been found in at least five other species in the genus (Moore, 1973
; Fedorov, 1974
; Goldblatt, 1981
).
Heliantheae
First reports are provided for species of Calea (2), Espeletia (1), Melanthera (1), Monactis (1), Montanoa (1), Oyedaea (1), Pappobolus (1), Psilostrophe (1), Smallanthus (3), Tilesia (1), Tridax (1), Verbesina (2), Viguiera (3), and Wedelia (3). Of these, only the reports for species of Calea and Smallanthus establish new chromosome numbers at the generic level. The new count here for Calea ternifolia (2n 21II + B's) also establishes a new number in Calea.
The chromosome number reported for populations of three varieties of Bidens odorata in Mexico and Central America is n = 12 (Ballard, 1986
). Our reports of 2n = 36II for Ecuadorean populations referred to this species raise some doubt as to the proper identity of our material. Ballard (1986)
discussed the occurrence of a form of B. pilosa in South America that closely resembles B. odorata. Although Ballard did not have access to material from South America, he reported that all plants of B. pilosa sampled in Mexico and Central America were hexaploid (n = 36). More collections and cytological determinations of Andean material will probably help clarify the taxonomy of B. pilosa and its close relatives in South America.
In Calea, chromosome numbers of n = 9, 10, 16, 17, 18, 19, 19 + 4–7 B, 24, 27, and 32 have been reported (Moore, 1973
; Fedorov, 1974
; Goldblatt, 1981
, 1984
, 1988
; Pruski and Urbatsch, 1984
; Goldblatt and Johnson, 1990
). To these we add 2n = 13II for C. angusta and C. pilosa and 2n 21II + B's for C. ternifolia. According to Urbatsch (personal communication, Lousiana State University), C. angusta and C. pilosa appear to form a subgroup within C. sect. Meyeria close to a group including C. abbreviata (see Pruski and Urbatsch, 1988
). Given the diversity of chromosome numbers already known in Calea, it appears that additional cytological determinations will be a valuable adjunct to any study aimed at clarifying infrageneric relationships in the genus.
Our report of 2n 39II for Coreopsis boliviana appears to be hexaploid based on x = 13, as apparently is the case in C. major (Moore, 1973
). The only other report for C. boliviana is 2n 23 (Goldblatt, 1985
). Additional collections and cytological determinations will be necessary to clarify the relationship between the numbers reported for C. boliviana.
Chromosome numbers of Melampodium have been rather extensively reviewed; accepted reports include n = 9, 10, 11, 12, 18, 20, 23, 25 ± 1, 27, 30, and 33 (Turner and King, 1962
; Stuessy, 1970
, 1971
). Intensive sampling of M. divaricatum has revealed only counts of n = 12 (Turner and King, 1962
; Stuessy, 1971
). Thus, our count of 2n 30II credited to this species based on a specimen from Brazil represents additional cytological complexity or perhaps taxonomic confusion. Likewise, our count of 2n = 12II + 1 B for M. cupulatum is at odds with previous reports of n = 10 for the species (Turner and King, 1962
; Stuessy, 1971
).
Our determinations for Smallanthus latisquamata (2n = 16II), S. pyramidalis (2n = 29II), and S. glabratus (2n 30II) match or closely approximate previously reported numbers for the genus (n = 16, 27–30, 29 + 1, and 34; Robinson et al., 1981
; Strother, 1983
). Our report of 2n = 32II in S. maculatus, together with earlier counts of n = 16, 16, and 34, seems to substantiate the occurrence of autopolyploidy in this species.
Our report of 2n = 9II + 
IV for Thelesperma megapotamicum matches an earlier report by Keil and Pinkava (1976)
and is also reconcilable with reports of n = 11, 22, and 18II + 2 IV for the same species (Strother, 1976
; Powell and Powell, 1977
). The extensive cytological evaluation of Thelesperma by Greer (1997)
confirmed the earlier counts and also documented additional cytogenetic peculiarities in T. megapotamicum and related species.
Previous reports for Viguiera include n = 8, 11, 12, 17, 18, 21, 32, 34, and 40 (Fedorov, 1974
; Robinson et al., 1981
; Goldblatt, 1984
, 1988
). Our first reports of 2n 42II for V. mucronata and 2n 51II for V. linearis are new numbers in the genus, perhaps representing a tetraploid based on x = 21 and a hexaploid based on x = 17, respectively.
Our report of 2n 28II for Wedelia angustifolia agrees with a report of n = 28 for W. pilosa Baker (Moore, 1973
). Other species of Wedelia share the presumed base number of x = 14 as well (Moore, 1974
; Goldblatt and Johnson, 1991
). Wedelia is the third generic home for this species; it was treated previously as Aspilia cupulata S. F. Blake, and originally as Oyedaea angustifolia Gardn. Cytologically, the species would fit well in Oyedaea, in which x = 14 appears to dominate. The new report of 2n 35II for Wedelia silphioides is not easily reconciled with the previous report of n = 23 for the species (as Aspilia silphioides; Goldblatt, 1988
). Our approximate count may reflect x = 12, a base number which is pervasive in Wedelia (Fedorov, 1974
; Moore, 1973
; Goldblatt, 1988
; Goldblatt and Johnson, 1991
, 1996
). The earlier count of n = 23 is matched in at least three other species of Wedelia (Fedorov, 1974
; Moore, 1974
; Goldblatt and Johnson, 1990
). The disparity in chromosome numbers attributed to Wedelia silphioides is most likely due to taxonomic confusion.
Inuleae
Our reports of 2n 14II for Cassinia longifolia, 2n = 11II for Feldstonia nitens, and 2n = 10II for Pseudoconyza viscosa represent the first for these genera. First reports for species of Achyrocline (1), Craspedia (1), Gnaphaliothamnus (1), Gnaphalium (2), Helichrysum (3), Helipterum (2), and Pluchea (3) are also provided. These counts establish new chromosome numbers at the generic level in Craspedia, Gnaphalium, and Helipterum. Our count of 2n = 12II in Waitzia aurea differs from the previous determination of n = 10 for the species but agrees with reports for at least three other species in the genus (Moore, 1973
).
Our count of 2n = 44 (mostly univalents) for Craspedia glauca is equivalent to the count of n = 22 for C. uniflora (Goldblatt, 1988
). An individual of C. glauca from another location yielded a count of 2n = 44II (with occasional univalents). This is a new number for Craspedia, apparently representing octoploidy based on x = 11, a number that occurs elsewhere in the genus (Moore, 1973
).
Chromosome numbers of n = 7, 8, 9, 10, 14, 21, 26, and 28 have been reported for Gnaphalium (Moore, 1973
; Fedorov, 1974
; Goldblatt and Johnson, 1990
, 1991
). Our report of 2n = 20II for G. purpurascens represents a new number for the genus, perhaps a dysploid derivative of the hexaploid based on x = 7 or a tetraploid based on x = 10.
In Helipterum, chromosome numbers of n = 5, 7, 8, 10, 11, 12, 14, and 24 are known (Moore, 1973
; Fedorov, 1974
). If the 2n = 13II alternative of our dubious report of 2n = 13II (or possibly 12II + IV) for H. hyalospermum proves correct, this would reflect a new number for the genus.
Pseudoconyza viscosa has had a checkered past, from Conyza (Astereae) to Pseudoconyza (Astereae) to Blumea (Inuleae) and back to Pseudoconyza (Inuleae). Cytologically, at 2n = 10II, it appears to be at home in Inuleae-Plucheeae, sharing the base number x = 10 with the genera Epaltes, Laggera, Pluchea, and Sphaeranthus (cf. Federov, 1974
; Goldblatt, 1981
).
Lactuceae
Our determinations for this tribe include a first report for Hieracium traillii (2n 18II). No new numbers are established at the generic level.
Liabeae
First reports are provided for species of Chrysactinium (1), Erato (1), and Liabum (3). Because of the difficulty in getting good cytological preparations in this tribe, as discussed by Robinson et al. (1985)
, reports have been rather confusing and difficult to interpret. Each of our new determinations is discussed in the context of previous reports and in line with the generic circumscriptions of Robinson et al. (1985)
.
Our first report of 2n = 12II for Chrysactinium acaule agrees with earlier reports for C. hieracioides and C. rosulatum, but for the latter two species other counts of n = 13–14, or even 15–16, were also acknowledged as possible, based on some of the cells sampled (Robinson et al., 1985
). Despite some confusion, n = 12 seems to be the recurrent number in all of the determinations of Chrysactinium to date (Robinson et al., 1985
; Goldblatt and Johnson, 1990
).
Three separate determinations for Erato vulcanica and one for E. polymnioides are n = 9 (Robinson et al., 1985
) but for the latter species n 11 has also been reported (Goldblatt and Johnson, 1990
). Our first report for E. soderoi is 2n = 9–12II or possibly 9II + B's or univalents. Additional determinations would obviously be helpful to evaluate the variation in chromosome numbers reported for Erato.
Chromosome numbers of n = 17–20, 18, 19, 19 + 2, 19 + 6 or 7 fragments, 20, and 20 + 1 or 2 fragments have been reported for species of Liabum (Robinson et al., 1985
; Goldblatt and Johnson, 1990
). To these we add first reports of 2n 38II for L. asclepiadeum and 2n = 18II + 5 micro B's for L. wurdackii. We also provide what may be the first count for L. bourgeaui of 2n = 36II. The possibly erroneous previous report of n = 9 for L. bourgeaui was discussed by Robinson et al. (1985)
. At this point it appears that x = 18 and 19 are firmly established and represented at both the diploid and tetraploid levels in the genus.
For species of Munnozia, available counts include n = 10, 11, 12, 12 +1, 13, and 24 (Robinson et al., 1985
). Our report of 2n = 13II for Munnozia jussieui confirms the earlier report of n 13 for this species. Collections from two different locations for M. hastifolia in Venezuela yielded 2n = 10II, in contrast with n = 11 and 12 reported earlier for specimens from Ecuador (Robinson et al., 1985
).
Mutisieae
Our count of 2n 23II for Gochnatia vernonioides is the first for the genus. First reports are also provided for species of Acourtia (1), Jungia (2), and Leucheria (1). Our determination of 2n = 26II for Acourtia rigida establishes a new number in the genus; previously only n = 27 and 54 have been reported for three other species (Goldblatt, 1981
, 1985
). Likewise, our first report for Leucheria achillaefolia of 2n = 19II is unique in the genus, differing from reports of n = 20 for a dozen other species (Goldblatt, 1981
, 1988
; Goldblatt and Johnson, 1991
). We also report a new number in Trixis; our count of 2n = 28II differs from the previous report of n = 27 for Trixis inula (Sundberg, Cowan, and Turner, 1986
) and eight other species in the genus (Moore, 1973
, 1977
; Fedorov, 1974
; Goldblatt, 1981
, 1985
, 1988
; Goldblatt and Johnson, 1990
).
Tageteae
Our determination of 2n = 8III for Thymophylla acerosa from New Mexico extends the occurrence of triploids in this species. The common occurrence of triploidy, apparently associated with apomixis, in T. acerosa was documented by Strother (1989)
, based on populations in Texas and Arizona.
Vernonieae
First reports are provided for one species each of Gutenbergia, Lychnophora, and Vernonanthura. The count of 2n = 9II + B's for Lychnophora phylicifolia establishes a new number in the genus in which previously only n = 17 had been reported (Goldblatt, 1984
). Our count of 2n = 17II for Eirmocephala brachiata disagrees with n = 16 previously reported for the species (as Vernonia; Goldblatt, 1988
).
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FOOTNOTES |
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONLITERATURE CITED |
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Fedorov, A. [ed.]. 1974 Chromosome numbers of flowering plants. Otto Koeltz, Koenigstein.
Goldblatt, P. [ed.]. 1981 Index to plant chromosome numbers 1975–1978. Monographs in Systematic Botany from the Missouri Botanical Garden 5: 1–553.
———. 1984 Index to plant chromosome numbers 1979–1981. Monographs in Systematic Botany from the Missouri Botanical Garden 8: 1–427.
———. 1985 Index to plant chromosome numbers 1982–1983. Monographs in Systematic Botany from the Missouri Botanical Garden 13: 1–224.
———. 1988 Index to plant chromosome numbers 1984–1985. Monographs in Systematic Botany from the Missouri Botanical Garden 23: 1–264.
Goldblatt, P., and D. E. Johnson [eds.]. 1990 Index to plant chromosome numbers 1986–1987. Monographs in Systematic Botany from the Missouri Botanical Garden 30: 1–243.
———, and ———. 1991 Index to plant chromosome numbers 1988–1989. Monographs in Systematic Botany from the Missouri Botanical Garden 40: 1–238.
———, and ———. 1994 Index to plant chromosome numbers 1990–1991. Monographs in Systematic Botany from the Missouri Botanical Garden 51: 1–267.
———, and ———. 1996 Index to plant chromosome numbers 1992–1993. Monographs in Systematic Botany from the Missouri Botanical Garden 58: 1–276.
Greer, L. F. 1997 Origins of polyploidy in Thelesperma (Asteraceae: Heliantheae, Coreopsidinae). Master's thesis, Sul Ross State University, Alpine, TX.
Heywood, V. H., and C. J. Humphries. 1977 Anthemideae—systematic review. In V. H. Heywood, J. B. Harborne, and B. L. Turner [eds.], The biology and chemistry of the Compositae, vol. 2, 851–898. Academic Press, London.
Källersjö, M. 1991 The genus Athanasia (Compositae-Anthemideae). Opera Botanica 106: 1–75.
Keil, D. J., and D. J. Pinkava. 1976 Chromosome counts and taxonomic notes for Compositae from the United States and Mexico. American Journal of Botany 63: 1393–1403.[CrossRef][ISI]
Moore, R. J. [ed.]. 1973 Index to plant chromosome numbers 1967–1971. Regnum Vegetabile 90: 1–539.
———. 1974 Index to plant chromosome numbers for 1972. Regnum Vegetabile 91: 1–108.
———. 1977 Index to plant chromosome numbers for 1973/74. Regnum Vegetabile 96: 1–257.
Norlindh, T. 1977 Chapter 33. Arctoteae—systematic review. In V. H. Heywood, J. B. Harborne, and B. L. Turner [eds.], The biology and chemistry of the Compositae, vol. 2, 943–959. Academic Press, London.
Ornduff, R. [ed.]. 1968 Index to plant chromosome numbers for 1966. Regnum Vegetabile 55: 1–126.
Powell, A. M., and S. A. Powell. 1977 Chromosome numbers of gypsophilic plant species of the Chihuahuan Desert. Sida 7: 80–90.
Pruski, J. F., and L. E. Urbatsch. 1984 Chromosome counts in Calea (Asteraceae-Heliantheae). Plant Systematics and Evolution 144: 151–153.[CrossRef][ISI]
———, and ———. 1988 Five new species of Calea (Compositae: Heliantheae) from planaltine Brazil. Brittonia 40: 341–356.[CrossRef][ISI]
Raven, P. H., O. T. Solbrig, D. W. Kyhos, and R. Snow. 1960 Chromosome numbers in Compositae. I. Astereae. American Journal of Botany 47: 124–132.[CrossRef][ISI]
Robinson, H., G. D. Carr, R. M. King, and A. M. Powell. 1997 Chromosome numbers in Compositae, XVII: Senecioneae III. Annals of the Missouri Botanical Garden 84: 893–906.[CrossRef][ISI]
———, A. M. Powell, R. M. King, and J. F. Weedin. 1981 Chromosome numbers in Compositae, XII: Heliantheae. Smithsonian Contributions to Botany 52: 1–28.
———. 1985 Chromosome numbers in Compositae, XV: Liabeae. Annals of the Missouri Botanical Garden 72: 469–479.[CrossRef][ISI]
Smith-White, S., C. R. Carter, and H. M. Stace. 1970 The cytology of Brachycome. I. The subgenus Eubrachycome: a general survey. Australian Journal of Botany 18: 99–125.[CrossRef][ISI]
Solbrig, O. T., L. C. Anderson, D. W. Kyhos, P. H. Raven, and L. Rüdenberg. 1964 Chromosome numbers in Compositae V. Astereae II. American Journal of Botany 51: 513–519.[CrossRef][ISI]
Strother, J. L. 1976 Chromosome studies in Compositae. American Journal of Botany 63: 247–250.[CrossRef][ISI]
———. 1983 More chromosome studies in Compositae. American Journal of Botany 70: 1217–1224.[CrossRef][ISI]
———. 1989 Chromosome numbers in Thymophylla (Compositae: Tageteae). Sida 13: 351–358.
———, L. E. Watson, and J. L. Panero. 1996 Documented chromosome numbers 1996.3. Chromosome numbers in some South African Compositae. Sida 17: 265–268.
Stuessy, T. F. 1970 Chromosome studies in Melampodium (Compositae, Heliantheae). Madroño 20: 365–372.
———. 1971 Chromosome numbers and phylogeny in Melampodium (Compositae). American Journal of Botany 58: 732–736.[CrossRef][ISI]
Sundberg, S., C. P. Cowan, and B. L. Turner. 1986 Chromosome counts of Latin American Compositae. American Journal of Botany 73: 33–38.[CrossRef][ISI]
Turner, B. L., and R. M. King. 1962 A cytotaxonomic survey of Melampodium (Compositae-Heliantheae). American Journal of Botany 49: 263–269.[CrossRef][ISI]
Watanabe, K., T. Denda, Y. Suzuki, K. Kosuge, M. Ito, P. S. Short, and T. Yahara. 1996 47. Chromosomal and molecular evolution in the genus Brachyscome (Asteraceae). In D. J. N. Hind and H. J. Beentje [eds.], Compositae: systematics, vol. 1, 705–722. Royal Botanic Gardens, Kew.
Watanabe, K., and P. S. Short. 1996 Chromosome number determination in Brachyscome Cass. (Asteraceae: Astereae) with comments on species delimitation, relationships and cytogeography. Muelleria 7: 457–471.
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| HOME | HELP | FEEDBACK | SUBSCRIPTIONS | ARCHIVE | SEARCH | TABLE OF CONTENTS |
). K numbers represent R. M. King collection numbers. Counts by Powell are identified as (Powell) after the collection number; all other counts are by Carr. Vouchers are in US and MO.