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2University Herbarium, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, California 94720-2465 USA; and 3School of Biological Sciences, Section of Integrative Biology, University of Texas, Austin, Texas 78712 USA
Received for publication January 18, 2000. Accepted for publication May 19, 2000.
| ABSTRACT |
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Key Words: Asteraceae chromosomes Compositae Heliantheae phylogeny systematics taxonomy
| INTRODUCTION |
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| MATERIALS AND METHODS |
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Our format here is the same as before (Strother and Panero, 1994
): each entry in Results and Discussion consists of name of taxon (first counts for species are marked *; counts that differ from other published counts are marked
); chromosome number [and meiotic configuration(s)]see Strother and Nesom (1997)
; collection information (state only); reference(s) to other reports of chromosome number, if any (see following); and discussion (if appropriate).
To provide access to original literature without having an unduly long Literature Cited here, for counts that confirm reports of chromosome numbers listed in standard indices, reference to earlier counts is made only by index citation. The index edited by Fedorov (1969)
is coded F. The various indices titled Index to plant chromosome numbers ... are coded as follows (according to last year covered): I-65 (Ornduff, 1967
), I-66 (Ornduff, 1968
), I-71 (Moore, 1973
), I-72 (Moore, 1974
), I-74 (Moore, 1977
), I-78 (Goldblatt, 1981
), I-81 (Goldblatt, 1984
), I-83 (Goldblatt, 1985
), I-85 (Goldblatt, 1988
), I-87 (Goldblatt and Johnson, 1990
), I-89 (Goldblatt and Johnson, 1991
), I-91 (Goldblatt and Johnson, 1994
), I-93 (Goldblatt and Johnson, 1996
), and I-95 (Goldblatt and Johnson, 1998
). Other references are cited directly, as appropriate.
| RESULTS AND DISCUSSION |
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Perezia carpholepis A. Gray). 2n = c. 54 (c. 27 II): Oaxaca, P 6759.
*Acourtia scapiformis (Bacigalupi) B. L. Turner (
Perezia scapiformis Bacigalupi). 2n = c. 54 (c. 27 II): Oaxaca, P 5549.
Our reports for acourtias are consistent with reports of 2n = 54 in other species of Acourtia (e.g., I-78, I-83; some reported as Perezia spp.).
*Ageratina josepaneroi B. L. Turner. 2n = 3436 (1718 II): Guerrero, P 6183.
*Ageratina pendula Panero & Villaseñor. 2n = 34 (17 II): Oaxaca, P 5686.
Counts of 2n = 34 have been recorded for ten or more other species of Ageratina (e.g., I-78, I-83).
*Alepidocline trifida (J. J. Fay) B. L. Turner (
Sabazia trifida J. J. Fay). 2n = 16 (8 II): Oaxaca, P 5006; 2n = 16 (8 II): Oaxaca, P 6783.
Strother and Panero (1994)
reported 2n = 8 II + 02 f for A. annua S. F. Blake.
Alloispermum scabrifolium (Hook. & Arn.) H. Rob. 2n = 32 (16 II): Jalisco, P 5592.
Our count differs from the count of "n = 17 ± 1" for this species reported by Solbrig et al. [1972; as Calea scabrifolia (Hook. & Arn.) Hemsl.] for a plant from Nayarit. Most counts for other species of Alloispermum have been 2n = 32.
*Brickellia problematica B. L. Turner (
Barroëtea glutinosa Brandegee). 2n = 18 (9 II). Oaxaca, P 6784.
Counts of 2n = 18 have been recorded for a host of other barroëteas and brickellias (e.g., I-66, I-71, I-72, I-74, I-78, I-81, I-83, I-87, I-89).
Calea megacephala B. L. Rob. & Greenm. 2n = 38 (19 II): Chiapas, P 4483. I-83, I-85, I-87.
Carminatia recondita McVaugh. 2n = 20 (10/10 in dyads): Guerrero, P 6193. I-87.
Chaetymenia peduncularis Hook. & Arn. 2n = 18 (9 II): Jalisco, B 978. I-93.
Chromolaena sagittata (A. Gray) R. M. King & H. Rob. 2n = 20 + 2 f (10 II + 2 f): Sinaloa, P 6178. I-66, I-78, I-87 (some as Eupatorium sagittatum A. Gray).
Coreopsis mutica DC. var. holotricha (S. F. Blake) S. F. Blake. 2n = c. 42 (c. 21 II): Oaxaca, P 6148.
Other counts for C. mutica (with or without variety indicated) have been 2n = 48 and 52 (as n = c. 24 and c. 26; Turner, Powell, and King, 1962
) and 2n = 56 (as "gam" = 28; I-72, I-81, I-85). Our report here of 2n = c. 42 is anomalous for the species.
*Cosmos juxtlahuacensis Panero & Villaseñor. 2n = 26 (13 II): Oaxaca, P 3506.
Counts of 2n = 22 and 24 (and polyploid numbers) have been recorded for other species of Cosmos. Our report here is evidently the first report of a species of Cosmos with 2n = 13 II. Counts of 2n = 20, 22, 24, 26, and 28 (and polyploid numbers) have been recorded for species in the sister genus Coreopsis (F, I-65, I-66, I-71, I-72, I-74, I-78, I-81, I-83, I-85, I-87, I-89, I-91, I-93, I-95).
*Dahlia pteropoda Sherff. 2n = 64 (32 II): Oaxaca, P 3598.
Most counts for other species of Dahlia have been 2n = 32 or 2n = 64 (e.g., I-71, I-72, I-78).
Desmanthodium fruticosum Greenm. 2n = c. 34 (c. 17 II): Michoacán, P 3625.
Other counts recorded for D. fruticosum have been 2n = 34 (I-89) and 2n = 36 (I-74, I-78, I-89).
*Digitacalia crypta B. L. Turner. 2n = 60 (30 II): Guerrero, P 6186.
Counts of 2n = 60 have been reported for D. chiapensis (Hemsl.) Pippen (I-83) and D. tridactylitis (B. L. Rob. & Greenm.) Pippen (I-71).
*Galinsoga crozieri Panero. 2n = 16 (8 II): Durango, G 6187.
Galinsoga longipes Canne. 2n = 16 (8 II): Michoacán, P 3624; 2n = 32 (16 II): Michoacán, P 3630.
Our report of 2n = 32 is evidently the first report of a tetraploid plant of G. longipes. Canne (1983)
reported n = 8 and 2n = 16 for each of two separate plants of G. longipes from the state of México.
Galinsoga parviflora Cav., vel aff. 2n = 48 (24 II): Oaxaca, P 6785.
Other reports for G. parviflora have been 2n = 16 or 2n = 32: I-65, I-66, I-71, I-72, I-74, I-78, I-81, I-83, I-85, I- 87, I-89, I-91, I-95.
Galinsoga quadriradiata Ruiz & Pavón. 2n = 32 (various IVs + IIs): Durango, H 7468. I-81, I-83, I-85, I-87, I-89.
Heliomeris obscura Cockerell. 2n = 26 (13 II): Oaxaca, P 3545.
The only other count reported for H. obscura was 2n = 28 for a plant from Puebla (Strother and Panero, 1994
). Counts for other species of Heliomeris (some reported as viguieras) have been 2n = 16 [Viguiera annua (M. E. Jones) S. F. Blake, I-78; V. ciliata (B. L. Rob. & Greenm.) S. F. Blake, I-78; H. hispida (A. Gray) S. F. Blake, I-89; H. longifolia (B. L. Rob. & Greenm.) Ckll., I-66, I-71, I-72, I- 81, I-85, I-89, Carr et al. (1999)
; H. multiflora (Nutt.) S. F. Blake, I-74, I-83; H. multiflora (Nutt.) S. F. Blake var. nevadensis (Nels.) Yates, I-72, I-81, I-89] but for one report of 2n = 8 II + 2 I and 2n = 9 II (for a plant of Viguiera multiflora Nutt., I-78) and one report of 2n = 16 II [for V. multiflora Nutt. var. nevadensis (Nels.) S. F. Blake, I-72].
*Hofmeisteria dissecta (Hook. & Arn.) King & H. Rob. 2n = 34 (17 II): Guerrero, P 6191.
*Hofmeisteria schaffneri (A. Gray) King & H. Rob. 2n = c. 36 (c. 18 II): Jalisco, P 5589.
*Hofmeisteria urenifolia (Hook. & Arn.) Walp. 2n = 36 (18 II): Jalisco, P 5593.
Counts of 2n = 34 or 36 (I-95), 36 (I-71, I-78), and c. 38 (I-74) have been reported for Hofmeisteria fasciculata (Benth.) Walp. We found no reports for other species of Hofmeisteria.
Jaegeria glabra B. L. Rob. 2n = 18 (9 II): México, P 3621. I-71, I-93, Carr et al. (1999)
.
Microspermum nummulariifolium Lag. 2n = 24 (12 II): Oaxaca, P 5168. I-71, I-87.
Montanoa revealii H. Rob. 2n = c. 114 (c. 57 II): Oaxaca, P 5067.
Prior counts for M. revealii have been "n = 56 + 1" and "n = 57 + 1" (I-81) and "n = 5657 + 1" (I-83); the species is evidently hexaploid (x = 19 for the genus).
*Oteiza acuminata La Llave. 2n = 16 (8 II): Michoacán, P 5299.
When we reported 2n = 34 (17 II + 03 f) for O. scandens (Strother and Panero, 1994
), we remarked that n = 17 is an unusual chromosome number for a member of Galinsoginae, for which chromosome numbers are usually based on x = 4, 8, or 9. Given that O. acuminata has 2n = 8 II, once again the thought that O. scandens may be an amphidiploid comes to mind.
Packera sanguisorbae (DC.) C. Jeffrey (
Senecio sanguisorbae DC.). 2n = c. 46 (c. 23 II): Morelos, P 7356.
Turner, Beaman, and Rock (1961)
reported "n = ca. 30" for P. sanguisorbae (as Senecio sanguisorbae); voucher: J. H. Beaman 2638 from Cerro Potosí, Nuevo León. Other packeras have been reported to have 2n = 46 or other numbers based on x = 23 (F, I-65, I-66, I-71, I-72, I-74, I-78, I-81, I-83, I-85, I-87; some as senecios).
Philactis zinnioides Schrad. 2n = 56 (28 II): Chiapas, P 5749.
Torres (1969)
suggested that the ultimate base chromosome number for Philactis is 14 and treated P. zinnioides s.s. (sensu stricto) (2n = 84, see I-71) as a presumed hexaploid distinct from the presumed tetraploid (2n = 56, see: F, I-71, I-78) species P. liebmannii (Klatt) S. F. Blake and P. nelsonii (Greenm.) S. F. Blake. We consider Philactis to include a single, chromosomally variable species, P. zinnioides.
Porophyllum punctatum (Mill.) S. F. Blake. 2n = 22 (11 II): Sinaloa, P 6179.
Other reports for P. punctatum have all indicated 2n = 24 (F, I-74, I-78, I-83). Counts of n = 22 have been reported for other species of Porophyllum.
*Porophyllum viridiflorum (Kunth) DC. 2n = 22 (11 II): Guerrero, P 6187.
Counts of 2n = 22 have been reported for some species of Porophyllum and 2n = 24 others.
*Psacaliopsis paneroi (B. L. Turner) C. Jeffrey var. paneroi. 2n = c. 60 (c. 30 II): Guerrero, P 3315.
Ours is evidently the first count for the genus. Counts of 2n = 60 have been recorded for species in genera closely related to Psacaliopsis (e.g., Digitacalia, Psacalium).
*Roldana mixtecana Panero & Villaseñor. 2n = c. 60 (c. 30 II): Oaxaca, P 5099.
Counts of 2n = 60 have been recorded for ten or more other species of Roldana (some recorded as species of Senecio).
Rumfordia guatemalensis (J. M. Coult.) S. F. Blake. 2n = 24 (12 II): Hidalgo, P 3734.
Strother (1983)
reported 2n = 13 II for a plant of R. guatemalensis from Chiapas. Sanders (1977) reported "n = 24" for a plant identified as R. floribunda DC. from Oaxaca.
Sabazia humilis (Kunth) Cass. 2n = 8 (4 II): Distrito Federal, P 3407. F, I-71, I-83, I-89.
Sabazia liebmannii Klatt var. hintonii Longpre, vel aff. 2n = c. 16 (c. 8 II): Guerrero, P 3321.
Prior counts for S. liebmannii have indicated 2n = 6x = 48 (F, I-71, I-78, I-83; all for var. liebmannii or variety not stated) with one exception. Sundberg, Cowan, and Turner (1986)
reported "2n = ca. 12 II" for var. hintonii; the voucher for that report is annotated "n = 12 II (circa)." Panero's 3321 differs in some traits from typical var. hintonii and may prove to belong to a different, perhaps unnamed, taxon.
Schistocarpha bicolor Less. 2n = 16 (8 II): Oaxaca, P 5524. F, I-87.
Sigesbeckia agrestis Poepp. 2n = 24 (12 II): Michoacán, P 3629; 2n = 22 (11 II): Oaxaca, P 3571.
Other counts for S. agrestis have all been 2n = 30 or 60 (I-78, I-81, I-83, I-85, I-87).
*Sigesbeckia blakei (McVaugh & Lask.) B. L. Turner. 2n = 26 (13 II): Oaxaca, P 4220.
Most species of Sigesbeckia have been reported to have 2n = 30. Our report of 2n = 26 for S. blakei is based on four or more, well-stained diakinetic cells.
Sigesbeckia jorullensis Kunth. 2n = 30 (15 II): Distrito Federal, P 3312. I-72, I-78 ("gam" = 15, 30), I-81, I-85, I-95 ("gam" = c. 16).
Sigesbeckia repens B. L. Rob. & Greenm. 2n = c. 30 (c. 15 II): Oaxaca, P 3577. F, I-87.
*Sinclairia ismaelis Panero & Villaseñor. 2n = 18 (9 II): Oaxaca, P 3572.
Reports for other species of Sinclairia have been: 2n = 34 or 36 [I-74, for S. discolor Hook. & Arn., as Liabum discolor (Hook. & Arn.) Hemsl.], 2n = 30 or 32 [I-85, for S. hypochlora (S. F. Blake) Rydb.], and 2n = 30, 32, or 34 [I-85, for S. sublobata (B. L. Rob.) Rydb.]. Robinson et al. (1985)
suggested x = 16 for Sinclairia.
Smallanthus maculatus (Cav.) H. Rob. 2n = 32 (16 II): Michoacán, P 3623.
Half a dozen or more counts for S. maculatus (some reported as Polymnia maculata Cav.) have indicated 2n = 32 for the species. Other numbers reported for S. maculatus have been "n = 17" (for a plant identified as P. cf. maculata; Turner, Powell, and King, 1962
), n = "ca. 16" and "ca. 34" (Robinson et al., 1981
), and "2n = 32 II" (Carr et al., 1999
).
Stenocarpha filiformis (Hemsl.) S. F. Blake. 2n = 18 (9 II): Sinaloa, P 3651.
Four or more reports for S. filiformis (
Galinsoga filiformis Hemsl.) have indicated 2n = 16 for the species (F, I-65, I-83, I-93). Solbrig et al. (1972)
reported 2n = 18 and Canne (1983)
reported n = 8 and 8 + 1 and 2n = 16 and 16 + 2 for plants from Sinaloa.
Stevia aschenborniana Schultz-Bip. ex Klotzsch var. aschenborniana. 2n = 24 (12 II): Guerrero, P 6168.
The only other report for S. aschenborniana (without indication of variety) is 2n = 22 by Keil and Stuessy (1977)
.
*Stevia lita Grashoff, 2n = 5 II. Oaxaca, P 3594.
One group of stevias is characterized by x = 11, another by x = 12, and a third by x = 17 (Grashoff, Bierner, and Northington, 1972
). Clearly 2n = 5 II for S. lita is a remarkable new number for the genus. Such intrageneric heterogeneity in chromosome numbers is very unusual but not unheard of. Hymenoxys Cass. provides a similar or parallel example: most hymenoxyses have 2n = 30, 28, 24, or 22; in H. texana (J. M. Coult. & Rose) Cockerell, plants with 2n = 16 (8 II) and others with 2n = 6 (3 II) are known (Strother and Brown, 1988
).
Stevia viscida Kunth. 2n = c. 34 (c. 17 II?, c. 17/17 each pole in dyads): Zacatecas, P 6173.
Stevia viscida is a chromosomally complex apomict and other reports (F, I-66, I-72) have indicated x = 11 for the species or complex. Grashoff, Bierner, and Northington (1972)
reported 2n = 34 I for a plant from Durango identified as "S. viscida > S. serrata var. serrata."
*Tetrachyron brandegeei (Greenm.) Wussow & Urbatsch. 2n = 30 (15 II): Oaxaca, P 3609.
Reports for other species of Tetrachyron (some as species of Calea) have been 2n = 32 (I-81, I-83, I-87) and 2n = 34 (I-95).
*Thymophylla aurantiaca (Brandegee) Rydb. [
Dyssodia aurantiaca Brandegee]. 2n = 16 (8 II): Oaxaca, 3597.
Strother (1989)
suggested that x = 8 is the base number for Thymophylla.
Tridax balbisioides (Kunth) A. Gray. 2n = 20 (10 II): Durango, H 7455. F, I-65, I-71, I-72.
Tridax bicolor A. Gray, 2n = 20 (10 II): Chihuahua, H 7451. I-65, I-71.
*Tridax brachylepis Hemsl. 2n = 20 (10 II): Oaxaca, P 5007.
*Tridax candidissima A. Gray. 2n = 18 (9 II): San Luis Potosí, P 6629.
*Tridax luisana Brandegee. 2n = 18 (9 II): Oaxaca, P 5034.
Tridax mexicana A. M. Powell. 2n = 18 (9 II): Oaxaca, P 5001. F, I-71, I-78, I-83.
*Tridax palmeri A. Gray. 2n = 20 (10 II): Oaxaca, P 6790.
*Tridax oaxacana B. L. Turner. 2n = 20 (10 II): Oaxaca, P 5049.
Tridax rosea Schultz-Bip. 2n = 20 (10 at each pole in dyads): Queretaro, H 7484. I-65.
Tridax trilobata (Cav.) Hemsl. 2n = 20 (10 II): México, P 7342. I-65.
According to Powell (1965)
, species referred to Tridax sect. Tridax have chromosome numbers based on x = 9 and species included in sect. Imbricata A. M. Powell have numbers based on x = 10. Our reports here are consistent with the pattern noted by Powell.
Trigonospermum melampodioides DC. 2n = c. 26 (c. 13 II): Oaxaca, P 3563.
Four other reports [F, I-74 (two), I-78] all indicated 2n = 30 for this species and other trigonospermums have numbers based on x = 15 as well. Of the four diakinetic cells we studied, two seemed clearly to have no more than 13 II and two were "sticky" and might have been interpreted as having 13 to 15 or 16 II.
Viguiera cordata (Hook. & Arn.) D'Arcy. 2n = c. 40 (c. 20 II + 01 f): Chiapas, P 2539; 2n = c. 68 (c. 34 II): Oaxaca, P 3559.
Most species of Viguiera s.s. have chromosome numbers based on 17 and our count of 2n = c. 20 II + 01 f for Panero 2539 is unusual for a viguiera. We are in good company, however, because Turner and Flyr (1966)
reported "n = 21" and Jansen and Stuessy (1980)
reported 4445 "bivalents" for plants identified as Hymenostephium cordatum (a synonym of V. cordata). Robinson et al. (1981)
reported two counts of "n = 40" and "n = c. 40" for V. cordata. Yet, Strother (1983)
reported 2n = c. 51 II (hexaploid on x = 17) and Sundberg, Cowan, and Turner (1986)
reported 2n = c. 34 II (tetraploid on x = 17) for V. cordata. The species, or the "Hymenostephium complex," clearly merits cytological and taxonomic review.
Viguiera grammatoglossa DC. var. huajuapana Panero & Villaseñor. 2n = c. 34 (c. 17 II): Oaxaca, P 6141. F, I-89 (three reports, all told, all as V. grammatoglossa without indication of variety).
Wedelia acapulcensis Kunth var. cintalapana Strother. 2n = c. 22 (c. 11 II): Chiapas, P 4478.
Strother (1991)
reported 2n = c. 11 II for W. acapulcensis var. parviceps (S. F. Blake) Strother and 2n = 26 (12 II + 1 II B's) for W. acapulcensis var. acapulcensis; Strother and Panero (1994)
reported 2n = 24 II for W. acapulcensis var. ayerscottiana (B. L. Turner) Strother. Counts of 2n = 22 have been recorded for species of Wedelia closely related to W. acapulcensis (Strother, 1991
).
*Wedelia hintoniorum B. L. Turner. 2n = 22 (11 II): Oaxaca, P 3496; 2n = c. 22 (c. 11 II): México, P 7352.
Counts of 2n = 22 have been recorded for species of Wedelia closely related to W. hintoniorum (Strother, 1991
).
| FOOTNOTES |
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[ed.]. 1977 Index to plant chromosome numbers for 1973/74. Regnum Vegetabile 96: 1257
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[ed.]. 1968 Index to plant chromosome numbers for 1966. Regnum Vegetabile 55: iiv + 1126
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. 1989 Chromosome numbers in Thymophylla (Compositae: Tageteae). Sida 13: 351358
. 1991 Taxonomy of Complaya, Elephandra, Iogeton, Jefea, Wamalchitamia, Wedelia, Zexmenia, and Zyzyxia. Systematic Botany Monographs 33: 1111
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Sundberg, S., C. P. Cowan, and B. L. Turner. 1986 Chromosome counts of Latin American Compositae. American Journal of Botany 73: 2238
Torres, A. M. 1969 Revision of the genus Philactis (Compositae). Brittonia 21: 322331[CrossRef][ISI]
Turner, B. L., J. H. Beaman, and H. F. L. Rock. 1961 Chromosome numbers in the Compositae. V. Mexican and Guatemalan species. Rhodora 63: 121129
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, M. Powell, and R. M. King. 1962 Chromosome numbers in the Compositae. VI. Additional Mexican and Guatemalan species. Rhodora 64: 251271
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