2 PHYTOLOGIA An international journal to expedite plant systematic, phytogeographical and ecological publication Vol.70 February 1991 No.2 CONTENTS 3 G.L. NESOM, Taxonomy of Jsocoma (Compositae: Astereae) ....69 S.D. JONES & A.M. REZNICEK, Carex bicknellii ‘‘Bicknell’s Sedge,’ new in Texas and a key to species of section ET A ORL IR OGL a OR SA Eat eye ee ne Oe eee 115 -M.H. MACROBERTS & B.R. MACROBERTS, The distribution of Sarracenia in Louisiana, with data on its abundance in the WWESPCELUT TLE RIE PS SEH M br. uy hha Soh aages meinpasnn’ Jody ors vaew en ¥9 119 B.L. TURNER, Recension of the Asplundianthus group of Leg eO PT) Tey RI ee SON. alee Meat oT Re a yy hy A EE Ne ee 126 - B.R. MACROBERTS & M.H. MACROBERTS, Floristics of three DOSS UV WESLEED: LOUMIATIA rely ta: wot caepanensh res) «saspng has tosh Ae. b35 : B.L. TURNER, Two new species of Verbesina (Asteraceae) eke POMP MELECEO MIE SIEG oY oye cere ee oehew ued occaee coectaka bdo eas 142 LIRRARY “ ben 2 x —* .y a APR 75 7831 Published by Michael J. Warnock 185 Westridge Drive Huntsville, Texas 77340 U.S.A. PHYTOLOGIA is printed on acid free paper. eee SE —<— - —ti‘(i‘i i; ’”. PHYTOLOGIA (ISSN 00319430) is published monthly by Michael J. Warnock, 185 Westridge Drive, Huntsville, TX 77340-8916. Second Class postage at Huntsville, TX. Copyright ©1990 by PHYTOLOGIA. Domestic individual Subscription (6 issues): $18.00. Domestic instiiutional subscription (6 issues): $20.00. Foreign and/or airmail postage extra. Single copy sales: Current issue and back issues volume 67 to present, $3.50; Back issues (previous to volume 67), $3.00 (add $.50 per copy postage and handling US [$1.00 per copy foreign]). Back issue sales by volume: $17.00 per volume 42-66 (not all available as complete volumes); $21.00 per volume 67-present; add $2.00 per volume postage US ($4.00 per volume foreign). POSTMASTER: Send address changes to Phytologia, 185 Westridge Drive,Huntsville, TX 77340-8916. j Phytologia (February 1991) 70(2):69-114. TAXONOMY OF JSOCOMA (COMPOSITAE: ASTEREAE) Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT The genus Jsocoma comprises sixteen species, three of which in- clude varietal taxa. Five previously undescribed species are recognized: I. azteca sp. nov., I. felgeri sp. nov., I. humilis sp. nov., I. telua- cana sp. nov., and I. tornentosa sp. nov.. The plants of the Pacilic coast previously identified as /. veneta (Kunth) E. Greene are recog- nized as I. menziesii comb. nov., including one previously undescribed infraspecific taxon, var. diabolica var. nov., and others requiring new varietal combinations: var. decumbens, var. sedoides, var. triden- tata, and var. vernonioides. Two new combinations are proposed within J. acradenia (E. Greene) E. Greene, var. bracteosa and var. eremophila, and one within J. coronopifolia (A. Gray) E. Greene, var. pedicellata. Lectotypes are designated for some of the taxa, and dis- tribution maps and keys are provided for all of the currently recognized ones. KEY WORDS: Isocoma, Haplopappus, Asteraceae, Astereae, México Isocomais a genus of sixteen species primarily endemic to northern México and the southwestern United States. The plants are small, usually glutinous shrubs with discoid heads usually in compact corymbs, goblet shaped corollas, achenes with a pappus of bristles, and a base chromosome number of z = 6. The genus was proposed originally by Nuttall, but the taxonomic structure of the group was provided primarily by E.L. Greene in two short discussions (1894, 1906). Hall (1907) originally treated these plants as [socoma but later (1928), in the only previous study of the whole group, submerged it as a section of Haplopappus. Some authors of floras of regions in the western United States have maintained the nomenclature in Haplopappus, but others (e.g., Jepson 1925; Shinners 1950; Correll & Johnston 1970; Turner 1972) have accepted [socoma as a separate genus. The study presented here is adjunct to the preparation of a treatment of /socoma for the forthcoming “Asteraceae of Mexico” by Turner & Nesom, since all but four of the species occur in Mexico. 69 70 PHYTOLOGIA volume 70(2):69-114 February 1991 Hall considered Jsocoma to be most closely related to Hazardia ( Haplopap- pus sect. Hazardia). Clark (1979), however, placed Hazardia closest to species of Haplopappus sect. Polyphyllus in South America as well as to some species of Machaeranthera and Xylorhiza. While he speculated that Hazardia brick- ellioides (S.F. Blake) Clark, an anomalous element in the genus with n = 6 pairs of chromosomes, might represent a “distant link” to /socoma, he sum- marized other opinions and affirmed that Jsocoma is more closely related to other North American genera with a base chromosome number of z = 6. Iso- coma, Grindelia, Prionopsis, Xanthocephalum, Olivaea, Stephanodoria, and the “Haplopappus phyllocephalus DC. group” form a group of closely related taxa (the “Xanthocephalum group;” Nesom, et al., submitted). Within this group of species, Hartman & Lane (1991) have documented the occurrence of natural hybrids between the widely divergent J. veneta (Kunth) E. Greene and X. humile Benth. On the basis of chromosome number, morphological similarity, and artificially produced hybrids, Jackson (1966) and Jackson & Dimas (1981) considered the H. phyllocephalus group to be a part of Isocoma (as Haplopappus sect. Isocoma); Hartman (see 1990 for summary and other references) has considered the H. phyllocephalus group to be a separate genus, although he has not formalized this view, and it is also excluded from [so- coma in the present treatment. Compared to /socoma, the former species are annual herbs with much broader, campanulate heads in a loosely associated capitulescence and achenes with nonresinous nerves. Shinners inexplicably transferred Aster palmeri A. Gray (= Ericameria austrotezana M.C. Johnston) to Isocoma, where it is conspicuously anomalous, but that species is now included in the genus Xylothamza (Nesom, et al. 1990). Diploids and tetraploids have been reported in three species of /socoma (IJ. acradenia |E. Greene] E. Greene, J. plurifolia (Torr. & A. Gray] E. Greene, and I. menziesii [Hook. & Arn.] Nesom, see references below), but the taxonomic significance of this, if any, has yet to be clarified. The variation patterns and the taxonomy of Jsocoma are most complex along the Pacific coast. In J. menziesi of that area, there are numerous in- tergrading forms to which names have been applied, and a more intensive and field oriented study may ultimately provide a more precise taxonomy for these plants. Many of the habitats of J. menziesii, however, have been eliminated by human expansion. With regard to the Californian plants of Jsocoma, Hall (1907) noted that “Endless forms ...might be described from the abundant material at hand, but they could be characterized only by various combinations of characters well known to be inconstant.” He referred all of these “endless forms” to two variable taxa of a single species (var. vernontoides (Nutt.) Jep- son and var. acradenia |E. Greene] Hall of J. veneta) but subsequently (1928) recognized each as a different species. Other students of the Californian plants have recognized a number of varieties in reach of two species, here identified as I. acradenia and J. menziesii. Nesom: Taxonomy of Isocoma 71 Most of the species of Jsocoma have geographic ranges allopatric, or at least parapatric, with all others (z.e., only a single species is found in any given area), although correlated geological or topographic boundaries are difficult to discern (Maps 1-6). This is a remarkable phenomenon, the distribution maps giving the appearance of a closely fitted patchwork quilt. Exceptions to this are noted in the discussions following several of the species. The occurrence of hybrids at contiguous points of their geographic ranges are noted for a number of the species. In most instances it seems clear that species of /socoma closely associated geographically also are most closely related to each other, but without further and more detailed study, the relatively few characters useful in distinguishing the taxa cannot support broader hypotheses regarding interspecific phylogenetic relationships. Type specimens for new taxa named by E.L. Greene are divided for the most part between US and ND-G. Except where noted in the discussions below, those chosen as lectotypes bear an annotation by Greene as “Type.” The species are treated in alphabetical order. Isocoma Nutt., Trans. Amer. Phil. Soc., ser. 2, 7:320. 1840. TYPE: /socoma vernonioides Nutt., Trans. Amer. Phil. Soc., ser. 2, 7:320. 1840. Hap- lopappus sect. Isocoma (Nutt.) Hall, Carnegie Inst. Washington Publ. 389:36. 1928. (H)Aplopappus sect. Aplodiscus DC., Prodr. 5:350. 1836. LECTO- TYPE (designated here): (H)Aplopappus discoideus DC. (= Iso- coma veneta [Kunth] E. Greene). DeCandolle also included one other species in his sect. Aplodiscus: (H)Aplopappus ramulosus DC. (= Baccharis pteronioides DC.). Perennial subshrubs from a woody base, often glutinous, the stems com- monly erect and few branched, sometimes decumbent. Leaves primarily oblance- olate, entire or with spinulose tipped teeth or lobes, uninerved, glandular punctate to papillate or stipitate glandular, usually glutinous. Heads discoid, turbinate to campanulate, sessile to short pedicellate in terminal corymbs, rarely solitary; phyllaries strongly graduated; receptacle deeply alveolate, rarely shallowly so. Corollas yellow, narrow, abruptly ampliate, the tube with at least a few glandular hairs, the lobes erect, usually 0.5-1.8 mm long, the tube elongating at maturity and elevating the corolla at anthesis, outer corollas prominently bent or leaning outward; anthers inserted at midpoint in the tube; style appendages narrowly triangular. Achenes moderately to densely sericeous, broadly to narrowly turbinate, mostly 4-10 ribbed, the ribs some- times very thick and resinous; pappus of numerous, thick, barbellate bristles of uneven length. Base chromosome number, z = 6. volume 70(2):69-114 February 1991 72 PHYTOLOGIA I. acradenia o var. acradenia © var. bracteosa @ var. eremophila 4 I. arguta @ |. felgeri 4 I. tenuisecta Map 1. Distribution of Isocoma acradenia, I. arguta, J. felgert, and /. tenut- secta. Nesom: Taxonomy of Isocoma 73 gw lI. azteca 4 I. humilis d) © I. plurifolia O I. rusbyi @ I. tomentosa \ IX Map 2. Distribution of /socoma azteca, |. humilis, 1. plurifolia, J. rusbyi, and 1. tomentosa. February 1991 volume 70(2):69-114 PRY T:O:L,06PA 74 eo O @ @ % IIpuowUInIp ‘| Blel[aoIped “eA BI[OJIdOUOIOD “IBA BI[OJIdOUOIOD *] v @ O Map 3. Distribution of Isocoma coronopifolia and J. drummondii. Nesom: Taxonomy of Isocoma 75 : < os % oO |. hartwegii | & Of. ,tehuacana e J. veneta Map 4. Distribution of Isocoma gypsophila, 1. hartwegzi, I. tehuacana, and J. veneta. 76 PHY TOLOGIA volume 70(2):69-114 February 1991 I. menziesii @ var. decumbens I. menziesii @® var. menziesii 4 var. tridentata Map 5. Distribution of [socoma menziesii (var. decumbens, var. menziesii, and var. tridentata). Taxonomy of Isocoma 77 Nesom: I. menziesii @ var. diabolica x var. sedoides @® var. vernonioides Map 6. Distribution of Jsocoma menziesii (var. dzabolica, var. sedoides, and var. vernonioides). 78 PRYTOLOGIA volume 70(2):69-114 February 1991 KEY TO THE SPECIES OF ISOCOMA 1. Plants of California (U.S.A.) and Baja California Norte and Sur (México). » hbciacatraevbe wired bile ea . .00.5 0... .sd040425e 0 oe I, acradenia 8. Leaves narrowly oblong to oblanceolate-oblong or linear, the margins entire to shallowly toothed, not pinnatifid. .................. ( 13) 8’ Leaves pinnatifid, ..........0 00.000. ces noe ee (9) 9. Plants sometimes glandular but otherwise completely glabrous. .... (11) Nesom: Taxonomy of Isocoma 79 9’ Plants minutely hispidulous or very sparsely puberulous, at least on the WP PEL SUEMISS SC cieiicrs ders Pera ae ee ol meets ME. CREED STO Ed cael (10) 10. Heads 8-12 (-15) flowered; involucres 4.0-5.5 mm long, 2.0-2.8 mm wide; northern Sonora, Arizona, New México. ....... I. tenuisecta 10’ Heads 13-22 flowered; involucres 5.0-6.7 mm long, 4.0-5.0 mm wide; central¢Mexiconr4 209% =. 2S aieike, Reet seats Paina. 2 I. hartwegu 11. Heads 12-15 flowered, 2.5-4.0 mm wide; Texas and northeastern Mexico (Coahuila, Nuevo Leon, Tamaulipas). ................. I. coronopifolia 11’ Heads 19-25 flowered, 5.0-7.5 mm wide; Arizona and New Mexico. Fe) Ne Ae aE hs Ne tem a Si ea high Fal oe a3 I, azteca 12. Stems and leaves sparsely hispidulous to villosulous or tomentose; leaves prominently toothed.” <.2.55'25 ais 2s oo oO. ee ote cee tes (17) 12’ Plants glabrous or the leaves merely ciliate; leaves entire or very shallowly toothed near thetapex:. © ..26s8/G,. 222/52). Steer. «5 (13) 13. Plants of New Mexico, Arizona, Colorado, and Utah. ............. (16) 13’ Plants of Vexas*and northern ‘Nlexico. 4, 2920 009 5 Bio AS on oars (14) 14. Heads 22-34 flowered, the involucres 4.5-5.5 (-7.0) mm wide, mostly 6F528 Camm longs ss See ek bbe Sane eaides sods I. drummondu 14’ Heads 8-15 flowered, the involucres 2.5-4.0 mm wide, mostly 4-6 CIVIL: LOM Bog Aas bec RW aah cs ETE ROG 2 2s =o 0.6 5 ae Oe (15) 15. Leaves sometimes stipitate glandular but never glutinous, minutely hispid- ulous along the margins, sometimes on the lamina as well; involucres 4-06.20 emo gs sae soe yale wee os Se thee aravslefe e's a oo atm SE a I. plurifolia 15’ Leaves never stipitate-glandular, almost always glutinous, completely glabrous; involucres: 5-% mim long:)42...4 ee. hk hese oe I. coronopifolia 16. Leaves usually minutely ciliate, stipitate glandular or with sunken, papillate glands but not glutinous; heads (8-) 11-15 flowered, the involucres 4.0-5.2 mm long, 2.5-4.0 mm wide; corolla lobes 1.1-1.8 mm long, equaling the throat length; achenes 1.5-1.9 mm long, the ribs Notmapicallys “horned.” _.... 6.00.5. 4se0c s. SUS eee I. gypsophila 18’ Heads 19-28 flowered; involucres 6-7 mm wide; phyllaries with nar- TOW SCALiOUS MALgINS! ~ 2. o... 6 Wisse «00 /salde eae eee I. humilis 19. Stems leaves, and phyllaries puberulent-tomentose. ....... I. tomentosa 19’ Stems and leaves minutely hispidulous to short puberulent; phyllaries GIADCOUG. ae. d5 iia Fate Be so bls ce 2 noses pig aye = ses oe in ee er (20) 20. Stems and leaves densely short puberulent; involucres 6-7 mm wide; corolla tubes sparsely sericeous; achenes 2.8-4.0 mm long. RE Se AC a A ee EES I. tehuacana 20’ Stems and leaves minutely hispidulous to nearly glabrous; involucres 4-5 mm wide; corolla tubes glabrous; achenes 1.6-2.8 mm long. aWbiiae Ceade bel sede dee valecs ce sos Ronee 2 I. veneta 1. Isocoma acradenia (E. Greene) E. Greene Small shrubs 0.6-1.5 m tall, with markedly whitish stems. Stems and leaves glabrous to minutely hispidulous. Leaves entire to pinnatifid or toothed, punc- tate papillate but rarely glutinous, often in axillary fascicles. Heads turbinate to narrowly obconic, 8-27 flowered; phyllaries narrowly oblong, glabrous or rarely somewhat papillate, yellowish white and indurated except for the green- ish or brownish apex, the apex bearing a single, thick, subepidermal resin pocket nearly as wide as the bract, or sometimes a central pocket and 2- several, smaller, lateral ones. Corollas 5-7 (-8) mm long. Achenes 8-10 ribbed, (2.0-) 3.0-3.5 (-4.0) mm long. Chromosome numbers, n = 6, 12 pairs (see below). Isocoma acradenia is distinctive in its whitish stems and narrow, whitish indurated, phyllaries with an apical resin pocket (or pockets). Three inter- grading varieties can be recognized. Key to the varieties 1. Heads 20-27 flowered; phyllary apices spinulose-aristate. .. var. bracteosa Nesom: Taxonomy of [socoma 81 1’ Heads 8-20 flowered; phyllary apices usually rounded and not at all spinu- lose, sometimes with weakly developed aristae. ..................-. (2) 2. Leaves entire, rarely the lower toothed or lobed; heads 8-18 flowered. A le, a a Gott RAR Clee de oP Fie Sh ata IRS Pe Pe var. acradenia 2' All leaves toothed to pinnatifid; heads 12-20 flowered. 5 SEER OE AMER OREN et = LE Ot ee var. eremophila a. Var. acradenia Isocoma acradenia (E. Greene) E. Greene var. acradenia BASIONYM: Bige- lovia acradenia FE. Greene, Bull. Torrey Bot. Club 10:126. 1883. LEC- TOTYPE (designated here): UNITED STATES. California: Mohave Desert, 6 Sep 1881, FE. Greene s.n. (ND-G!; Isolectotypes: CAS, GH!). Aster acradenius (EK. Greene) O. Ktze., Rev. Gen. 318. 1891. Isocoma veneta (Kunth) E. Greene var. acradenia (E. Greene) Hall, Univ. Cali- fornia Publ. Bot. 3:64. 1907. Haplopappus acradenius (E. Greene) S.F. Blake, Contr. U.S. Natl. Herb. 25:546. 1925. The ND-G specimen, a duplicate ez CAS, is annotated by Greene as Bigelovia acradenia Greene but not specifically as the “Type.” Isocoma limitanea Rose & Standl., Contr. U.S. Natl. Herb. 16:18. 1912. TYPE: MEXICO. Sonora: village of Sonoyta, 14 Nov 1907, D.T. MacDougal 14 (HOLOTYPE: US!). Leaves 2-3 cm long, entire, or sometimes in Arizona the lower toothed or lobed. Heads 8-18 flowered, the involucres 5-7 mm high; phyllaries not spinulose-aristate. Achenes 2.0-2.4 mm long. [Putative differences between this and var. eremophila (EK. Greene) Nesom in head size and flower number do not hold, although both range larger in var. eremophila. Chromosome number, n = 12 pairs (Raven, et al. 1960; Urbatsch 1974). Mojave Desert of California, adjacent Nevada, and Arizona, coastal Sonora; most commonly in salt shrub communities, often with Larrea, 650-1100 m; (June-) September-November. See comments regarding putative intermediates with /socoma tenuisecta I. Greene following that species. b. Var. bracteosa (E. Greene) Nesom, comb. nov. Isocoma acradenia (E. Greene) E. Greene var. bracteosa (E. Greene) Nesom. BASIONYM: Isocoma bracteosa E. Greene, Leafl. Bot. Observ. Crit. 1:170. 1906. LECTOTYPE (designated here): UNITED STATES. 82 PHY TO.LO G EA volume 70(2):69-114 February 1991 California: Tulare Co., 27 Aug 1889, C.S. Sheldon (US!). Haplopappus acradenius (E. Greene) S.F. Blake subsp. bracteosus (E. Greene) Hall, Carnegie Inst. Washington Publ. 389:233. 1928. Haplopappus acrade- nius (E. Greene) S.F. Blake var. bracteosus (E. Greene) McMinn, /Ilustr. Man. Calif. Shrubs 574. 1939. Lower leaves usually shallowly toothed, those on the upper (10-) 15-20 cm of the stem spreading to slightly deflexed, mostly 5-15 mm long, entire to serrulate. Heads 20-27 flowered, the involucres 7-8 mm high; phyllaries dis- tinctly spinulose-aristate at the apex. Achenes 2.5-3.0 mm long. Chromosome number, n = 6 pairs (Raven, et al. 1960). California, at the northwestern portion of the range of the species, primarily in the San Joaquin Valley, but extending, at least historically, to San Francisco (see comments below); August-October. Var. bracteosa (E. Greene) Nesom comprises a geographically and morpho- logically distinct group of populations within /socoma acradenia. Compared to var. acradenia, to which it is geographically adjacent, var. bracteosa has heads with a greater number of flowers and phyllaries with spinulose-aristate apices. The putative distinction regarding distribution and sizes of cauline leaves, emphasized in previous treatments, is not at all consistent, since many plants of var. acradenia are habitally identical to those of var. bracteosa. The two varieties differ in relatively few features, but intermediates between them apparently are rare in the zone of contact. A collection from the “Bay of San Francisco,” made by the Wilkes Expe- dition (no. 1668, NY, US) has been annotated and mapped as var. bracteosa, but in their few flowered (13-14) heads and more extensively chlorophyllous phyllary apices, they are more like Jsocoma arguta E. Greene. They have entire leaves, however, and cannot be placed with the latter species. c. Var. eremophila (E. Greene) Nesom Isocoma acradenia (E. Greene) E. Greene var. eremophila (E. Greene) Nesom. BASIONYM: Isocoma eremophila E. Greene, Leafl. Bot. Ob- serv. Crit. 1:170. 1906. LECTOTYPE (designated here): UNITED STATES. California: San Diego Co., southwestern part of the Col- orado Desert, 1 Nov 1890, C.R. Orcutt 2223 (US!; Isolectotype: GH!). Haplopappus acradenius (E. Greene) S.F. Blake subsp. eremophilus (E. Greene) Hall, Carnegie Inst. Washington Publ. 389:233. 1928. /Hap- lopappus acradenius (E. Greene) S.F. Blake var. eremophila (E. Greene) Munz, Man. S. Calif. 523. 1935. Isocoma acradenia (E. Greene) E. Greene subsp. eremophila (E. Greene) Beauchamp, Phytologia 59:437. 1986. Nesom: Taxonomy of Isocoma 83 Leaves mostly 2.5-5.0 cm long, pinnatifid to pinnately toothed with 2-4 (-5) pairs of narrow lobes or teeth. Heads 12-20 flowered, the involucres 6-8 mm high, phyllary tips with a resin pocket or sometimes merely glandular-papillate but swollen, not spinulose aristate. Achenes 3.5-4.0 mm long. Chromosome number, n = 6 pairs (De Jong & Montgomery 1963). Baja California Norte, northwestern Sonora, southern California, south- western Arizona, southern Nevada, Utah(?); sandy soil, alkaline or salt flats, desert dunes, often with Saguaro-Prosopis-Larrea, Larrea, Atriplex, Yucca, Pachycereus, or mixed scrub; (-15-) 0-900 (-1050) m; (May-) August-November (-December). A single collection of Jsocoma acradenia var. eremophila is la- beled as having been made in Utah (no other locality data, 1870, Palmer 21-GH, US), but this species has not been recorded from Utah in more recent floristic treatments. Var. eremophila is distinguished from var. acradenza by its shallowly lobed to serrate crenate leaves. More strongly pinnatifid leaved plants occur along the coast of northeastern Sonora as well as in northwestern Arizona and scat- tered in southern California. For the most part, the two varieties are clearly separated in geography; plants in the apparently isolated system of var. er- emophila in Nevada do not differ from others in the range. In some areas (e.g. in southwestern Nevada), intergradation between var. eremophila and var. acradenia appears to be nearly complete, but in others (e.g. western San Bernadino Co., particulary southeast of Barstow) the two morphological forms appear to be contiguous with little or no intergradation. In the latter area, each of the two varieties has been reported at a different ploidy level. In the area around Dixieland (Imperial Co., California) there is a distinctive population system of var. eremophila with heads borne on long pedicels. In a few plants of otherwise typical morphology scattered through the range, the resin pockets characteristic of the phyllary apices may be reduced or absent, the apices appearing merely punctate glandular, though often still remaining thickened. These are particularly common in Riverside Co. in the area of Hemet and San Jacinto. The plants from northwestern Sonora consistently tend to have less strongly developed resin pockets and are also similar in their linear leaves to the putative intermediates between var. acradenia and Isocoma tenuisecta (see discussion following the latter). 2. Isocoma arguta E. Greene Isocoma arguta E. Greene, Man. Bot. San Franc. Bay Reg. 175. 1894. LEC- TOTYPE: UNITED STATES. California:: Solano Co., Morning Light, [“subsaline plains east of the Vaca Mountains,” in the protologue] 16 Sep 1891, W.L. Jepson s.n. (ND-G!; Isolectotypes: JEPS, US-2 sheets!; Probable isolectotype [without collection data]: ND-G!). /socoma veneta 84 PHY POLO GPA volume 70(2):69-114 February 1991 (Kunth) E. Greene var. arguta (E. Greene) Jepson, Fl. W. Mid. Calif. 500. 1901. Haplopappus venetus (Kunth) S.F. Blake var. argutus (E. Greene) Keck, Aliso 4:103. 1958. Neither the US nor ND-G specimens is specifically marked as “Type.” Shrubs 1-3 dm tall, glabrous or the stems lightly villous near the very base. Leaves mostly 1.0-2.5 cm long, 3-7 mm wide, only slightly reduced in size up- wards, serrate to pinnatifid with 3-5 pairs of aristate teeth or lobes arranged from base to apex, viscid, glabrous to minutely hispidulous and stipitate glan- dular. Heads short pedicellate in dense clusters, 10-13 flowered, turbinate with a narrowly acute base, the involucres 5-7 mm high, 4-5 mm wide; phyllaries glutinous, with a sharply demarcated, densely punctate and thickened apical . area (sometimes as a small resin pocket), spinulose-aristate at the apex, the lower 2/3-3/4 white indurated; receptacles with short, triangular alveoli with filiform apices. Corollas 4.0-5.0 mm long, the lobes deltate, 0.4-0.5 mm long. Achenes 3.0-3.5 mm long, densely sericeous, with ca. 6-8 resinous ribs. California, apparently localized in Solano and Contra Costa counties; low hills, subsaline plains; August-October. Isocoma argutais distinctive in its short stature, glabrous, small, thickened, regularly toothed to short lobed leaves, few flowered heads, and narrowly ob- long or obovate, white indurated phyllaries with small, sharply defined apical areas and spinulose-aristate tips. It has been treated as a variety of J. men- ziesit, but it is clearly more similar to /. acradenia var. bracteosa in its phyl- lary morphology. Further, in its inland distribution, /. argutais geographically more similar to and nearly continuous with /. acradenia, adding a northern segment above the range of var. bracteosa. The plants of /. arguta, however, are consistent in their morphology, different from those of var. bracteosa in their leaf size and margins and their fewer flowered heads; plants that could be considered intermediate with any other taxon have not been observed. 3. Isocoma azteca Nesom. Isocoma azteca Nesom, sp. nov. TYPE: UNITED STATES. New Mex- ico: McKinley Co., 3.4 mi by road SW of Ojo Encino on slopes below NE-facing “badlands,” 11 Aug 1976, R. Spellenberg, et al. 4344 (HOLO- TYPE: TEX!; Isotypes: NMC!, NY!). Isocomae rusbyi E. Greene similis sed foliis pinnatisectis differt. Shrubs 2-4 dm tall, forming clumps up to 6 cm wide. Stems glabrous, yellowish green to whitish, rarely slightly villous. Leaves glabrous, punctate, slightly resinous, 2-5 cm long, 4-12 mm wide, narrowly oblong to narrowly oblanceolate, shallowly to deeply pinnatifid with 3-8 pairs of aristate tipped Nesom: Taxonomy of Isocoma 85 lobes evenly arranged from base to apex. [leads campanulate, basally rounded, 18-23 flowered, the involucres 7-8 mm high, 5-7 mm wide; inner phyllaries with broad scarious margins, the apices green, barely to prominently punctate. Corollas 5-6 mm long, the tube 3.0-3.5 mm long, the lobes triangular, 0.7-1.0 mm long. Achenes 2-4 mm long, with 5-8 thick resinous ribs, forming small “horns” at the apex, densely sericeous. Chromosome number, n = 6 pairs (Jackson 1959, as Haplopappus hartwegz |A. Gray] E. Greene). Northeastern Arizona and northwestern New Mexico; river edges, slopes, sandy to clay soil, gypseous or saline, commonly with Atrzplez, badlands in pinyon pine-juniper woodlands; 1500-1800 m; July-September. Additional collections examined: UNITED STATES. Arizona: Apache Co., northeastern Arizona (Moki Reservation) and Little Colorado River, 1 Aug-5 Sep 1896, Hough 114 (US); Apache Co., Hopi Ind. Res., 5000 ft, 7 Aug 1937, Whiting 854/2693 (ARIZ). New Mexico: Cibola Co. [labeled Valencia Co.], extreme W part of the county, near the Arizona border, 16 Aug 19838, McIntosh s.n. (NY); San Juan Co., Botanical Station, Gallegos Wash, Horn Canyon Quadrangle, 5340 ft, deep sand at edge of wash, 24 Jul 1974, Blanken- horn 89B (ARIZ); San Juan Co., 10 mi S of area complex, Navajo Mine, 4 Corners Power Plant, 16 Aug 1986, Kass 2567 (NY); San Juan Co., Navajo Project area 13 mi S of Farmington, 6000 ft, 28 Jul 1978, Pase 2328 (TEX); San Juan Co., 5.5 mi by road §S of Bisti, first draw S of De-na-zin Wash, 5840 ft, 20 Aug 1976, Reztzel & McKinney 4390 (NMC, NY); San Juan Co., flood plain of Chaco River ca. 7 mi SE of Shiprock, 9 Sep 1977, Spellenberg 4856 (NMC, NY, TEX); San Juan Co., ca. 17 mi S of Fruitland, 15 mi NNE of Burnham Trading Post, in Cottonwood Arroyo at Cottonwood Springs, 10 Sep 1977, Spellenberg 4870 (NMC, TEX); San Juan Co., ca. 10 air mi SE of Shiprock, E side of Hogback near S end of Ist ridge S of San Juan River, clay soil, 10 Aug 1981, Spellenberg & Ward 6091 (NMC, NY); San Juan Co., 6 mi N of Farmington, 1 mi S of Jackson Lake, W of Hwy 170, 26 Jul 1977, Welsh, et al. 15645 (NY). The plants from Cibola Co., New Mexico (AMcJ/ntosh s.n.) are atypical in their vestiture (stems lightly villous and leaves minutely stipitate glandular) but they are otherwise typical of Jsocoma azteca Nesom. On the labels of his collection 4856 of Isocoma azteca, Richard Spellenberg noted that there were “nearly entire-leaved to (more commonly) pinnatifid- leaved plants” at the single locality, and examples of both morphological forms are mounted on each of the three duplicate sheets cited above. The branches, however, bear either nearly entire leaves or deeply pinnatifid ones, and in- termediates, if present at all, must have been less common than these two extremes in morphology. As interpreted here, the population at this locality comprises individuals of both species, which meet there at the margin of each of their geographic ranges (Map 2). In Apache Co., Arizona, typical plants of both species have been collected in relatively close proximity (J. rusbyz E. 86 PHYTOLOGIA volume 70(2):69-114 February 1991 Greene at Tanner’s Crossing of Little Colorado, 13 Nov 1899, Ward 7-US; J. azteca, Hough 114). One plant of /. rusby: with few lobed lower leaves (the upper entire) has been collected near the western edge of its geographic range (Coconino Co., near Tuba, 15-31 Jul 1920, Clute 120-NY), but all other plants studied have completely entire leaves. Should further collections demonstrate that a broader area of intermediates exists between these two taxa, they may be combined into a single species. See additional comments regarding possible intermediates with J. plurifolia following that species. The relationship of Jsocoma azteca to [. rusbyi appears to be analogous to that of J. hartwegiz to I. veneta. In both cases, the two taxa appear to be sister species but each member of the pair has a sharply defined geographic range, allopatric or parapatric with its close relative, and few if any intermediates are formed between them. The relationship of /. acradenia var. acradenza (with entire leaves) to var. eremophila (with toothed to pinnatifid leaves), that of I, menziesii var. menziesw to var. vernonioides (Nutt.) Nesom, and that of I. coronopifolia (A. Gray) E. Greene var. pedicellata (E. Greene) Nesom to var. coronopifolia is also similar to that of J. rusbyi to J. azteca. In each of these three pairs of taxa, however, there is considerable intergradation between them, at least along part of their contiguous ranges, and the rationale for maintaining them as separate species is weakened accordingly. 4. Isocoma coronopifolia (A. Gray) E. Greene Shrubs 0.3-1.2 m tall. Leaves pinnatifid with 1-3 pairs of spreading, linear, spinescent tipped lobes, less commonly entire or only the lowermost leaves pinnatifid, the blade 1.5-3.0 (-5.0) cm long, 0.5-2.0 (-2.5) mm wide, glutinous, completely glabrous, somewhat fleshy and drying with a characteristic “alliga- tor skin” pattern. Heads 12-15 flowered, usually narrowly cuneate at the base, the involucres 5.0-6.0 (-7.0) mm long, 2.5-4.0 mm wide (pressed); phyllary apex markedly thickened, somewhat glandular, thickened, rarely punctate, tips of the innermost mostly flat and not glandular; receptacles with lanceo- late alveoli. Corollas 4.5-6.0 mm long, the tube 2.5-3.5 mm long, lobes 0.8-1.0 mm long, 1/3-2/5 the length of the limb. Achenes 1.8-2.2 mm long, with 4-6 resinous ribs, sometimes with additional, interspersed, thinner nerves, lightly sericeous. Chromosome number, n = 6 pairs (Turner, Powell, & Watson 1973, as I. heterophylla [A. Gray| E. Greene; Urbatsch 1975; Powell & Powell 1977, 1978). Chihuahua?, Coahuila, Tamaulipas, Nuevo Leon, and Texas; gypsum or alkaline flats, calcareous substrate (Texas), dunes or sandy habitats, matorral or shrublands; 250-1100 m; (May-) June-October. The single collection of this species attributed to Chihuahua (var. coronopifolia, Schott s.n. - US) has no locality data other than “roadside.” Map 3 shows the ranges of the two varieties, which are essentially similar in habitat. Nesom: Taxonomy of Isocoma 87 Key to the varieties of /. coronopifolia ft. At least the upper leaves-entire. 22/5... . 6.0 denns essences var. pedicellata MoAT leaves pin atiha. 35. e< bdaets ora seas «accuses sae var. coronoptfolia a. Var. coronopifolia Isocoma coronopifolia (A. Gray) E. Greene var. coronopifolia. BASIONYM: Linosyris coronopifolia A. Gray, Pl. Wright. 1:96. 1852. TYPE: UNITED STATES. Texas: [Kinney Co., probably at Brackettville], Sep 1849, C. Wright 289 (HOLOTYPE: GH!, NY-photo!, US-photo!); not (H)Aplo- pappus coronopifolius DC., 1836. Bigelovia coronoprfolia (A. Gray) A. Gray, Proc. Amer. Acad. Arts 8:638. 1873. Isocoma coronopzfolia (A. Gray) E. Greene, Erythea 2:111. 1894. b. Var. pedicellata (KE. Greene) Nesom. Isocoma coronopifolia (A. Gray) E. Greene var. pedicellata (E. Greene) Nesom, comb. et stat. nov. BASIONYM: Isocoma pedicellata E. Greene, Leafl. Bot. Observ. Crit. 1:170. 1906. LECTOTYPE (designated here): UNITED STATES. Texas: [LaSalle Co.], “Guadaloupe, a mail station 105 mi SW of San Antonio,” 17-18 Apr 1879, 2. Palmer 486 (US!; Isolec- totypes: GH!, NY!, US!). Plants of /socoma coronopifolia with entire leaves on at least the upper parts of the stems occur in two separate regions within the range of the species (Map 3). Although it is not clear whether or not these entire leaved population systems have arisen independently, they are sharply distinct from the typical plants with all leaves nearly pectinately dissected. For consistency with other infraspecific taxa within the genus with analogous variation in leaf morphol- ogy (see comments following /. azteca), the two are treated here as separate varieties. Plants with pinnatifid leaves on the lower parts of the stems and en- tire ones on the upper parts occur in both regions and are perhaps genetically intermediate, but they are mapped as var. pedicellata. Entire leaved plants of Jsocoma coronopifolia (var. pedicellata) can be dis- tinguished from /. plurifolia by their fleshy, completely glabrous and differently textured leaves with narrower blades, longer involucres, and their more shal- lowly cleft corolla lobes. The two taxa do not come into geographical contact, and there is no evidence of intergradation between them, although some forms of each may be superficially very similar. In Brooks Co., Texas, /. coronopifolia var. coronopifolia has been noted in label data as “growing with and distinct from” J. drummondu (Torr. & A. Gray) E. Greene (4 mi S of Falfurrias, John- ston 541482-TEX). 88 PHYTOLOGIA volume 70(2):69-114 February 199] 5. Isocoma drummondii (Torr. & A. Gray) E. Greene Isocoma drummondii (Torr. & A. Gray) EB. Greene, Erythea 2:111. 1894. BASIONYM: Linosyris drummondi Torr. & A. Gray, Fl. N. Amer. 2:233. 1842. TYPE (probable holotype): UNITED STATES. Texas: [Austin Co.?, San Felipe-see comments below], 1833-34, T. Drummond 223 (GH not located, GH-photo!; Duplicate: K). Bigelovia drummondu (Torr. & A. Gray) A. Gray, Proc. Amer. Acad. Arts 8:639. 1873. Aster berlandieri O. Ktze., nom. nov., Rev. Gen. 318. 1891. Chondrophora drummondii (Torr. & A. Gray) Heller, Contr. Herb. Franklin & Marshall College 1:101. 1895. (H)Aplopappus drummondii (Torr. & A. Gray) S.F. Blake, Contr. U.S. Natl. Herb. 23:1491. 1926. The GH specimen cited by Hall (1928), and photographed, was not relocated in the present study. Isocoma megalantha Shinners, Field & Lab. 23:24. 1955. TYPE: UNITED STATES. Texas: Karnes Co., 1.3 mi SE of Harmony School, 26 Nov 1954, J.C. Johnson 1638 (HOLOTYPE: SMU!; Isotypes: NY!, TEX!). Shrubs ca. 0.5-1.0 m tall, completely glabrous, glutinous. Leaves narrowly oblong to oblanceolate oblong, 2.0-4.5 mm wide, 1.3-5.0 cm long, entire or less commonly with 1-3 pairs of shallow teeth. Heads 22-34 flowered, campanu- late with rounded base, the involucres (5.5-) 6.5-8.0 mm long, 4.5-5.5 (-7.0) mm wide; apices of phyllaries rounded to obtuse, with a sharply demarcated and strongly glandular punctate apical area ca. 1/3-1/2 the phyllary length; receptacles weakly to strongly developed, triangular alveoli. Corollas 5.8-7.5 mm long, the lobes triangular, 0.8-1.0 mm long. Achenes 2.0-2.8 (-4.0) mm long, very lightly sericeous, with 6-9 thin, sometimes whitish ribs, these not forming apical horns. Chromosome number unknown. Southern Texas and immediately adjacent northeastern Tamaulipas; coastal habitats, brushy prairie, scrub woods, often with mesquite, sand or clay, some- times on beach dunes; 0-20 m; (March-) May-July. According to Hall (1928), one of Drummond’s collections of this species at K carries the notation “San Felipe,” but I have neither seen more recent collections from this area nor personally been able to relocate the species there, which is apparently considerably to the northeast of its primary range (the presumed type locality shown on Map 2). Drummond also collected at other sites in southeastern Texas (Geiser 1948), and it is likely that the type locality was further southwest and nearer the coast than San Felipe. Similarly, attempts to relocate populations in the area of the type locality of /socoma megalantha Shinners have been unsuccessful, as the plants there apparently have been extirpated by agriculture. Isocoma drummondii has sometimes been considered to include J. rusbyi, but the latter is widely separated from J. drummondit in range and has some- Nesom: Taxonomy of Isocoma 89 what smaller heads with shorter corollas and larger, more densely pubescent achenes with much thicker, resinous ribs. In its somewhat fleshy leaves, and achene morphology, J. drummondz is more similar to the geographically adja- cent I. coronopifolia (var. coronopifolia, with divided leaves). 6. Isocoma felgert Nesom. Isocoma felgeri Nesom, sp. nov. TYPE: MEXICO. Sonora: 5 mi by road E of town of Bahia Kino, crest of slight rise in desert flat, sandy soil, cactus scrub with Pachycereus pringlez, 19 Oct 1963, R.S. Felger 9051 (HOLOTYPE: TEX!; Isotypes: ARIZ!, GH!, MEXU!). Isocomae acradeniae (E. Greene) E. Greene similis morphologia phyllariis sed foliis profunde pinnatisectis lobis linearibus et lobis corollarum longioribus differt. Shrubs 0.5-1.2 m tall. Stems and leaves glutinous, sparsely and minutely hispidulous. Leaves mostly 1.5-3.5 cm long, deeply pinnatisect, with the cen- tral portion and lobes linear to linear oblanceolate, 3-12 mm long, strongly divergent, in 1-2 pairs mostly near the apex. Heads sessile to short pedicel- late in terminal clusters, 3.5-5.0 mm wide, 8-11 flowered; phyllaries strongly graduated in 4-5 series, white indurated thickened, all except the inner with a sharply delimited, apical resin pocket, the innermost 4.0-5.5 mm long, with thin hyaline margins and an acute apex, the outer oblong with a rounded apex. Corollas 4.5-5.5 mm long, the tube 2.5-3.5 mm long, the lobes triangular, 1.0- 1.5 mm long, ca. half the length of the limb. Achenes ca. 2 mm long, with 5-6 resinous ribs, densely sericeous. Chromosome number unknown. Additional collections examined: MEXICO. Sonora: 8 mi by road E of village of Bahia Kino, nearly flat desert plain, sandy soil, cactus scrub with Pachycereus pringlei, 29 Jan 1964, Felger 9841 (ARIZ, MEXU, NY, TEX); 8.9 mi by road E of Bahia Kino (village), desert plain, “cardonal” cactus scrub with Pachycereus pringlez and dense cover of ephemeral forbs and grasses, 17 Dec 1966, Felger 15206 (ARIZ, MEXU, TEX); 5.1 mi by road NE of town of Bahia Kino, low saline flats with silty soil, near sea level, cactus scrub, 19 Dec 1966, Felger 15285 (ARIZ, MEXU, TEX, US); desemboque del Rio de la Concepcion, vicinity 30° 33’ N, 113° 00’ W, desert scrub in high shifting beach dunes, locally common, 27 Dec 1967, Felger 16781 (ARIZ, ENCB, GI, MEXAU, NY, TEX, UC) US): Isocoma felgert Nesom appears to be endemic to the area around Bahia Kino along the coast of Sonora. The plants are similar to [. acradenia in their oblong, whitish, indurated thickened phyllaries, each with a sharply delimited resin pocket at the apex, but plants of /. acradenia have neither such deeply 90 PHYTOLOGIA volume 70(2):69-114 February 1991 cut corolla lobes nor similarly dissected leaves. /socoma tenuzsecta has simi- lar corollas but a much denser vestiture and its phyllaries typically lack the strongly defined resin pockets. /socoma acradenia var. eremophila also has pinnately toothed or lobed leaves but the leaf segments are much broader, although a few collections with narrow, short lobes have been made from the north end of the Gulf of California. Var. acradenia occurs sporadically along the coastal region of Sonora to near Games and has been collected near Bahia Kino, apparently completely surrounding the range of J. felgerz. Plants of var. acradenia, however, produce entire, narrowly obovate leaves, although the heads are of nearly the same size as I. felgeri. The new species is named for Dr. Richard Felger, its sole collector and a relentless explorer of arid habitats in the southwestern United States and adjacent México. 7. Isocoma gypsophila B. ‘Turner Isocoma gypsophila B. Turner, Sida 5:24. 1972. TYPE: MEXICO. Nuevo Leén: 15 mi S of San Roberto Junction, B.L. Turner 6213 (HOLOTYPE: TEX!; Isotype: NY!). Similar to Jsocoma veneta and occurring sympatrically in the northern- most portion of its range, but different in its smaller stature, villosulous upper stems, solitary (or rarely double) and larger heads with 30-40 flowers (the in- volucres 6-7 mm long, 7-9 mm wide), and its broader inner and outer phyllaries with broad, translucent margins and green, barely glutinous, apical portions; receptacles with short, lacerate-lanceolate alveoli. Chromosome number, n = 6 pairs (Turner 1972). Local in Nuevo Leon and adjacent northern Zacatecas; saline flats in gy pse- ous soil, hills, with other gypsophilous subshrubs and mesquite, juniper, op- untia; 1600-1800 m; August-October. In the original description Turner noted that typical /socoma veneta oc- curred at the type locality intermixed with plants of J. gypsophila B. Turner. Plants of I. gypsophila from Zacatecas are large headed with villous vestiture, but the heads are sometimes paired rather than solitary. All collections exam- ined for this relatively poorly known species are cited here: MEXICO: Nuevo Leon: topotype, 6 Aug 1971, Reveal 2655 (GH, LL). Zacatecas: no other data, 1908, Lloyd 19 (GH, US); hills near Cedros, 30 Oct 1907, Lloyd & Kirkwood 148 (GH); E of San Juan de los Cedros, ca. 35 mi W of Mex Hwy 54 near Concepcion del Oro, 22 Sep 1973, Reveal 3361 (GH, NY, TEX, US). ; Nesom: Taxonomy of Isocoma 91 8. Isocoma hartwegii (A. Gray) E. Greene Isocoma hartwegi (A. Gray) E. Greene, Erythea 2:111. 1894. BASIONYM: Bigelovia hartwegit A. Gray, Synopt. Fl. N. Amer. 1(2):143. 1884. LECTOTYPE (selected by McVaugh 1984): MEXICO. Jalisco: Lagos, Hartweg 114 (K, NY-photo!; Isotype: GII according to Hall 1928, but not relocated in the present study). Aster hartwegu (A. Gray) O. Ktze., Rev. Gen. 318. 1891. (H)Aplopappus hartwegu (A. Gray) S.F. Blake, Contr. U.S. Natl. Herb. 23:1492. 1926. Haplopappus venetus (Kunth) S.F. Blake var. hartwegi (A. Gray) McVaugh, Contr. Univ. Michigan Herb. 9:364. 1972. Subshrubs, at least the upper stems, but commonly stems and leaves, minutely hispidulous to stiffly puberulous. Leaves oblanceolate, usually nar- rowly so, pinnatifid with 1-4 pairs of slender lobes, 5-20 mm long, the blades 0.5-1.5 (-2.0) mm wide, 2-8 mm wide from lobe tip to tip, 8-20 times longer than wide. Heads 13-22 flowered, broadly turbinate to campanulate, rounded at base, the involucres 5-7 mm high, 4-5 mm wide; phyllaries greenish, barely thickened at the apex, sometimes gland-dotted, without prominent scarious margins; receptacles strongly triangular-alveolate. Corollas 4.5-6.0 mm long, the tube 2.0-3.0 mm long, the lobes deltate to deltate-triangular, 0.6-1.0 mm long, 1/4-1/3 the length of the limb. Achenes broadly turbinate, 1.3-2.3 mm long, densely short sericeous, with 4-6 low, broad, resinous ribs. Chromosome number, n = 6 pairs (Jackson & Dimas 1981, as Haplopappus venetus |[Kunth| 5.F. Blake). Zacatecas, San Luis Potosi, Aguascalientes, Jalisco, Guanajuato, and Hi- dalgo; gravelly or sandy loam, low hills, saline alluvium, clay flats, or over limestone, gypsum, sandstone, or rhyolite, Larrea-Prosopis, matorral; 1800- 2400 m; (May-) June-November (-February). Isocoma hartwegi has been treated as only varietally distinct from /. veneta by McVaugh (1984). Although it is likely that the two probably are related as sister taxa, they may reasonably be considered as separate species, consistent in treatment (in the present study) with other closely related taxa in the genus. /socoma hartwegu has shorter, pinnatifid (vs. merely toothed) leaves and longer corolla lobes than J. veneta. The two taxa are parapatric (Map 4), and in east central Zacatecas and southwestern San Luis Potosi, where both occur, plants that might be regarded as intermediate are rare among the numerous specimens representing the two extremely well collected species. The single collection known from Hidalgo (Tula, Viereck 1279-US) is typical in morphology. A collection of /. veneta from the same area, however (Tula, Pringle 6405-US 2 sheets), comprises branches typical of the latter as well as one branch apparently intermediate between /. veneta and J. hartwegw. Both branches on Pringle 6405 from ND-G are intermediate, and two branches on a MO sheet are typical /. veneta. 92 PHY TOLOGIA volume 70(2):69-114 February 1991 Within his concept of Jsocoma hartwegzi, Ifall (1928) included J. coronopi- folia as well as J. tenuisecta. The latter has pinnatifid leaves and consistently hispidulous vestiture like /. hartwegi, but it has smaller heads with fewer flow- ers, phyllaries with scarious margins and thicker apices, longer achenes, and it is distantly separated in geography. 9. Isocoma humilis Nesom. Isocoma humilis Nesom, sp. nov. TYPE: UNITED STATES. Utah: Wash- ington Co., Zion National Park, sandstone, 25 Sep 1971, W.R. Leverich 1045A (HOLOTYPE: TEX!). Isocomae rusbyo KE. Greene similis capitulis latis multifloribus et phyllariis ovatis marginibus scariosis apicibusque tantum punctati- glandulosis sed habitu humili acervato, foliis integris, vestimento villoso, et acheniis minoribus costis vadosioribus minus resinosis differt. Shrubs apparently with a low, moundlike habit, with short, densely and highly branched stems mostly 4-8 cm high; young stems and leaves moderately villous with short, crisped, white hairs. Leaves gland dotted but not resinous, narrowly oblanceolate, 5-10 (-18) mm long, 1.5-3.0 (-5.0) mm wide, sometimes entire but usually with 1-2 (-3) pairs of pinnately arranged teeth or shallow lobes, the teeth and leaf apices with a thick, short, white, spinulose claw. Heads 19-28 flowered, solitary or in pairs, on bracteate pedicels 3-10 mm long, the involucre campanulate, basally rounded, 6-7 mm wide, 5-6 mm high; phyl- laries narrowly ovate, in 3-4 strongly graduated series, white indurated with narrow but prominently scarious margins, with a sharply delimited, gland dot- ted apical portion, without resin pockets. Corollas 4.0-5.0 mm long, the tube 2.3-2.8 mm long, the lobes triangular, 0.5-0.8 mm long. Achenes 1.5-2.0 mm long, obovate, sericeous, with 6-8 slightly raised ribs; longest pappus bristles 3-4 mm long. Chromosome number unknown. Additional collections examined: UNITED STATES. Utah: Washington Co., Dixie State Park, Snow Canyon, near St. George, red sandstone canyon, siliceous sandy soil, 25 Sep 1971, Leverich 1042A (TEX); Washington Co., 15 mi W of Zion National Park on Utah 15, sandstone, sand, 25 Sep 1971, Leverich 1044A (TEX). Isocoma humilis Nesom is recognized by its low stature and rounded habit, villous vestiture, small, toothed leaves, and small corollas and achenes. It occupies a geographic position between that of J. rusbyi and J. acradenia (Map 2), but its relationships are difficult to perceive. It is similar to J. acradenia var. eremophila in its toothed leaves and its indurated phyllaries with sharply delimited apical portions but more similar to J. rusbyi in its broad, many Nesom: Taxonomy of Isocoma 93 flowered heads and ovate phyllaries with scarious margins and merely punctate glandular apical portions. It differs from both species in its habit, vestiture, small leaves, and small achenes, although /. acradenia produces achenes that rarely range as small. All three collections of Isocoma humilis were made by William Leverich, at the time a graduate student in botany at the University of Texas who had begun an investigation of the systematics of Jsocoma. He did not complete his studies but his eye for plants of the genus produced interesting collections and observations. The occurrence of /socoma in Washington Co. was not recorded by Welsh (1983). 10. [socoma menziesu (Hook. & Arn.) Nesom. Isocoma menziesii (Hook. & Arn.) Nesom. BASIONYM: Pyrrocoma men- zvestt Hook. & Arn., Bot. Beechey Voy. 351. 1838. Variable in habit, vestiture, and leaf morphology. Heads turbinate, basally obtuse to acute, with (15-) 18-25 (-28) flowers, the involucres (6-) 7-9 mm high, 6-8 mm wide; phyllaries oblong oblanceolate to oblong lanceolate, usually the lower 2/3 strongly white indurated, lanceolate acute to rounded at the apex, mostly with green punctate apical areas, small resin pockets sometimes developed in var. menziesii and var. decumbens (E. Greene) Nesom; receptacles with long, narrowly to broadly lanceolate alveoli. Corollas 5.0-6.5 (-7.0 in var. diabolica Nesom) mm long, the tube 3-4 mm long, the lobes 0.5-0.8 mm long, deltate, 1/5-1/3 the length of the limb. Achenes 2.3-3.6 (-4.0) (-5.0 in var. diabolica) mm long, with 5-6 orange resinous ribs, or sometimes with up to 10-11 thinner, apparently nonresinous nerves, sericeous, teretish to somewhat flattened. Isocoma menziesii comprises a series of morphologically disparate but inter- grading taxa, which have been treated as varieties or subspecies of J. veneta in many previous studies. The chief dissenter was the chief architect of the genus, E.L. Greene himself, who regarded each of these as a separate species. Indeed, each variety is as singular in its typical morphology as other taxa of [socoma recognized as distinct species in the present treatment. Numerous intergrades (as noted below), however, have deterred most students of the group, including the present author, from maintaining more than one species, but field studies and more detailed analyses of morphology may yet show some of these taxa to be isolated to a greater degree than hitherto suspected. One such taxon, /. arguta, is segregated as a distinct species in the present treatment. All of the taxa included here in Jsocoma menziesii, however, are distinct from typical J. veneta and none of them intergrade with it. Compared to the plants of central México, all those of the Pacific coast (J. menzzesiz) differ in their vestiture, longer involucres, and their longer achenes. As in many of the 94 PHY TOLO'G TA volume 70(2):69-114 February 1991 species of Isocoma, these two have parallel tendencies of variation, but they are no more alike than other presumably closely related but distinct species. Further, J. veneta and J. menziesii are strongly different in habitat and disjunct more than 1100 kilometers at point of closest approach to each other. Key to the varieties of /. menziesi 1. Leaves obovate, coarsely serrate, fleshy thick; stems and leaves glabrous; plants decumbent. 0:00.20. 25°. 3... 22 eee eee var. sedozdes 1’ Leaves obovate to oblanceolate, entire to toothed, not fleshy thick, or if so then at least the stems prominently villous; stems and leaves glabrous to villous, tomentose, or glandular; plants erect to decumbent. ..... (2) 2. Herbage stipitate glandular, essentially without other vestiture. (3) 2' Herbage glabrous to villous or tomentose, not prominently glandular. [vines cndilabtcas$iheves (4) 3. Plants strictly erect; leaves 4-10 mm wide; capitulescence distinctly corym- boid; achenes mostly 4-5 mm long; San Benito and Santa Clara cos. Ske na Re Re Soca hs Rac te Ae nc Lae Ln ote .... var. diabolica 3’ Plants decumbent to erect; leaves 2-4 mm wide; capitulescence corymboid to loosely paniculate; achenes mostly 2.3-3.6 mm long; San Diego Co. to Baja California; Norte:: s...2.8. 5. foe...) 3 Sa eee var. decumbens 4, Plants glabrous or slightly hairy, sometimes resinous. .......... (5) 4’ Plants prominently tomentose or villous. ..................++-- (8) 5. At least the lower leaves with serrate margins. ..............+...+0: (6) 5’ Leaves entire or few toothed at the apex. .......1: 2.0... sence (7) 6. Corollas 6-7 mm long; achenes (3.8-) 4.5-5.0 mm long. sate wb Vols d See Rb ofek atts s dan, Gee var. diabolica 6’ Corollas 5-7 mm long; achenes 2.3-3.6 mm long. . var. vernonzoides 7. Leaves distinctly spreading toothed or lobed at the apex; phyllary apices usually spreading to slightly reflexed. .................. var. tridentata 7’ Leaves entire to shallowly serrate at the apex; phyllary apices erect, rarely SPLCAGING.. de ois v cabrones sm Syeremete's sine eae doce var. menzvesit 8. Herbage villous with spreading-crisped, relatively thick based hairs; leaves oblanceolate to oblong oblanceolate, commonly toothed along most of the margins; stems usually erect. ....... var. vernonioides Nesom: Taxonomy of /socoma 95 8’ Herbage finely and closely gray tomentose; leaves narrowly oblance- olate, entire to few toothed; stems decumbent to erect. AD They BLS cx thd STP eS ET? Sy phe eed oat ad ce per etave var. decumbens a. Var. decumbens (EF. Greene) Nesom. Isocoma menziesii (Hook. & Arn.) Nesom var. decumbens (E. Greene) Nesom, comb. nov. BASIONYM: Jsocoma decumbens E. Greene, Leafl. Bot. Observ. Crit. 1:172. 1906. LECTOTYPE (designated here): UNITED STATES. California: San Diego Co., clay depression on mesas, near San Diego, 13 Sep 1903, 7.5. Brandegee 3405 (ND-G!; Isolectotypes: GH!, LL!, NY!, US!). Haplopappus venetus (Kunth) S.F. Blake var. de- cumbens (E. Greene) Munz, Man. S. Calif. Bot. 522. 1935. The com- bination /socoma veneta (Kunth) E. Greene var. decumbens (E. Greene) Jepson (Man. Fl. Pl. S. Calif. 1029. 1925) was not legitimately made, as it was based on an apparently unpublished name, “Bigelowia veneta var. decumbens Bdg.” The lectotype has a handwritten label as “Jsocoma decumbens Greene;” the duplicates have a printed label distributed by C.F. Baker as “Jsocoma decumbens (Brand.) Greene.” Bigelowia furfuracea E. Greene, Bull. Calif. Acad. Sci. 1:87. 1885. TYPE: Probably MEXICO. Baja California Norte (according to Greene): collection data not specified (UC, see Hall 1928 for com- ments). Haplopappus venetus (Kunth) S.F. Blake subsp. furfuraceus (E. Greene) Hall, Carnegie Inst. Washington Publ. 389:226. 1928. Haplopappus venetus (Kunth) S.F. Blake var. furfuraceus (E. Greene) Munz, Man. S. Calif. Bot. 523. 1935. Isocoma veneta (Kunth) E. Greene var. furfuracea (E. Greene) Beauchamp, Phytologia 59:437. 1986. Nearly prostrate to decumbent or somewhat erect shrubs, closely arachnoid and minutely glandular, sometimes one or the other. Leaves narrowly oblance- olate, mostly entire or with 1-2 pairs of teeth near the apex, 5-22 mm long, 2-4 mm wide, commonly densely arranged, often in axillary fascicles. Heads turbinate-campanulate, usually arranged in a loose capitulescence with numer- ous (1-) few headed clusters at the ends of slender, often lax branches; phyl- laries narrowly oblanceolate-oblong, strongly white indurated, usually with an orange midvein from top to bottom, acute to rounded at the apex, the middle and inner sometimes short aristate, with a sharply delimited, villosulous apical area, punctate or often with several small but distinct resin pockets. Chromo- some number, n = 12 pairs (Raven, ef al. 1960, as Haplopappus venetus var. vernonioides). Localized in southern San Diego Co., San Clemente and Santa Catalina Islands, and immediately adjacent Baja California Norte; sandy flats or slopes, commonly in disturbed sites, 10-50 (-175) m; July- November. 96 PHYTOLOGIA volume 70(2):69-114 February 1991 Plants of var. decumbens may be erect to decumbent in habit, commonly with slender stems and small, sometimes crowded leaves, but they are partic- ularly distinguished by their vestiture. The stems, leaves, and phyllaries most commonly produce a close, arachnoid tomentum of hairs finer than those in var. vernonioides, and beneath this, a layer of minute, stipitate glands. Tomen- tose plants are the more common, but the development of both the tomentum and the glands is variable and some plants may produce only one or the other. The type of Bigelovia furfuracea E. Greene apparently produces only minute glands (Hall 1928), lacking other types of trichomes. The narrow, strongly in- durated phyllaries, sometimes with resin pockets (also found in vars. menziesi2 and tridentata |E. Greene] Nesom), are similar to those of J. acradenia, and the possibility of genetic influx from the latter should be investigated further. The boundaries between var. decumbens and both var. menziesi and var. vernonioides are blurred by intermediates, although there does appear to be some degree of isolation. Collections of var. decumbens and var. menziesiz, including apparent intermediates, have been in San Diego Co. at Lindo Lake ( Youngberg 29 and 29a-LL) and at Chula Vista (various collectors). Many of these prominently tomentose plants appear to be otherwise identical in mor- phology to var. menziesii. In most cases, | have arbitrarily identified plants as var. menziesii if they are completely glabrous, even though they are otherwise similar to var. decumbens. b. Var. diabolica Nesom. Isocoma menziesii (Hook. & Arn.) Nesom var. diabolica Nesom, var. nov. TYPE: UNITED STATES. California: San Benito Co., Diablo Range, 9 mi SE of turnoff to Pinnacles Natl. Monument along Calif. Rte. 25, openly wooded slope with Quercus turbinella, Juniperus californica, and assorted shrubs, 1250 ft, locally abundant, many branched shrub, 5 Oct 1985, D.J. Keil 19042 (HOLOTYPE: TEX!; Isotypes: OBI, UCR!). Isocomae menziesii (Hook. & Arn.) Nesom var. vernonioidi (Nutt.) Nesom similis sed vestimento dense stipitati-glanduloso plerumque sine villis, corollis ac acheniis longioribus, et habition- ibus montanis differt. Erect shrubs 4-6 dm tall, with distinctly whitish stems, the stems, leaves, and phyllaries densely viscid with stipitate glandular hairs, sometimes eglan- dular but then densely resinous viscid, without other pubescence except some- times very sparsely villous in the leaf axils or along stems. Leaves obovate to narrowly oblanceolate or narrowly elliptic-oblanceolate, the largest lower 2-4 cm long, 5-10 mm wide, with the margins shallowly serrate, the upper sharply reduced in size, entire. Heads 20-26 flowered, in corymboid capitulescences, turbinate, the involucres 7-9 mm high; phyllaries with an apical area 1/2-1/3 Nesom: Taxonomy of Isocoma 97 the total length, gradually developed from the base, punctate, the tip slightly reflexed or crisped, often short-spinulose. Corollas 6.0-7.0 mm long. Achenes (3.8-) 4.3-5.0 mm long, narrowly oblong-oblanceolate, somewhat compressed, densely sericeous. San Benito and Santa Clara cos., apparently most abundant in the Diablo Range of the former; open slopes and cliffs, mostly in foothill woodlands, 15- 400 m; August-October. Additional collections examined: UNITED STATES. California: San Ben- ito Co., Tres Pinos—Paicines, 22 Sep 1920, Abrams 7661 (NY); San Benito Co., near Emmett’s Station, Panoche Pass Road, exposed cliffs, 25 Sep 1927, Ferris 6889 (LL); San Benito Co., along Rte. 25 SE of Hollister, 2.5 mi NW of Paicines, foothill woodland area, 600 ft, 4 Sep 1983, Keil 17924 (UCR). Santa Clara Co., Calaveras Road 2 mi E of Milpitas, 50 ft, open hills, 10 Oct 1955, Rose 55183 (TEX). These plants were included by Hall (1928) for the most part within his concept of /socoma veneta var. vernonioides, but they are distinctive in mor- phology as well as geography. They are the only ones of J. menzzesz to inhabit primarily inland, montane sites. Var. diabolica differs from var. vernonioides in its resinous, usually densely stipitate glandular vestiture (with villous hairs lacking or sparsely present along the stems), and in its consistently reduced, somewhat bractlike upper cauline leaves, longer corollas, and longer achenes. In vestiture and leaf morphology, as well as the inland habitats, these plants are reminiscent of /. acradenia var. bracteosa, which occurs only slightly to the south and east, but the phyllaries of var. diabolica lack well developed resin pockets. Although the plants of var. dzabolica are consistent in morphology, some of var. vernonioides, including the type of /. villosa E. Greene, also are stipitate glandular. Further, some plants from San Diego Co. (e.g., Brandegee 1633-NY, US-2 sheets), as well as a number of collections from around King City in Monterrey County (e.g., Rose 36735-GII, US), may be very similar to var. diabolica in habit and vestiture, although in achene and corolla size they are more typical of other varieties of /. menzzesw (identified here as var. decumbens and var. vernontoides, respectively). Populations of Jsocoma acradenia var. bracteosa occur in close proximity to those of var. dzabolica (e.g., San Benito Co., Griswold Hills, ca. 3 mi S of jet of Idria road with Panoche Pass road, Ferris & Ernst 13083-NY). c. Var. menztesii Isocoma menziesii (Hook. & Arn.) Nesom var. menzzesiz BASLONYM: Pyrro- coma menztesu Hook. & Arn., Bot. Beechey Voy. 351. 1838. TYPE: UNITED STATES. California: [near San Diego?|, 1786-89, A. Menzies (K, GH-photo!, US-photo!). (H)Aplopappus menziesi (Hook. & Arn.) Torr. & A. Gray, Fl. N. Amer. 2:242. 1842. Bigelovia menziesii (Hook. 98 PHYTOLOGIA volume 70(2):69-114 February 1991 & Arn.) A. Gray, Proc. Amer. Acad. Arts 8:638. 1873. See comments by Hall (1928) regarding the type specimens of this taxon and /. vernon- 1oides. Isocoma ozyphylla E. Greene, Leafl. Bot. Observ. Crit. 1:171. 1906. LECTOTYPE (designated here): UNITED STATES. California: San Diego Co., Jamul Valley, near San Diego, 1875, E. Palmer 134 (US!; Isolectotypes: BM, UC). Haplopappus venetus (Kunth) S.F. Blake subsp. oryphyllus (E. Greene) Hall, Carnegie Inst. Wash- ington Publ. 389:225. 1928. Haplopappus venetus (Kunth) S.F. Blake var. oryphyllus (E. Greene) Munz, Man. S. Calif. Bot. 523. 1935. Isocoma veneta (Kunth) E. Greene var. oryphylla (E. Greene) Beauchamp, Phytologia 59:438. 1986. Haplopappus fasciculatus Vasey & Rose, Proc. U.S. Natl. Mus. 11:530. 1888. TYPE: MEXICO. Baja California Norte: San Quentin Bay, Jan 1889, E. Palmer 635 (HOLOTYPE: US!). Shrubs 0.5-2.5 m tall, glabrous and usually resinous, erect or in spreading clumps, rarely decumbent. Leaves narrowly oblanceolate, 15-30 (-40) mm long, 3-8 mm wide, entire or with 1-2 pairs of apical teeth or shallow lobes, axillary fascicles commonly produced. Phyllaries sometimes reflexed or spreading at the apex, especially in San Diego Co., commonly with a thin, orange midvein extending from base to tip, small resin pockets sometimes present. Chromo- some number, n = 12 pairs (Pinkava & Keil 1977, as H. venetus [Kunth] S.F. Blake subsp. furfuraceus |E. Greene] Hall). Baja California Norte and Baja California Sur, Orange and San Diego cos., San Clemente and Santa Catalina Islands, largely replaced northward by var. vernonioides, except for apparently disjunct populations in the vicinity of San Francisco (see below); chaparral, scrub communities, dunes or sandy flats, sandy arroyos, stream banks, edges of saline ponds, rocky canyon walls; 2-350 (-1100) m; (April-) August-January. Plants of var. menziesii are similar to those of var. decumbens but different in their glabrous herbage, larger leaves, and heads commonly in more compact and distinctly corymboid capitulescences. The type of [socoma menziesii, with its glabrate herbage and narrowly lanceolate leaves with a few shallow teeth near the apex, is more similar to plants previously known as J. ozyphylla E. Greene than any other taxon. In fact, similar plants occur through nearly the whole range of var. “ozyphylla.” To the north of San Diego and Orange counties, plants (var. vernonioides) have a tendency to produce a villous vestiture and obovate leaves evenly toothed along most of the margin. The two taxa, however, appear to be completely in- tergrading, and many plants are necessarily identified arbitrarily. In San Diego County, nearly all the plants identified here as var. menziesii are morpholog- Nesom: Taxonomy of Isocoma 99 ically influenced to some degree either by var. decumbens or by var. vernon- ioides. In the present study, I have identified glabrous or glabrate plants with leaves prominently toothed along the margins as well as plants prominently villous but with more weakly toothed leaves as var. vernonioides. Glabrous or glabrate plants with entire to apically few toothed leaves are var. menziesii. Clearly, these two taxa are essentially the same biologically as those perceived by Hall, but I have associated the type of J. menzzesi with his “subsp. ozy- phylla” rather than J. vernonzozdes. From Encinitas to La Jolla (San Diego Co.), a number of collections have been made of decumbent plants otherwise mostly similar to erect plants of var. menziesi. In Baja California N and S, spreading to somewhat decumbent plants are relatively common, and the significant variability in leaf shape in that area needs to be investigated further. Haplopappus fasciculatus is a form with densely arranged leaves but otherwise within the geographical and mor- phological range of var. menziesiz, which commonly produces axillary fascicles of small leaves. The plants identified as var. menziesi from the vicinity of San Francisco (e.g., Brandegee s.n.-NMC, NY; Cannon s.n.-GH) produce sparsely villous stems but nearly glabrous leaves with a few, shallow teeth only on the distal margins, and they are nearly identical to many collected from around San Diego. The San Francisco populations are far disjunct from others of var. menziesii and they appear to be separated from those of var. vernonzozdes as well; at least I have seen no collections of the latter from Santa Cruz Co., and those from Monterrey Co. are strongly divergent in morphology. d. Var. sedoides (E. Greene) Nesom. Isocoma menziesii (Hook. & Arn.) Nesom var. sedoides (E. Greene) Ne- som, comb. nov. BASIONYM: Bigelowia veneta var. sedozdes E. Greene, Bull. Calif. Acad. Sci. 2:400. 1887. TYPE: UNITED STATES. Cal- ifornia: [Santa Barbara Co.], Santa Cruz Island, “edges of low cliffs overhanging the sea, on the north side of the island,” other data not specified (CAS). Isocoma sedoides (E. Greene) E. Greene, Leafl. Bot. Observ. Crit. 1:172. 1906. Jsocoma veneta (Kunth) E. Greene var. se- doides (E. Greene) Jepson, Man. Fl. Pl. Calif. 1029. 1925. Haplopappus venetus (Kunth) S.F. Blake var. sedoides (E. Greene) Munz, Man. S. Calif. Bot. 522. 1935. See comments following the citation of J. latzfolza. Plants glabrous or nearly so; stems stout, decumbent, prostrate, or pen- dent. Leaves clearly succulent, broadly obovate to oblanceolate, coarsely ser- rate. Heads in a dense, capitate cluster. California (Santa Rosa, Santa Cruz, San Miguel, Anacapa, and Santa Catalina Islands, and adjacent coast from the vicinity.of Newport Beach north 100 PHYTQLOGHA volume 70(2):69-114 February 199] to Morro and Cambria), northwestern Baja California Norte; dunes, coastal scrub, talus slopes, commonly pendulous on cliff edges, 0-20 m; (April-) June- December. Var. sedoides (E. Greene) Nesom appears to be somewhat distinct in habi- tat though sympatric in distribution with var. vernonizoides, and intermediates between the two have been collected at every locality where var. sedozdes oc- curs. Some identifications have to be made arbitrarily. The label of a collection from Piedras Blancas Point in San Luis Obispo Co. (Leverich 1072-TEX) notes that a mixed population occurred there: upright plants (var. vernoniozdes), with glabrous to pubescent herbage, and a “low form in tangled woody clumps up to 6 ft in diameter,” with succulent leaves (var. sedoides). Plants of the single collection from México here identified as var. sedozdes (Baja California Norte, ca. 1 mi N of mouth of Arroyo Santo Tomas on coastal bluffs, Wiggins & Thomas 406-US) were noted as being “shrubby.” e. Var. tridentata (KE. Greene) Nesom. Isocoma menziesii (Hook. & Arn.) Nesom var. tridentata (E. Greene) Nesom, comb. nov. BASIONYM: Bigelovia tridentata E. Greene, Bull. Torrey Bot. Club 10:126. 1883. TYPE: MEXICO. Baja California Norte: Cedros Island, 1885, E. Greene s.n. (HOLOTYPE: UC). Jsocoma tridentata (E. Greene) E. Greene, Erythea 2:111. 1894. (H)Aplopappus tridentatus (E. Greene) S.F. Blake, Contr. U.S. Natl. Herb. 23:1493. 1926. Haplopappus venetus (Kunth) S.F. Blake subsp. tridentatus (E. Greene) Hall, Carnegie Inst. Washington Publ. 389:225. 1928. Linosyris dentata Kellogg, Proc. Calif. Acad. Sci. 2:16. 1863. TYPE: MEXICO. Baja California Norte: Cedros Island, Veatch (UC-see comments by Hall 1928). Distinguished by its narrowly oblanceolate leaves 15-30 mm long, 2-5 mm wide, consistently with 1-2 (-3) pairs of prominently divergent teeth or lobes near the apex and its phyllaries with triangular-lanceolate-attenuate, reflexing or spreading apices. Erect shrubs 0.8-1.5 m tall, completely glabrous, usually glutinous; phyllaries often with small resin pockets; achenes 2.3-2.6 mm long. Cedros Island of Baja California Norte and immediately adjacent coastal mainland, from near Punta Prieta (Baja California Norte) to the Vizcaino region (Baja California Sur); coast or upper strand, dunes, arroyos, pond edges, rocky areas, 1-125 (-200) m; October-May. Var. tridentata is similar to var. menziesii and apparently intergrades with it on the mainland, particularly in the region of Punta Prieta, but numerous and relatively uniform collections of these distinctive plants have been made within the small area of its range, both on Cedros Island and the mainland. The spreading-reflexing phyllary apices are distinctive although similar ones Nesom: Taxonomy of Isocoma 101 sometimes occur in plants of var. menziesii, particularly those from San Diego County and vicinity. f. Var. vernonioides (Nutt.) Nesom. Isocoma menziesii (Hook. & Arn.) Nesom var. vernonioides (Nutt.) Ne- som, comb. nov. BASIONYM: Isocoma vernonioides Nutt., Trans. Amer. Phil. Soc., ser. 2, 7:320. 1840. HOLOTYPE: UNITED STATES. Cal- ifornia: Santa Barbara, in marshes near the sea, Apr-May [1836], T. Nuttall s.n. (BM, GH-photo!; Probable isotypes: GH!, NY!). /socoma veneta (Kunth) E. Greene var. vernoniordes (Nutt.) Jepson, Fl. W. Mid. Calif. 560. 1901. Haplopappus venetus (Kunth) S.F. Blake subsp. ver- nonioides (Nutt.) Hall, Carnegie Inst. Washington Publ. 389:224. 1928. The branch on the NY sheet closely matches the BM plant and descrip- tion; that on the GH sheet is significantly different. Hall (1928) also cited a duplicate at K. Isocoma leucanthemifolia E. Greene, Leafl. Bot. Observ. Crit. 1:171. 1906. LECTOTYPE (designated here): UNITED STATES. Cali- fornia: San Diego Co., Warner’s Ranch, 21 Oct 1889, C.R. Orcutt (US!). Isocoma microdonta E. Greene, Leafl. Bot. Observ. Crit. 1:171. 1906. LECTOTYPE (designated here): UNITED STATES. California: [Santa Barbara Co.], Santa Maria, Nov 1893, Mrs. Blochman s.n. (ND-G!). Isocoma latifolia E. Greene, Leafl. Bot. Observ. Crit. 1:172. 1906. LECTOTYPE (designated here): UNITED STATES. California: [Santa Barbara Co.], Santa Cruz Island, Jul-Aug 1886, l.L. Greene s.n. (ND-G!; Isolectotype: GH!). On the ND-G sheet (marked by Greene as “/Jsocoma latifolia, Greene Type!”) are four branches, two of which are prominently villous, matching Greene’s descrip- tion of this taxon. The other two branches are nearly glabrous and are referable to var. sedoides in the present study. The GH speci- men is var. sedoides. The glabrous branches in this collection may ultimately be found to represent elements of the type of Greene’s Isocoma sedoides. Isocoma villosa E. Greene, Leafl. Bot. Observ. Crit. 1:172. 1906. LEC- TOTYPE (designated here): UNITED STATES. According to Greene, “sent from southern California” and “grown in the Uni- versity Botanic Garden” at Berkeley, where gathered in Nov 1893 by Mr. Davy (ND-G!; Isolectotype: ND-G!). The label data on the two type sheets correspond exactly with Greene’s published data. Three branches are preserved and probably are from the same plant; 102 PHYTOLOGIA volume 70(2):69-114 February 1991 the stems are villous-pilose and the leaves are densely stipitate glan- dular and sparsely pilose. Erect or ascending to somewhat decumbent subshrubs 0.5-1.0 m tall, glab- rate to hirtellous- or villous-pilose or densely gray tomentose with long, vitre- ous, flattened hairs, sometimes stipitate glandular as well. Leaves glabrate to villous, linear to oblanceolate or spatulate-oblong, 1-4 cm long, 2-6 (- 9) mm wide, pinnately toothed or shallowly lobed. Phyllaries acute atten- uate to deltate or nearly oblong with a blunt apex, not aristate or rarely only slightly so, usually glabrous. Achenes lightly sericeous, usually with 5- 6 orange-resinous nerves but sometimes with 10-11 thin, light colored nerves. Chromosome number, n = 12 pairs (Raven, et al. 1960; De Jong & Montgomery 1963; Pinkava & Keil 1977; Keil 1979; Semple, et al. 1989, as J. veneta). Northern Baja California Norte, abundant in California primarily along the coast from San Diego north to Santa Barbara and on all the Channel Islands, more infrequent northward; coastal bluffs and dunes, sandy flats, borders of salt marshes, and occasionally on dry slopes, 5-400 m in California, 650-900 m in Mexico; (April-) July-December. In the southern area of their range (particularly San Diego and Riverside Cos.), plants of var. vernonioides produce small leaves that are often nearly glabrous, perhaps reflecting the influence of genes from var. menziesu. Among putative intermediates with var. menziesi, plants with regularly serrate leaf margins have been arbitrarily identified here as var. vernonioides. Several similar but atypical collections from the area of Pasadena (e.g., Grinnell s.n..NY, US; McClatchie s.n-NY) are apparently decumbent with loose, long pedicellate capitulescences and have phyllaries with large, sharply delimited apical areas. See var. sedoides for other comments on variation in var. vernoniozdes; further detailed comments regarding variation in the latter were provided by Hall (1928). plan Flot, ! 11. [socoma plurifoha (Torr. & A. Gray) E. Greene Isocoma plurifolia (Torr. & A. Gray) E. Greene, Erythea 2:111. 1894. BA- SIONYM: Linosyris plurifolia Torr. & A. Gray, I'l. N. Amer. 2:233. 1842. TYPE: UNITED STATES. [Sources of the Canadian”], from the Long Expedition, 1820, Dr. E. James (HOLOTYPE: NY!, see com- ments below regarding the collection locality and type). Bigelovia plu- rifolia (Torr. & A. Gray) A. Gray, Proc. Amer. Acad. Arts 8:638. 1873. Haplopappus plurifolius (Torr. & A. Gray) Hall, Carnegie Inst. Washing- ton Publ. 389:237. 1928. Nesom: Taxonomy of Isocoma 103 Linosyris wrightu A. Gray, Pl. Wright. 1:95. 1852. TYPE: UNITED STATES. Texas: [El Paso Co.], valley of the Rio Grande, 60 or 70 mi below El Paso, Sep 1852, C. Wright 284 (HOLOTYPE: GIi!, NY-photo!; Isotypes: GH!, US!). Bzgelovia wrightii (A. Gray) A. Gray, Proc. Amer. Acad. Arts 8:639. 1873. Isocoma wrighti (A. Gray) Rydb., Bull. Torrey Bot. Club 33:152. 1906. Linosyris heterophylla A. Gray, Pl. Wright. 1:95. 1852. TYPE: UNI- TED STATES. Texas: [Reeves Co.?], valley of the Pecos, Aug 1849, C. Wright 283 (HOLOTYPE: GH!; Isotype: ND-G!). Aster hetero- phyllus (A. Gray) O. Ktze., Rev. Gen. 318. 1891. Isocoma hetero- phylla (A. Gray) E. Greene, Erythea 2:111. 1894. (H)Aplopappus heterophyllus (A. Gray) S.F. Blake in Tidestrom, Contr. U.S. Natl. Herb. 25:546. 1925. Linosyris hirtella A. Gray, Pl. Wright. 1:95. 1852. TYPE: UNITED STATES. Texas: [Jeff Davis Co.], “valley of the Limpia,” Aug 1849, C. Wright 285 (HOLOTYPE: GH!; Isotype: GH!). Bigeloua wrighta (A. Gray) A. Gray var. hirtella (A. Gray) A. Gray, Synopt. Fl. N. Amer. 1(2):142. 1884. Isocoma hirtella (A. Gray) Heller, Muhlenbergia 1:6. 1900. Isocoma ozylepis Woot. & Standl., Contr. U.S. Natl. Herb. 16:180. 1913. TYPE: MEXICO. Chihuahua: Mexican boundary line near White Water, 11 Sep 1893, E.A. Mearns 2288 (HOLOTYPE: US!). Isocoma halophytica B. Turner, Sida 5:23. 1972. TYPE: MEXICO. Chihuahua: S end of Laguna Jaco, 9 Sep 1940, /.M. Johnston & C.H. Muller 1090 (HOLOTYPE: MICH; Isotypes: GH!, TEX!). Shrubs 0.5-1.0 (-1.5) m tall, with whitish stems. Leaves spreading-ascend- ing, often curving upwards, oblanceolate to narrowly oblong oblanceolate or nearly linear, mostly 1-4 (-5) cm long, 3.0-5.0 (-9.0) mm wide, the margins en- tire or uncommonly with | (-3) pairs of shallow teeth or lobes, often sparsely and minutely ciliate along the margins, rarely more densely hirtellous, gray green, usually not glutinous, the blade with imbedded glandular papillae or less commonly short stipitate glandular, otherwise glabrous to sparsely hispidu- lous. Heads in dense, corymboid capitulescences, (8-) 11-17 (-21) flowered, campanulate to broadly turbinate, the involucres 3.2-5.5 mm high, 2.5-4.0 mm wide; phyllary apex barely differentiated, or if green and thicker, then surrounded by a scarious-translucent margin, not glutinous; receptacles with narrowly triangular to deltate alveoli. Corollas 5-6 mm long, the tube 3.0- 4.0 mm, glabrous, the lobes 1.0-1.8 mm long, 2/5-3/5 the length of the limb. Achenes 1.5-1.9 (-2.8) mm long, with 6-8 thick, resinous ribs. Chromosome number: n = 6 pairs (Powell & Turner 1963, as J. heterophylla, Reeves Co.; Weedin & Powell 1978, as H. wright, Culberson Co., Tex.; Powell & Powell 104 PHYTOLOGIA volume 70(2):69-114 February 1991 1977, as H. wright, Culberson and Reeves cos., Tex.); n = 12 pairs (Jack- son 1959, as H. plurifolia, Rio Arriba Co., N.M.; Turner, Powell, & Watson 1973, as I. wrightii, Brewster Co., Tex.; Powell & Powell 1977, as H. wraghtu, Hudspeth Co., Tex.). Chihuahua, Coahuila, Texas, New Mexico, Arizona; igneous or calcareous substrate, less commonly over gypsum, dunes, sandy or clay loam, commonly with Larrea- Prosopis; 400-1400 (-1600) m; (April-) July-October. The type specimen was first identified by Torrey as Chrysocoma graveolens Nutt. (Ann. Lyceum Nat. Hist. New York 2:211. 1828.), from a series of col- lections sent to him by the naturalist Dr. E.P. James of the Long Expedition. The authorship of this name was later incorrectly attributed to Torrey him- self, but in 1842, Torrey & Gray recognized that the plant was not the same species as Nuttall’s (a species of Chrysothamnus) and named it as a member of Linosyris. Torrey & Gray, as well as Gray in later publications, placed Chrysocoma graveolens (sensu Torrey) as a synonym of Linosyris plurifolia Torr. & A. Gray. The type collection of Linosyris plurifolia evidently was made at the north- ern edge of the range of the species; the label gives no specific information but Torrey & Gray noted that it might have come from the “Upper Missouri or Platte?.” Part of Long’s expedition, including Long & James, left Colorado in late July and August of 1820 and traveled southeastward into what they believed to be “sources of the Canadian” River. Osterhout (1920), however, thought it more likely that these were actually tributaries of the Cimarron in southeastern Colorado and adjacent New Mexico. In any case, the expedition continued to travel eastward and the type probably was collected either in the Texas panhandle or the northeastern corner of New Mexico at the upper extremity of the range of the species. No records outside the Texas panhandle are shown in the Atlas of the Flora of the Great Plains (GPFA 1977), but it is recorded here for northeastern New Mexico in Harding Co. (A.H. Wright s.n._NMC), and it has been reported for Union Co., New Mexico (Martin & Hutchins 1981, as Haplopappus heterophyllus; added on Map 2). Isocoma plurtfolia is recognized by its relatively small, few flowered heads, deeply cut corolla lobes, and usually entire, mostly glabrous, nonglutinous but papillate or stipitate glandular leaves commonly with sparsely ciliate margins. The species is variable in vestiture but its geographic boundaries appear to be well defined and there is no reasonable way to formally recognize the variants. See further comments below and following the description of J. tomentosa. Plants of /socoma plurifolia with shallowly toothed or lobed leaves are relatively common in central New Mexico along the western margin of the range of the species, where it approaches J. azteca. In Sandoval Co., New Mexico, Arsene 16486 (LL) from the vicinity of San Ysidro is typical /. plu- rifolia, but Hartman 3387 (LL) from 21 mi NW of Bernalillo bears three separate branches, all small headed like J. plurifolia and producing deeply in- Nesom: Taxonomy of Isocoma 105 cised corolla lobes, but the lower leaves on two of the branches are shallow pinnately toothed. Similar plants have been collected from another locality in Sandoval Co. (Puerco River, Losure 197-ARIZ 2 sheets), in Rio Arriba Co. (Chamita, Eggleston 20470-GH, NY-2 sheets), Santa Fe Co. (e.g., 13 mi SW of Santa Fe, Plowman & Kilham AP139-GH), and Socorro Co. (near Ft. Craig, Rusby 2286-ND-G). It is possible that these plants show the genetic influence of I. azteca, but other plants with few toothed or shallowly lobed leaves, in- cluding the type of Linosyris wrightu A. Gray, occur sporadically throughout the range of I. plurifolia, in most cases apparently significantly removed from possible sources of genetic influence from any other species. The prominently lobed leaf plants are mostly along the western margin of the species and might ultimately be recognized as a weakly defined variety. Isocoma plurifolia was reported by Kearney & Peebles (1951, as Haplopap- pus heterophyllus) as widespread in Arizona; some of the Arizona plants pre- viously identified as this species are J. rusbyz, which differs in its larger heads with more flowers and longer achenes with much thicker, apically extended ribs, and some apparently are forms of J. acradenza with only weakly devel- oped resin pockets in the phyllaries. Few specimens have been recorded in the present study that document the occurrence of J. plurifolia in Arizona: Cochise Co., 9 mi SE of Cochise, 8 Aug 1936, Anderson 1266 (NMC); Gra- ham Co., Sulphur Springs Valley below Fort Grant, 4 Sep 1919, Eggleston 15929-GH; [Pima Co.], Empire Ranch, 20 Sep-4 Oct 1902, Griffiths & Thorn- ber 278 (NY); “southern Arizona,” Sep 1874, Rothrock 694-GH, NY); “alka- line plains,” 11 Sep 1884, Pringle s.n.-GH, NY-2 sheets. These plants have deeply cut corolla lobes and other relatively features of J. plurifolia, except for the densely hispidulous leaves and the stems, which vary from hispidulous to sparsely short-puberulous or -villous. A plant from southwestern New Mexico (Luna Co., Columbus, Hershey s.n._NMC) also belongs with these. The An- derson collection, however, is glabrous, and except for it, these plants appear to form a natural unit; all are best associated with J. plurzfolia and may even- tually warrant formal taxonomic recognition, based primarily on their stem vestiture. The type of Linosyris hirtella A. Gray has densely hirtellous stems and leaves, and similar scattered collections, though mostly with less dense vesti- ture, have been made in southwest Texas and southern New Mexico, as well as in Arizona. It is possible that this reflects an ancestral relationship be- tween Isocoma plurifolia and I. tenuisecta, which also has hirtellous herbage, dense capitulescences, and deeply cut corolla lobes. The two species, however, apparently are sympatric in southeastern Arizona, and no evidence of intergra- dation in leaf shape or phyllary morphology has been noted. A contrasting hypothesis regarding the closest relative of /. tenuzsecta follows its description. The type collection of Isocoma halophytica B. Turner was made in the southeastern corner of the range of the species (almost certainly in Chihuahua 106 PHEYTOLOGIA volume 70(2):69-114 February 1991 rather than “Coahuila” as noted on the label, though very near the state line). These plants have densely short stipitate glandular stems and leaves but otherwise fall within the variability of J. plurifolia. Two other collections of the same highly glandular morphotype have been made near Laguna Jaco in the vicinity of the type locality (Stewart & Johnston 1955-GH, LL; Stewart 667-GH), but nearly identical forms occur scattered through the range of the species (e.g., Reeves, Brewster, Presidio, and Hudspeth cos. in Texas, and in Eddy and Chaves counties, New Mexico, particularly in the vicinity of Roswell in the latter). These glands apparently are papillae raised above the leaf surface, perhaps in response to a highly gypseous substrate, such papillae otherwise occurring imbedded on leaf surfaces of many plants lacking a glandular appearance. Both diploids and tetraploids have been reported within Jsocoma plurifo- lia, but these ploidy levels do not appear to be correlated with differences in morphology or geography. 12. Isocoma rusby1 E. Greene Isocoma rusby: E. Greene, Leafl. Bot. Observ. Crit. 1:170. 1906. LECTO- TYPE (designated here): UNITED STATES. Arizona: [Navaho Co.], Holbrook, 20 Aug 1883, H.H. Rusby 651 (US!; Isolectotypes: GH!, ND- G!, NY-2 sheets!, UC). Shrubs 45-90 cm tall. Leaves narrowly elliptic oblong to elliptic obovate, entire, mostly 2-4 mm wide (-10 mm in Coconino Co.), completely glabrous and without marginal cilia, punctate glutinous or less commonly papillate, Ileads 19-25 flowered, the involucres (5.5-) 6.0-9.5 mm long, 5.0-7.5 mm wide; phyllaries narrowly triangular-lanceolate with acute apices, apically glutinous, the hyaline margins often very broad and often minutely ciliate fringed; recep- tacles with long, lanceolate-attenuate, lacerate alveoli. Corollas 5.0-6.5 mim long, the tube 2.8-3.8 mm long, lobes triangular, 0.8-1.1 mm long. Achenes 2.8-3.5 mm long, 8-10 ribbed, the ribs thick, resin filled, and often forming “horns” at the apex. Chromosome number, n = 6 pairs (Solbrig, et al. 1964, as Haplopappus drummondii; Keil 1979, as I. drummondi; Turner & Flyr 1966, as I. heterophylla). Arizona, Utah, New Mexico, and Colorado; rocky or sandy soil, less com- monly in clay, desert shrub communities, usually in saline soil, sometimes with scattered junipers, 750-1500 m; August-October (-November). This distinctive species has been mistakenly identified both as Jsocoma drummondii and J. plurifolia. See comments following J. azteca regarding putative intermediates between it and J. rusbyz. Only one specimen of Isocoma rusby: from Colorado has been seen in the present study (“S.W. Colorado, Nesom: Taxonomy of Isocoma 107 near the Utah line,” Aug 1875, Brandegee 1208-NY), although it may be more abundant in that state. 13. Isocoma tehuacana Nesom. Isocoma tehuacana Nesom, sp. nov. TYPE: MEXICO. Puebla: Tehua- can, Dec 1841, Liebmann 526 (HOLOTYPE: NY!; Isotypes: GH!, GH- tracing and fragment ez herb. Klatt!, MO!). Isocomae venetae (Kunth) E. Greene similis sed vestimento pu- berulo caulium ac foliorium, tubis sericeis corollarum, et acheniis longioribus differt. Subshrubs 2-3 dm tall, the stems and leaves densely short puberulous with hairs with thick, orange resinous bases quickly tapered to filiform, whitish, crisped apices. Leaves narrowly oblanceolate to oblanceolate, 1-2 cm long, 2-3 mm wide, with 1-3 pairs of aristate teeth. Heads broadly turbinate, basally obtuse, ca. 20 flowered, the involucres 4.5-6.0 mm high, 6-7 mm wide; phyl- laries glabrous, each with a distinct, dark greenish, punctate glandular apical area, the apex blunt to rounded, the margins with a very narrow, scarious rim, sometimes distally ciliate fimbriate; receptacles with lanceolate-lacerate alve- oli. Corollas 5.5-7.0 mm long, the tube 3.0-4.0 mm long, prominently sericeous, the lobes deltate, 0.6-0.9 mm long. Achenes 2.8-4.0 mm long, densely sericeous, with 4-6 resinous ribs. Chromosome number unknown. Known only from the type collection. The plants represented in this collection are from slightly outside the south- ern periphery of the range of [socoma veneta (Map 4) and they differ from it in the puberulent vestiture of their stems and leaves, sericeous corolla tubes, and longer achenes. Eleven branches are included on the type sheets, and the Klatt tracing shows yet another apparently from a European herbarium. It is unusual that this taxon is known from only a single, historical collection, but it is clearly outside the range of morphological variation in its widespread and much more common relative, J. veneta. Cronquist 11243(NY) from near San Sebastian El Seco, Puebla, has narrow leaves and somewhat puberulent stems, but the leaves are hispidulous and in all other features as well, it is typical of I. veneta. This is the only collection of the latter species observed during this study that has even slightly puberulent vestiture. It might be interpreted as intermediate in this respect, but overall, it clearly belongs with /. veneta rather than the plants from Tehuacan. 108 PHYTOLGGT:A volume 70(2):69-114 February 1991 14. Isocoma tenuisecta E. Greene Isocoma tenuisecta E. Greene, Leafl. Bot. Observ. Crit. 1:169. 1906. LEC- TOTYPE (designated by Benson 1940): UNITED STATES. Arizona: mesas about Tucson, 10 Sep 1867, C. Smart (ND-G; Isolectotype: US!). (H)Aplopappus tenuisectus (E. Greene) S.F. Blake ez Benson, Amer. J. Bot. 27:188. 1940. The ND-G specimen chosen as the lectotype by Benson has not been relocated in the present study. Isocoma fruticosa Rose & Standl., Contr. U.S. Natl. Herb. 16:18. 1912. TYPE: MEXICO. Sonora: MacDougal Pass, near the Pinacate Mts, 14 Nov 1907, D.T. MacDougal s.n. (HOLOTYPE: US). (H)Aplopappus fruticosus (Rose & Standl.) S.F. Blake, Contr. U. S. Natl. Herb. 23:1493. 1926. Plants minutely hispidulous on the stems and leaves, usually densely so, commonly strongly glutinous. Leaves pinnatifid, 2.0-3.5 cm long, the blade and lobes 0.5-2.0 mm wide. Heads 8-12 (-15) flowered, the involucres 4.0-6.5 mm long, 2.0-2.8 mm wide; phyllaries narrowly oblong-lanceolate, the inner with broad, scarious margins, apices with a small, sharply delimited, nonaristate, green resinous area, often distinctly thickened and approaching a resin pocket; receptacles with low, broad, but acute alveoli. Corollas 4.5-6.0 mm long, the tube 3.0-3.5 mm long, the lobes triangular, 0.8-1.2 mm long, 1/3-1/2 the length of the limb. Achenes (2.0-) 2.5-3.1 mm long, with 6-8 thin, resinous ribs, lightly to densely sericeous. Chromosome number unknown. Southern Arizona, New Mexico, and northern Sonora; sandy soil, gravelly hills, grasslands, most commonly in matorral or stands of Larrea; 750-1600 m; September-November. Isocoma tenuisecta is known from México only by the type collection of J. fruticosa. “Haplopappus hartwegii var. tenuisecta” and “Bigelovia hartwegii var. tenuisecta,” names appearing on a few collections and both attributed to A. Gray, were apparently never published. See further comments following /. plurifolia. Isocoma tenuisecta resembles J. plurifolia in its relatively few flowered heads in dense, corymboid capitulescences and its long corolla lobes. On the other hand, the minutely hispidulous vestiture and phyllary morphology are more similar to that of J. acradenia than that of J. plurifolia. Although J. tenuisecta produces thickened phyllary apices throughout its range, often approaching resin pockets, they appear to be more strongly developed where the species is contiguous in geography with J. acradenia var. acradenia in southwestern Ari- zona, particularly in the area of Organ Pipe National Monument (Pima Co.). For example, the leaves of Felger 87-272 (TEX) and Clark 10994 (GI) are glutinous and densely hispidulous as in J. tenuisecta, but they are primarily linear with minute, shallow teeth or short, linear spreading lobes, or on some Nesom: Taxonomy of [socoma 109 plants the upper leaves are entire, the lower pinnatifid. Further, these individ- uals have phyllaries with distinctive resin pockets. Similar plants have been collected to the north in Pima Co. in the vicinity of Cubo (Clark 11111-GH, 11119-GH) and slightly further in Maricopa Co. around Sentinel (Hall 11032- GH; Jones 25086-LL, NY, US); typical var. acradenia also occurs in these areas. In the early stages of this study, some of the specimens from Sentinel were annotated as /. acradenia var. eremophila, but it seems more reason- able to regard them as intermediates between var. acradenza and J. tenutsecta. They are mapped (Map 1) as J. tenuzsecta. 15. Isocoma tomentosa Nesom. Isocoma tomentosa Nesom, sp. nov. TYPE: MEXICO. Chihuahua: Bajos de San Diego, 1.8 km E of San Diego de Alcala, ca. 16 km NE of jct of local road with Hwy 45 (the jct 19.5 km SE of jct of Hwy 45 and road to Aquiles Serdan, SE of Cd. Chihuahua); ca. 1200 m, abundant on gyp hill, most past flower; 14 Oct 1986, G. Nesom 5478 with L. Vorobik (HOLO- TYPE: TEX!; Isotypes: ARIZ!, ASU!, CIIDIR!, COLO!, ENCB!, GH!, KANU!, MEXU!, MO!, NMC!, NY!, RM!, S!, TEX!, UC!, US!, WIS!). Isocomae plurifoliae (Torr. & A. Gray) E. Greene similis sed vestimento dense albo-tomentoso caulium foliorum ac phyllario- rum, foliis marginibus profunde dentatis, capitulis majoribus, corol- larum tubis sparsim sericeis, et acheniis longioribus differt. Caespitose subshrubs 1.5-7.0 dm tall, not at all glutinous. Stems mod- erately to densely and closely tomentose-puberulous with distinctly whitish hairs. Leaves oblanceolate to narrowly oblanceolate, 10-25 mm long, 2-6 mm wide, with (1-) 2-8 pairs of spreading, blunt to aristate teeth, tomentose- puberulous, less commonly hirtellous, sometimes sparsely and minutely stip- itate glandular. Heads arranged in a relatively few headed corymboid capit- ulescence, campanulate, basally rounded, with 17-27 flowers per head, the in- volucres 4.0-7.0 mm long, 4.5-7.0 mm wide; phyllaries relatively thin textured, the apical area green, not punctate, the apex rounded, not aristate, densely to moderately or sparsely white puberulous; receptacles with low, circular alve- oli. Corollas 5.5-7.0 mm long, the tube 2.8-3.5 mm long sparsely sericeous, the limb 2.0-3.2 mm long, the lobes 1.0-1.5 mm long, ca. (1/3-) 2/5-1/2 the length of the limb. Achenes narrowly turbinate, 2.0-3.0 mm long, densely sericeous, with 6-8 thin, slightly resinous ribs; pappus of numerous, slender, white, barbellate bristles 4.5-5.5 mm long. Chromosome number unknown. Known only from the type locality at Banos de San Diego, Chihuahua, on a hill of exposed gypsum with numerous other gypsophilic perennials, just W of an area of hot springs and small streams flowing into salt flats, ca. 1230 m at the hill crest. 110 PHY TOLOGHA volume 70(2):69-114 February 1991 Isocoma tomentosa occurs at the southern periphery of the geographical range of J. plurifolia and is similar to it in its deeply cut corolla lobes. On the basis of geography, the two would be suspected to be related as sister taxa, but the new species is unexpectedly different in morphology. Jsocoma tomentosa differs from /. plurifoha in the white pubescent vestiture of its stems, leaves, and phyllaries, toothed leaves, larger heads with more flowers, arranged in much smaller capitulescences, low circular alveoli, corollas with sparsely sericeous tubes, and longer achenes. The type collection represents 39 separate plants, including one entire plant (mounted as a TEX isotype) and 38 additional branches, each of the latter removed at the base from a different plant and selected to show the range of variation in vestiture and leaf morphology in the population. The species is characterized by a closely white tomentose vestiture and regularly serrate leaves, but a low percentage of the plants (3 of the 39) have entire, hirtellous leaves, very similar to some forms of I. plurifolia. Even these, however, which have been annotated as /. tomentosa “ > plurifolia,” have sparsely puberulous stems and 18-21 flowered heads; in Arizona the stems of J. plurifolia may be sparsely puberulous, but the large heads of J. tomentosa would be unusual anywhere in the range of J. plurifolia. Collections of typical Jsocoma plurifolia have bees made in the area of San Diego (plains near San Diego, 10 Sep 1891, Hartman 759-GH; 31 km N of Julimes, Johnston, et al. 12344-LL; Meoqui, 18 Aug 1935, Te Sucang 180- GH, TEX). The plants of all of these have very small heads with few flowers and small achenes (phyllaries glutinous, the longest 3 mm long, flowers 10- 12 per head, achenes 1.5-1.8 mm long, with pappus 3-4 mm long) and are clearly outside the range of any of the plants of J. tomentosa. It is possible, however, that variation among the plants of J. tomentosa toward I. plurifolia is influenced by genes from these nearby populations of the latter. 16. [socoma veneta (Kunth) E. Greene Isocoma veneta (Kunth) E. Greene, Erythea 2:111. 1894. BASIONYM: Bac- charis veneta Kunth, Nov. Gen. Sp. [folio] 4:53. 1818; [quarto] 4:68. 1820. TYPE: MEXICO. [Morelos]: near Cuernavaca, Humboldt & Bon- pland s.n. (HOLOTYPE: P, P-fiche!). Aster venetus (Kunth) O. Ktze., Rev. Gen. 318. 1891. (H)Aplopappus venetus (Kunth) S.F. Blake, Contr. U.S. Natl. Herb. 23:1492. 1926. (H)Aplopappus discoideus DC., Prodr. 5:350. 1836. LECTOTYPE (designated here): MEXICO. 1831, Alaman s.n. (G-DC fiche!, photo-US!). DeCandolle also cited several other specimens from different collectors, all clearly shown on the fiche of the G-DC col- lections. Nesom: Taxonomy of Isocoma 111 Linosyris mezicana Schlecht., Linnaea 14:Litt.-Ber. 128. 1840. TYPE: MEXICO. Illustration in Ind. Sem. Hort. Hal. 1839:9, pl. 4. 1840. Subshrubs 0.3-0.7 m tall, with stems and leaves sparsely to moderately minutely hispidulous. Leaves fleshy, mostly obovate-cuneate, 2-6 mm wide, with 1-4 pairs of shallow, short spinescent teeth. Heads 17-26 flowered, hemi- spheric to broadly turbinate, rounded to acute at base, the involucres 5-7 mm long, 4-5 mm wide; phyllaries glabrous, the lower part indurated stramineous, the apex greenish, glandular punctate and glutinous. Corollas 4.2-6.0 mm long, the tube 2.0-3.5 mm long, the lobes deltate to deltate triangular, 0.6-1.0 mm long. Achenes 1.6-2.8 mm long, with 3-6 thick ribs and several, mostly subepidermal nerves on the faces, densely sericeous. Chromosome number, n = 6 pairs (Anderson, et al. 1974; Powell & Turner 1963). Coahuila, Nuevo Leon, Tamaulipas, Zacatecas, San Luis Potosi (northern system), and Hidalgo, México, Distrito Federal, Morelos, Tlaxcala, Puebla, Veracruz, [Oaxaca?, Guerrero?, see comments below] (southern system); sandy loam, volcanic or limestone derived soil, saline flats, grassland, matorral; 2250- 2850 m; June-November (-December, -February). Isocoma veneta comprises two population systems slightly separated from each other in central México. Plants from the southern system (including the type locality) have leaves that range larger than those from the north. Leaves of northern system are 5-16 mm long, 2.4-5.0 times longer than wide; those of southern system are 10-35 mm long, 4.2-6.0 (-8) times longer than wide, but if 8 times then the leaf barely lobed and on a branch with relatively wider leaves. Several collections from around Esperanza, Puebla (e.g. Purpus 2633-GH, MEXU, MO, US) have more deeply incised leaves, approaching the morphology of IJ. hartwegii, but then others from the same vicinity grade into the typical form. A specimen noted as having been collected in Guerrero (Cerro del Pino, 14 Jul 1940, Miranda 460, MEXU) was probably instead taken in the state of México (see, for example, collections from Cerro de los Pinos, Edo. Mexico, MEXU, NY, US). A collection of typical Jsocoma veneta is reported on the label to have been collected in Oaxaca (Tlapujahua, 1850, Keerl s.n.-GH). No other collections from Oaxaca of this species have been discovered, and, like that from Guerrero, the Keerl record needs to be corroborated. Additional comments regarding the relationship of Jsocoma veneta to I. menziesi and I, hartwegii follow the latter two species. ACKNOWLEDGMENTS I thank B.L. Turner and P.O. Karis for reviews of the manuscript, Barney Lipscomb for his help in obtaining pertinent literature, Rich Spellenberg for 112 PHYTOLOGIA volume 70(2):69-114 February 1991 information regarding localities in New Mexico, John Pruski for his essential assistance in clarifying the identity of [socoma plurifola, and Barbara Hel- lenthal for help in studying /socoma in the ND-G herbarium of E.L. Greene. I thank Dr. John Kartesz for pointing out a potential nomenclatural prob- lem. Loans of specimens from ARIZ, GH, MO, MU, ND-G, NMC, NY, UCR, and US are appreciated; additional information has been recorded from the specimens of Jsocoma at MEXU during a visit there. In all, more than 1600 collections have been examined in this study. LITERATURE Anderson, L.C., D.W. Kyhos, T. Mosquin, A.M. Powell, & P.H. Raven. 1974. Chromosome numbers in Compositae. IX. Haplopappus and other Aster- eae. Amer. J. Bot. 61:665-671. Benson, L. 1940. Taxonomic contributions. Amer. J. Bot. 27:186-190. Blake, S.F. 1926. Aplopappus in P.C. Standley, Trees.and Shrubs of Mezico. Contr. U.S. Natl. Herb. 23:1486-1495. Clark, W.D. 1979. The taxonomy of Hazardia (Compositae: Astereae). Madrono 26:105-127. Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Teras. Texas Research Foundation, Renner, Texas. De Jong, D.C.D. & F.H. Montgomery. 1963. Chromosome numbers in some Californian Compositae-Astereae. Aliso 5:255-256. Geiser, S.W. 1948. Naturalists of the Frontier (ed. 2). Southern Methodist Univ. Press. Dallas, Texas. Great Plains Flora Association. 1977. Atlas of the Flora of the Great Plains. Iowa State Univ. Press, Ames. Greene, E.L. 1894. Observations on the Compositae.— VII. Erythea 2:105- 112. . 1906. New species of Isocoma. Leafl. Bot. Observ. Crit. 1:169-173. Hall, H.M. 1907. Compositae of Southern California. Univ. California Publ. Bot. 3:3-296. 1928. The genus Haplopappus (Section 13. Isocoma). Carnegie Inst. Washington Publ. 389:222-243. Nesom: Taxonomy of Isocoma 113 Hartman, R.L. 1990. A conspectus of Machaeranthera (Asteraceae: Aster- eae). Phytologia 68:439-465. & M.A. Lane. 1991. A natural intergeneric hybrid in the z = 6 group of Astereae. Sida 14(3), in press. Hoover, R.F. 1970. The Vascular Plants of San Luis Obispo County, Cali- fornia. Univ. of California Press, Berkeley. Jackson, R.C. 1959. Jn “Documented chromosome numbers of plants.” Madrono 15:49-52. . 1966. Some intersectional hybrids and relationships in Haplopap- pus. Univ. Kansas Sci. Bull. 46:475-485. & C.T. Dimas. 1981. Experimental evidence for systematic place- ment of the Haplopappus phyllocephalus complex (Compositae). Syst. Bot. 6:8-14. Jepson, W.L. 1925. A Manual of the Flowering Plants of California. Asso- ciated Student’s Store, University of California, Berkeley. Kearney, T.H. & R.H. Peebles. 1951. Arizona Flora. Univ. California Press, Berkeley. Keil, D.J. 1979. In IOPB chromosome number reports LXIII. Taxon 28:271- Bia: Martin, W.C. & C.R. Hutchins. 1980. A Flora of New Mezico. J. Cramer, Vaduz. McVaugh, R. 1984. Compositae. Flora Novo-Galiciana 12:1-1157. Nesom, G.L., D.R. Morgan, Y. Suh, & B.B. Simpson. 1990. Xylothamia (Asteraceae: Astereae), a new genus related to Euthamia. Sida 14:101- 116. Nesom, G.L., Y. Suh, & B.B. Simpson. Submitted. Phylogenetic position of the genus Stephanodoria (Compositae: Astereae) with evidence from chloroplast DNA, chromosome number, and morphology. Brittonia. Osterhout, G.E. 1920. Rocky Mountain botany and the Long Expedition of 1820. Bull. Torrey Bot. Club 47:555-562. Pinkava, D.J. & D.J. Keil. 1977. Chromosome counts of Compositae from the United States and Mexico. Amer. J. Bot. 64:592-596. 114 PHY TOL OG volume 70(2):69-114 February 1991 Powell, A.M. & B.L. Turner. 1963. Chromosome numbers in the Compositae. VII. Additional species from the southwestern United States and Mexico. Madrono 17:128-140. Powell, A.M. & S.A. Powell. 1977. Chromosome numbers of gypsophilic plant species of the Chihuahuan Desert. Sida 7:80-90. . 1978. Chromosome numbers in Asteraceae. Madrono 25:160- 169. Raven, P.H., O.T. Solbrig, D.W. Kyhos, & R. Snow. 1960. Chromosome numbers in Compositae. I. Astereae. Amer. J. Bot. 47:124-132. Semple, J.C. 1985. Chromosome number determinations in fam. Compositae tribe Astereae. Rhodora 87:517-527. , J.F. Chmielewski, & M.A. Lane. 1989. Chromosome number de- terminations in fam. Compositae, tribe Astereae. III. Additional counts and comments on generic limits and ancestral base numbers. Rhodora 91:296-314. Shinners, L.H. 1950. Notes on Texas Compositae. IV. Field & Lab. 18:25-32. Solbrig, O.T., L.C. Anderson, D.W. Kyhos, P.H. Raven, & L. Rudenberg. 1964. Chromosome numbers in Compositae. V. Astereae II. Amer. J. Bot. 51:513-519. Turner, B.L. 1972. Two new species of Isocoma (Compositae—Astereae) from north-central Mexico. Sida 5:23-25. Turner, B.L. & D. Flyr. 1966. Chromosome numbers in the Compositae. X. North American species. Amer. J. Bot. 53:24-33. Turner, B.L., A.M. Powell, & T.J. Watson. 1973. Chromosome numbers in Asteraceae. Amer. J. Bot. 60:592-596. Urbatsch, L.E. 1974. In IOPB chromosome number reports XLV. Taxon 23:619-624. 1975. First chromosome number reports for some Compositae. Southw. Nat. 19:283-285. Weedin, J.F. & A.M. Powell. Jn IOPB chromosome number reports LX. Taxon 27:223-231. Welsh, S.L. 1983. Utah Flora: Compositae (Asteraceae). Great Basin Nat. 43:179-357. Phytologia (February 1991) 70(2):115-118. CAREX BICKNELLI, “BICKNELL’S SEDGE” (CYPERACEAE): NEW TO TEXAS, WITH A KEY TO TEXAS SPECIES OF SECTION OVALES ‘Stanley D. Jones & *Anton A. Reznicek 'S.M. Tracy Herbarium (TAES), Department of Range Ecology and Management, Texas A&M University, College Station, Texas 77843 U.S.A. & ?University of Michigan Herbarium (MICH), Ann Arbor, Michigan 48109-1057 U.S.A. ABSTRACT Carez bicknellii Britt., section Ovales (Cyperaceae), previously un- substantiated in Texas, has been found in the following two northeast Texas counties: Delta and Lamar. KEY WORDS: Carez, Carer bicknellu, section Ovales, Cyper- aceae, Texas Carer bicknellit Britt. “Bicknell’s Sedge” is one of eleven species in sec- tion Ovales Kunth of the subgenus Vignea found in Texas. They are Carer alata Torr., C. albolutescens Schwein., C. athrostachya Olney, C. bicknelliz, C. brevor (Dewey) Mackenzie ez Lunell, C. brittoniana Bailey, C. festucacea Schkuhr ez Willd., C. hyalina Boott, C. longit Mackenzie, C. reniformis (Bai- ley) Small, and C. tribuloides Wahl. Section Ovales, represented by about seventyfive species (Mackenzie 1931), is the largest group of carices in North America. Correll & Johnston (1970) state that a report of Carex bicknelli in Texas was based on a specimen of C. reniformis. This is consistent with the authors’ findings of Texas specimens labeled C. bicknelliz. However, recent specimens have been collected that support its existence in Texas. Mackenzie (1931) gives the distribution for C. bicknelliz from the valley of Penobscot River, Maine to Saskatchewan, Canada and southward to Delaware, Arkansas, and Oklahoma. He goes on to list specimens, mentioning New Mexico but not Texas. Fernald (1950), Steyermark (1968), and Correll & Correll (1972) all mirror Mackenzie’s Ags 116 PAY O01 G TA volume 70(2):115-118 February 1991 distribution. Neither Mahler (1988) nor Hatch, et al. (1990) list this species as occurring in Texas. Specimens collected: UNITED STATES. Texas: Delta Co.: 14 May 1989, S. & G. Jones 2887 & T. Powell (MICH, TAES). The collection site is 0.1 of a mile south on Farm Road 1529 from its junction with Hwy 154, E of Cooper. The habitat is an open hydric roadside ditch with the soils in the Kaufman series. They are moderately well drained, dark gray to black in color and slightly acid. The geology of the site is of the Marlbrook Marl (kmb) formation (Cretaceous). Associated species include Carex crus-corvi Shuttlw. ez Kunze, C. hyalinolepis Steud., C. franki Kunth, C. triangularis Boeckl., Scirpus pendulus Muhl., Hordeum pusillum Nutt., Oenothera speciosa Nutt., and Lolium perenne L. Lamar Co.: 14 May 1989, S. & G. Jones 2882 & T. Powell (MICH). The collection site is 5.2 miles north on Farm Road 1184/1497 from its junction with the extension of Farm Road 1184 at Auds Creek, S of Paris. The habitat is an open mesic-hydric roadside ditch with the soils in the Houston Black-Leson-Heiden Series. They are moderately well drained, dark gray in color and moderately alkaline. The geology of the site is of the Gober Chalk (kg) formation (Cretaceous). Associated species are Carez crus-corvi, C. vulpinoidea Michx., Scirpus pendulus, Lolium perenne, Physostegia angustifolia Fern., Salix nigra Marsh., Daucus carota L., Juncus spp., Rumez sp., and Vicia spp. The following is a key to the section Ovales in Texas. Mature specimens are needed for correct identification. KEY TO THE SECTION OVALES OF TEXAS 1. Spikelets reddish brown; lower inflorescence bract leaflike and longer than the inflorescence, 2. 22 hen22). wegen, a eee C. athrostachya / . . . . . I’ Spikelets green to stramineous; lower inflorescence bract inconspicuous and much shorter than the inflorescence. 2. Perigynia with oblanceolate bodies mostly less than 1.5 mm wide. PROM Cr Re ES Ce Ee ITO od dickens scarce C. tribuloides 2’ Perigynia with ovate, obovate, orbicular, or even reniform bodies 1.5-6.0 mm wide. 3. Pistillate scales (at least the upper) scabrous awned; perigy- nia widest well above the middle; inner bands of sheaths green. \. see a etek: See + Ghee C. alata ese 3’ Pistillate scales obtuse to + acuminate, but never scabrous awned; perigynia and sheaths various. 4. Perigynia with several prominent nerves over the achene on the inner (ventral) surface. Jones & Reznicek: Carex bicknellit in Texas alee 5. Perigynium beaks less than 1.5 (-1.8) mm long; peri- gynia usually more than 30 per spike; spikes 3-8. 6. Longest perigynia 5-6 mm long. ...... C. bicknellia 6’ Longest perigynia 2.6-4.6 mm long. 7. Perigynia widest above the middle, the bodies + obovate. 8. Styles straight to somewhat sinuous; peri- gynium beaks + gradually tapering from wid- est point of body, appressed. ..... C. longi 8’ Styles abruptly contorted just above the a- chene; perigynium beaks abruptly tapered to a long tip, spreading. ..... C. albolutescens 7’ Perigynia widest at or below the middle; the bo- dies Erorbiculamwoa. 26G0.8 ee oe C. festucea 5’ Perigynium beaks 1.5-2.5 mm long; perigynia 15-25 per spike: spikes 22°47 020. cone cance s Se seee C. hyalina 4’ Perigynia nerveless or rarely with 1-3 faint nerves over the achene on the inner (ventral) surface. 9. Perigynia papillose (at 30 X); bodies usually wider than long, at least on lower perigynia. .... C. reniformis 9’ Perigynia not papillose; bodies usually as wide as long or narrower. 10. Perigynia 1.5-3.5 mm wide. 11. Achenes 1.7-2.0 mm long, perigynia 3.4-4.5 mm long and 2.4-3.2 mm wide. ....... C. brevior 11’ Achenes 1.3-1.7 mm long; perigynia 2.6-3.6 mm long and 1.5-2.4 mm wide. .... C. festucacea 10’ Perigynia 3.7-6.0 mm wide. ........ C. brittoniana In our area, Carez bicknellit superficially resembles C. reniformis and C. brevior. However, a closer examination will differentiate C. bicknelli: from those two taxa. The perigynia of C. reniformis is at least as wide as long, while those of C. brevior are about as wide as long, both lack nerves on the ventral surface over the achene. Carez bicknellu differs in having perigynia longer than wide with nerves on both faces. Steyermark (1968) provides the following diagnostic traits for C. bicknelli, “This is a fairly tall-growing Carez, attaining a height of 1 meter with the flowering culms, which greatly exceed the narrow leaves. The thin, transparent, large perigynia, mostly 5.5-7.5 mm long by 2.7-4.8 mm wide, many-nerved on each face, and the silvery brown 118 PHY POL OGIA volume 70(2):115-118 February 1991 or straw-buff light colored scales and perigynia are marks of recognition.” However, we have seen perigynia no longer than 6 mm. Based on Hermann’s (1972) description of Carez bicknelli var. opaca F .J. Herm. from Prairie and Lonoke counties, Arkansas, ours is of the typical va- riety, C. bicknelliz var. bicknelliz. Carex bicknelli var. opaca differs by having the perigynia strongly concave rather than flat. The perigynia are opaque (except for the outer margin), faintly if at all nerved ventrally, corky between the achene and wing, and the inner margin green, with only the outer hyaline. An unusual specimen Kessler 3331 (TAES) from Harris County, TX will key to C. bicknellii in our key, but differs in having perigynia with ovate bodies only 1.8-2.4 mm wide. Its identity is, as yet, unclear. ACKNOWLEDGMENTS We extend our appreciation and thanks to Larry E. Brown (SBSC), Charles T. Bryson (USDA, SWSL), Gretchen D. Jones, and J.K. Wipff (TAES) for their critical reviews of this manuscript. LITERATURE CITED Correll, D.S. & H.B. Correll. 1972. Aquatic and Wetland Plants of South- western United States. Stock #5501-0177. Environmental Protection Agency Research and Monitoring, Washington, D.C. Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Tezas. Texas Research Foundation, Renner, Texas. Fernald, M.L. 1950. Gray’s Manual of Botany, 8th ed. Van Nostrand and Reimhold Company, New York. Hatch, S.L., K.N. Gandhi, & L.E. Brown. 1990. Checklist of the Vascular Plants of Texas. Tex. Agric. Exp. Sta. Bull. MP-1655. Hermann, F.J. 1972. A new variety of Carez bicknellii from Arkansas. Sida 5(1):49. Mackenzie, K.K. 1935. North American Flora (Poales) (Cyperaceae). 18:393- 478. The New York Botanical Garden, New York. Mahler, W.F. 1988. Shinner’s Manual of the North Central Texas Flora. S.M.U. Herbarium, Dallas, Texas. Steyermark, J.A. 1968. Flora of Missouri. The lowa State University Press, Ames, Iowa. Phytologia (February 1991) 70(2):119-125. THE DISTRIBUTION OF SARRACENIA IN LOUISIANA, WITH DATA ON ITS ABUNDANCE IN THE WESTERN PART OF THE STATE M.H. MacRoberts & B.R. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. ABSTRACT Sarracenia occurs in bogs in southeastern and western Louisiana. Bogs are locally common but few have escaped damage and the vast majority have been destroyed or are degraded. KEY WORDS: Bog, pitcher plants, Sarracenia, Sarraceniaceae, Louisiana INTRODUCTION There has been almost nothing published on the distribution, abundance, or condition of bogs in Louisiana. So little attention has been paid to this plant community that as late as 1977, Folkerts, in his extensive review of en- dangered and threatened carnivorous plants of North America was unable to determine whether or not Sarracenia alata Wood, the sine qua non of bog indicator species in Texas and Louisiana, was endangered west of the Missis- sippi River delta — a point reiterated more recently by Frost, et al. (1986). As part of a continuing study of bogs, we present information on the distribution and abundance of Sarracenia in Louisiana (MacRoberts & MacRoberts 1988, 1990a, 1990b). METHODS We examined all specimens of Sarracenia (254 sheets) in Louisiana herbaria and in several national collections (DUKE, GH, LAF, LSU, LSUE, LSUS, LTU, MO, NATC, NLU, NO, NOLS, SFRP, US, USLH), and studied the literature on Sarracenia distribution (McDaniel 1971; Murry & Urbatsch 1979). In order to estimate abundance, we surveyed bogs in Natchitoches Parish. 119 120 PHRYTOLOGIA volume 70(2):119-125 February 1991 RESULTS AND DISCUSSION Figure 1 shows the distribution of Sarracenia alata and S. psittacina Michx. in Louisiana. Sarracenia alata occurs in five western and three southeastern parishes, and S. psittacina occurs in two southeastern parishes (Appendix 1). In the west, pitcher plant populations occur in southern Natchitoches Parish and are scattered throughout Vernon and Beauregard parishes. Sarracenia alata also occurs in a few localities in southern Sabine Parish. Other popu- lations occur in three localities in northern Natchitoches and in one locality in southern Bienville Parish. In southeastern Louisiana, S. alata occurs in Tangipahoa, Washington, and St. Tammany parishes. Sarracenia psittacina occurs in Tangipahoa and St. Tammany parishes. Only Sarracenia alata occurs in western Louisiana and reports of other taxa can be discounted. In a few bogs in the vicinity of Highway 117 near the Natchitoches- Vernon Parish boundary, several Sarracenia species (S. psittacina, S. minor Walt., S. leucophylla) have been introduced. The report of S. flava L. from Lincoln Parish (Roland 1966) is an error. We examined these speci- mens (Rabum s.n. [LTU]) and found that they are all S. alata that had been planted on a pond bank (Hartsell 1980; Don Rhodes, pers. comm.). The S. flava reported by Correll & Correll (1941) for Natchitoches and Tangipahoa parishes (Correll & Correll 9762, 10540 |DUKE]}) are both S. alata. Feather- man (1872) reported S. flava from Calcasieu Parish (see Murry & Urbatsch 1979). We examined this specimen (Featherman s.n. [LSU]); it is S. alata. Further, although the specimen is labeled “Calcasieu,” in 1871 when it was collected, Calcasieu included what today comprises four parishes: Calcasieu, Beauregard, Allan, and Jefferson Davis. As no Sarracenia have been found more recently in Calcasieu Parish (Joe Bruce, pers. comm., Steve Orzell, pers. comm.), the Featherman specimen probably came from what is now Beau- regard Parish. Pitcher plants have been reported also from Rapides Parish (Allen, et al. 1988). However, we could find no voucher specimen for that parish, and Allen (pers. comm.) acknowledged the report to have been in er- ror although what appears to be habitat suitable to Sarraceniais not lacking in Rapides Parish. In southeastern Louisiana, Brown (1972) reports S. psittacina from Washington Parish and MacRoberts (1989) reiterates. However, we could find no voucher specimen for that parish. We have had verbal reports of S. alata occurring in Red River and Rapides parishes, but we were unable to find voucher specimens or to verify its occurrence in the field at reported locations. There are two voucher specimens of Sarracenia purpurea L. for Louisiana (MacRoberts & MacRoberts 1988). Both were collected in the last century and come from the southeastern part of the state. One comes from St. Tammany Parish but, unfortunately, the exact location of the other one is not known. How common is Sarracenia alata in Louisiana? In the western part of the state, it is confined to bogs (see Appendix 2). Bridges & Orzell (1989) surveyed Fig. 1. Distribution of Sarracenia. Kee Rie &. 122 PHY TOLOGIA volume 70(2):119-125 february 1991 99 relatively high quality (less disturbed) bogs in the longleaf pine region of southeastern Texas and southwestern Louisiana and found S. alata in 68% of them. We are conducting a survey of bogs in the Kisatchie District of the Kisatchie National Forest in southern Natchitoches Parish and found S. alata in 33 (24%) of 140 bogs as of 11 February 1991. The difference in frequency of S. alata in bogs in these two studies is probably due to a combination of different surveying methods, variation in definitions of “bogs,” and the fact that our survey has been confined to the extreme northern range of the species. Bogs are small, ranging in Natchitoches Parish up to 4 ha but averaging less than 1 ha. The 33 bogs in Natchitoches Parish that have Sarracenia are about average in size. The bogs examined by Bridges & Orzell (1989: 254) appear to be about the same size as these. Therefore, bog habitat and bogs with Sarracenia are not particularly common. More important, however, as Bridges & Orzell (1989) emphasize, there are only a few relatively undisturbed (high quality) bogs left, the majority having been either destroyed or degraded beyond recovery. Of those that are in good condition, many are threatened by ditching, farming, grazing, logging, and fire suppression. Fortunately, recent conservation efforts in both Louisiana and Texas have resulted in the preser- vation of some of the best examples of bogs (Anon. 1990; Fritz & Alford 1986; Parvin 1989). There is no published information on the status of Sarracenia populations in southeastern Louisiana (Folkerts 1982). From our own observations and from talking with botanists and naturalists, the savannah and hillside bogs where Sarracenia grow are vanishing rapidly from the same causes that have affected the western bogs (Rebertus & Barker 1984). Efforts should be made to secure some of this habitat. ACKNOWLEDGMENTS We gratefully acknowledge the staff of the Kisatchie National Forest for their cooperation in all phases of our work and the curators and staff of herbaria who provided specimens, photocopies, and information on Sarracenia and other plants. Our thanks to the many individuals who helped us during the study, most especially Charles Allen, B. Bergeron, Edwin Bridges, Joe Bruce, Jim Caldwell, Ella Edwards, Chip Ernst, Tom Fair, Florence Givens, Walter Holmes, Tony Hough, Jessie Johnson, Richard Johnson, D.T. Mac- Roberts, Ben Martin, Nelwyn McInnis, Smith Mullens, Robert Murry, Steve Orzell, Don Rhodes, Latimore Smith, R. Dale Thomas, and Lowell Urbatsch. MacRoberts & MacRoberts: Sarracenia in Louisiana 123 APPENDICES Appendix 1. The following is a parish list of Sarracenia alata with one voucher specimen cited per parish: Beauregard, Thomas 24021 (NLU); Bienville, Holmes 3852 (NLU); Natchitoches, Lynch 2033 (LSU); Sabine, Carroll 1777 (NLU); St. Tammany, Kral 16512 (LAF); Tangipahoa, Cruz & Landry s.n. (LAF); Ver- non, Givens 2000 (LSU); Washington, Adams 1724 (LTU). The parishes with documented Sarracenia psittacina are: St. Tammany, Thieret 23374 (LAF); and Tangipahoa, Brown 18370 (LSU). Appendix 2. Bog habitat terminology has not been standardized in Louisiana or in the Southeast, and it probably will not be for some time (see Craig, et al. 1987; Smith 1988; Bridges & Orzell 1989; Frost, et al. 1986). We do not undertake to add to terminological problems here, but lump all types and proposed types of bog under the simple label “bog.” LITERATURE CITED Allen, C.M., C.H. Stagg, & S.D. Parris. 1988. Analysis of the vegetation in pitcher plant bogs in two baygalls at Fort Polk in west central Louisiana. Proc. Louisiana Acad. Sci. 50:1-6. Anon. 1990. Landmarks: A report on Conservancy actions and accomplish- ments. The Nature Conservancy Magazine 40(2):22. Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the west gulf coastal plain. Natural Areas Journal 9:246-263. Brown, C.A. 1972. Wildflowers of Louisiana and Adjoining States. Louisiana State University Press, Baton Rouge. Correll, D.S. & H.B. Correll. 1941. A collection of plants from Louisiana. Amer. Midl. Nat. 26:30-64. Craig, N., L.M. Smith, N.M. Gilmore, G.D. Lester, & A.M. Williams. 1987. The natural communities of coastal Louisiana: Classification and de- scription. Louisiana Natural Heritage Program, Louisiana Department of Wildlife and Fisheries. 124 PHY TO10'G1A volume 70(2):119-125 February 1991 Featherman, A. 1872. Third annual report of botanical survey of southwest and northwest Louisiana made during the year 1871. /n Annual Report of Prof. D.F. Boyd, superintendent, Louisiana State University, for the year 1871. Pp. 101-161. Folkerts, G.W. 1977. Endangered and threatened carnivorous plants of North America. In Extinction is Forever: The Status of Threatened and En- dangered Plants of North America. Eds. G.T. Prance & T.S. Elias, pp. 301-313. New York Botanical Garden, Bronx, New York. ______. 1982. The gulf coast pitcher plant bogs. Amer. Sci. 70:260-267. Fritz, E.C. & J. Alford. 1986. Realms of Beauty: The Wilderness Areas of East Tezas. University of Texas Press, Austin. Frost, C.C., J. Walker, & R.K. Peet. 1986. Fire-dependent savannas and prairies of the Southeast. In Wilderness and Natural Areas in the East- ern United States: A Management Challenge. Eds. D.L. Kulhavy & R.W. Conner. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas. Pp. 348-357. Hartsell, D.C. 1980. A preliminary survey of the vascular flora of Lin- coln Parish, Louisiana. M.S. thesis, Louisiana Tech University, Ruston, Louisiana. MacRoberts, B.R. & M.H. MacRoberts. 1988. Floristic composition of two west Louisiana pitcher plant bogs. Phytologia 65:184-190. 1990a. Vascular flora of two west Louisiana pitcher plant bogs. Phytologia 68:271-275. MacRoberts, D.T. 1989. A Documented Checklist and Atlas of the Vascular Flora of Louisiana. Bull. Mus. of Life Sciences 7-9, Louisiana State University, Shreveport. MacRoberts, M.H. & B.R. MacRoberts. 1988. A note on Sarracenia purpurea L. in Louisiana. Phytologia 65:191-194. . 1990. Size distribution and density of trees in bogs and pine wood- lands in west central Louisiana. Phytologia 68:428-434. McDaniel, S. 1971. The genus Sarracenia (Sarraceniaceae). Bull. Tall Tim- bers Research Station, No. 9, 1-36. Murry, R.E. & L.E. Urbatsch. 1979. Preliminary reports on the flora of Louisiana. III. The families Droseraceae and Sarraceniaceae. Castanea 44:24-27. MacRoberts & MacRoberts: Sarracenia in Louisiana 125 Parvin, R.W. 1989. Reclaiming a Big Thicket gem. The Nature Conservancy Magazine 39(3):22-26. Rebertus, A.J. & N.G. Barker. 1984. Louisiana’s fascinating bogs. Louisiana Conservationist 36:18-21. Roland, J.A. 1966. A floristic study of the vascular aquatic plants of Lin- coln Parish, Louisiana. M.S. thesis, Louisiana Tech University, Ruston, Louisiana. Smith, L.M. 1988. The natural communities of Louisiana. Louisiana Natu- ral Heritage Program. Louisiana Department of Wildlife and Fisheries. Unpublished. Phytologia (February 1991) 70(2):126-134. RECENSION OF THE ASPLUNDIANTHUS GROUP OF EUPATORIUM, s.\. B. L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A taxonomic treatment of the Andean genus Asplundianthus King & H. Robins. is rendered. Seven species are recognized in the complex. Brief descriptions, complete synonymy, a key to species, and maps show- ing distribution are presented. KEY WORDS: Asteraceae, Eupatorieae, Eupatorium, Asplundi- anthus. King & Robinson (Monographs Syst. Bot., Missouri Bot. Gard. 22:346- 348. 1987.) recognized ten species in their segregate genus Asplundzanthus, all of these restricted to relatively high elevations (2500-3000 m) of the Andean regions of Colombia, Ecuador, and Peri. The present treatment recognizes seven species, all of these having been previously treated within Eupatorium, s.l., by yet other workers. Two recently described species, Asplundianthus pseudostuebelii King & H. Robins. and A. sagasteguit King & H. Robins., are treated as synonyms of the widespread Eupatorium stuebeliz Hieron. In their treatment of the tribe Eupatorieae, King & Robinson erected the genus Asplundianthus to accommodate a group of montane Andean shrubs or clambering vines with mostly deciduous involucral bracts and slender stylar appendages. Asplundianthus was positioned within the subtribe Critoniinae between Aristeguretia King & H. Robins. and Austrocritonia King & H. Robins. It is distinguished from the former by its narrow style branches and from the latter by its rather evenly barbellate pappus bristles (vs. bristles barbellate below but smooth and tapered apically). In its total characters, however, it appears to stand somewhat closer to Austrocritonia, especially if stylar characters are emphasized. 126 Turner: Recension of Asplundianthus group of Eupatorium 127 KEY TO THE ASPLUNDIANTHUS GROUP OF EUPATORIUM RCAVER ESSE. CANORIINAG sca at's foie ans aan ngs cle Mea ten we E. densum Pe onenves With petioles 2-30 mm, longo? eo... eee ies gas aed (2) 2. Leaves mostly 2-4 cm long; Colombia. ................. E. arcuans o eaves tnostly 4-10 cutlong . 222. aasg. &cehowe-8 eG ateuens (3) 3. Leaves pinnately nervate, glabrous beneath; petioles mostly 2-8 mm long; LRA I yo aR PN sn ccrctic sad le Sie denim Ses GR A Ricca va RR pS E. toroi 3’ Leaves trinervate to subpinnately nervate, variously pubescent to glabrous heneath; petioles mostly 6-30 mimi long. | 2.20.0. 0.8. gece ieee ween nels (4) 4. Heads arranged in closely clustered ovoid corymbs; florets 4-7 per head; leaves subglabrous, pubescent mainly along the veins; Ecua- FIBES, oc end Ckn> Rane wd cdeae ao eeee cas E. pseudoglomeratum 4’ Heads arranged in cymose panicles, if in ovoid corymbs then the branches which bear them widely divaricate; florets 7-10 per head; leaves glabrous to densely pubescent. .. 2)... 5.0.0 0c0ccceanee (5) 5. Leaves glabrous or nearly so; blades trinervate from or near the base; Colombia, northern Ecuador, Venezuela. ............... E. smilacinum 5’ Leaves variously pubescent; blades trinervate from above the base, or else RIDIGMAUELY BELVGRGS . 0's iin cp «de. c neh ene Ohne ed na saa x = «ee eee (6) 6. Leaves scabridulous beneath, harsh to the touch; Peru (Dept. Hua- cc Pe See eee oe ee ee Pee ee Ee E. trachyphyllum 6’ Leaves with appressed or matted hairs beneath, soft to the touch; Colombia, Beundor, Perd. . j...0c.04 0210) SR ee E. stuebeliu Eupatorium arcuans B.L. Robins. Eupatorium arcuans B.L. Robins., Proc. Amer. Acad. Arts 54:237. 1918. Asplundianthus arcuans (B.L. Robins.) King & H. Robins., Phytologia 30:224. 1975. TYPE: COLOMBIA. Andean range near Bogota, 2600 m, 1851-57, J. Triana 1191 (HOLOTYPE: K; Photoholotype: GH!; Isotypes: GH!, fragment NY!; Photoisotypes: LL!, MO!). Eupatorium gongorae Cuatr., Trab. Mus. Madrid Bot. 29:17. 1935. TYPE: COLOMBIA. Vieja: “Quebrada de la Cuatrecasas,” with- out date, Cuatrecasas 2937 (HOLOTYPE: MA; Photoholotype: MO!) . 128 PY TCL GTA volume 70(2):126-134 February 1991 Shrubs with very leafy, arcuate branching, 2-3 m high. Stems densely rusty tomentose to nearly glabrate. Leaves mostly 1.5-3.5 cm long, 1.0-1.5 cm wide; petioles 3-6 mm long; blades ovate, trinervate, sparsely pubescent only along the major veins beneath, the upper surfaces smooth and seemingly somewhat viscid, the margins serrate. Heads 10-50, arranged in rounded terminal cy- mules, the ultimate peduncles 0-2 mm long. Involucres mostly 5-7 mm high, the bracts 16-18, 3-4 seriate, graduate, glabrous. Florets 10-12 per head, the corollas glabrous, ca. 4.5 mm long, the limb ca. 2 mm long. Achenes ca. 2.5 mm long, glabrous or a few hispid hairs near apices, the pappus of 40-50 moderately barbellate white bristles ca. 4.5 mm long, the apices slender. DISTRIBUTION (Fig. 1): Known only from the vicinity of Medellin and Bogota, Colombia, in Paramo vegetation, 2800-3000 m; July-August. REPRESENTATIVE SPECIMENS: COLOMBIA. Antioquia: vicinity of Medellin, 20 Aug 1927, Toro 470(NY). Boyaca: Paramo de la Rusia, Boyaca, 10,000 ft, 12 Jul 1968, Barkley 38C129 (TEX). Cundinamarca: Guadalupe, near Bogota, without date, Bro. Ariste-Joseph A250 (GH); Boqueron, 2850 m, 30 Jan 1925, A. Schultze 118 (GH). N. Grenada, without specific locality or date, Linden 16 (MO, NY). According to label data (Barkley 38C129), the plant is a “tree of 3 meters with lavender flowers.” Eupatorium densum Benth. Eupatorium densum Benth., Pl. Hartw. 200. 1845. Asplundianthus densus (Benth.) King & H. Robins. Phytologia 30:225. 1975. TYPE: COLOM- BIA. Cundinamarca: near Bogota, without date, Hartweg 1105 (HOLO- TYPE: K; Photoholotype: GH!, NY!; Isotype: NY!; Sketch of isotype: GH!; Photoisotype: MO!). Erect smooth shrub to 2 m high. Stems sparsely puberulent to glabrate. Leaves sessile, ovate-lanceolate, mostly 6-10 cm long, 1.5-2.5 mm wide, pin- nately veined, glabrous or nearly so, the margins entire. Heads sessile and numerous in closely packed terminal corymbose panicles. Involucres 4-5 mm high, the bracts ca. 18, 3-4 seriate. Florets 5-8 per head, the corollas glabrous, ca. 3 mm long, the weakly differentiated limb ca. 1.7 mm long. Achenes ca. 2 mm long, glabrous, the pappus of 30-40 weakly barbellate bristles 2-3 mm long, the apices slender. DISTRIBUTION (Fig. 1): Known only from the vicinity of Bogota, Colom- bia at about 2700 m; May-July. SPECIMENS EXAMINED: COLOMBIA. Cundinamarca: Slopes of Salto de Tequendama, 6 Jul 1929, Chardon 649 (GH); “Prov. de Bogota,” 2650 m, May-Jul 1855, Triana 1230 (also number 55), NY, F (fragment. from P). Turner: Recension of Asplundianthus group of Eupatorium et es oe VENEZUELA Fizi: \ ey See ee ae } ———— —- / % = i oa TS x So Beery Ce COLOMBIA *» fails => . t = poner’ 4 i i sy ~ 5 A arcuans Rie tease i 4 densum PERU 2? 0D smilacinum - O stuebelii f @ toroi Se Q@achyphyttum ‘ J ’ = ‘ > s \ i oes ow ne or , ’ a = oad ‘ == = i ae ‘ == = i) ‘ ="= = ' ‘ SSS ~ . — = = = ‘ a. == ES = i] sy = ————— ate AP v] ert. ———) a , “s => SS =! ‘ = — = \ oom — =z ’ a — ee 7 | ll yt a lll es) O (hs < > d a == & = = / a == = —9 ~——= “ SE ie mir Os eee - -—=— = 4 3 Eh eae / re — —— a g = = = ‘ —— — as ‘ —— ——— 2 ——= =! = ie aaa a = Fig. 1. Distribution of Asplundianthus species. aie @ pseudoglomeratum 130 P BY-T.OL'O GIA volume 70(2):126-134 February 1991 Vegetatively this species is superficially similar to the poorly known Fupa- tortum toroz B.L. Robins., but lacks the petiolate, glandular punctate, leaves of the latter. Eupatorium pseudoglomeratum Hieron. ez Sod. Eupatorium pseudoglomeratum Hieron. ez Sod., Bot. Jahrb. Syst. 29:8. 1900. Asplundianthus pseudoglomeratus (Hieron. ex Sod.) King & H. Robins. Phytologia 30:225. 1975. TYPE: ECUADOR. without locality, 1897, Sodiro 6/2 (Lectotype fragment and Photolectotype: {B], GH!). Three collections of Sodiro were cited in the protologue (6/2, 6/4, 6/10); col- lection 6/2 is selected here as the lectotype. Weak stemmed nearly glabrate suffruticose herbs or shrubs to 2 m high. Stems puberulent, glabrate with age. Leaves mostly 4-14 cm long, 2-5 cm wide; petioles 0.8-3.0 cm long; blades ovate deltoid to ovate, trinervate from or near the base, glabrous above, puberulo-hispid beneath, the margins ser- rate. Heads numerous, arranged in clusters of terminal globose corymbs, the ultimate peduncles 0-1 mm long. Involucres 5-6 mm high, the bracts 12-16, 3-5 seriate, glabrous. Florets 4-7 per head, the corollas ca. 3.5 mm long, the weakly differentiated limb ca. 1.5 mm long. Achenes ca. 2 mm long, glabrous, the pappus of 40-50 sparsely barbellate bristles ca. 3 mm long. DISTRIBUTION (Fig. 1): Known only from Colombia (?) and Ecuador, Prov. Pichincha, 2800-3000 m; January-March. REPRESENTATIVE SPECIMENS: ECUADOR. Pichincha: ca. 3 km SW Chillogallo, ca. 10,800 ft, 17 Jan 1974, King 6510 (F); ca. 10 km W of Quito, ca. 10,000 ft, 7 Feb 1974, King 6732 (MO). Yunguillo, 2800-3000 m, 14 Mar 1987, Zak 1830 (F, MO, NY); “about Tambillo, Pifo,” etc., 2000-3000 m, without date, Mille 540 (GH, MO, NY). COLOMBIA (?): Ocana: “in cours des maisons,” 3500 m, Nov 1851, Schliim 331 (F). Eupatorium smilacinum H.B.K. Eupatorium smilacinum H.B.K., Nov. Gen. & Sp. 4:87. 1818. [ed. folio] As- plundtanthus smilacinus (H.B.K.) King & H. Robins. Phytologia 30:225. 1975. TYPE: COLOMBIA. Tolima: Quindio Mountains, near Alto de Guayabal & Quebrada de Toche, 1280-1830 m, without date, Humboldt & Bonpland s.n. (HOLOTYPE: P; Photoholotype: GH!, LL!, MO!). Clambering nearly glabrous shrubs. Stems pubcrulent, glabrate with age. Leaves mostly 5-12 cm long, 2-5 cm wide; petioles 6-30 mm long; blades ovate Turner: Recension of Asplundianthus group of Eupatorium 131 to ovate-elliptic, glabrous above and below, trinervate from or near the base, the margins dentate, sometimes coarsely so. Heads sessile and numerous, arranged in rounded corymbs, these in turn arranged in widely or divaricately branched corymbose panicles. Involucres 4-6 mm high, the bracts ca. 16, graduate, 3-4 seriate, glabrous. Disk florets 6-8 per head, the corollas glabrous, 4-5 mm long, the limb weakly defined, ca. 2 mm long. Achenes ca. 2 mm long, glabrous except for a few hispid hairs near the apex, the pappus of ca. 40-50 white sparsely barbellate bristles 4-5 mm long. DISTRIBUTION (Fig. 1): Venezuela, Colombia, and Ecuador, montane rain forests and shrub zone (paramillo), 2100-3000 m; February-June (August). SPECIMENS EXAMINED: ECUADOR. Napo-Pastaza: Coyuja, ca. 57 km ESE of Quito, ca. 2900 m, 21 May 1947, Fosberg 27542 (MO). COLOMBIA. Cauca: Moscopan, 2600 m, Mar 1943, Kjell von Sneidern 4311 (LL); Moscopan, 2000 m, Aug 1944, Kjell von Sneidern 4684 (LL); Mount El Trueno, 2700-3000 m, 29-30 Jun 1972, Pennell 7519 (GH). Tolima (Quin- dio): Moral en el Quindio, 2100 m, Feb 1854, Triana 1192 (GH). VENEZUELA. Tachira: Slopes along Quebrada Agua Azul, 14 km SE of Las Delicias, 2150-2300 m, 22-23 Jul 1979, Steyermark & Leisner 118279 (MO). This taxon superficially resembles Eupatorium arcuans B.L. Robins., the latter having smaller, pubescent leaves, and capitulescences not divaricately branched. Eupatorium stuebelii Hieron. Eupatorium stuebelii Hieron., Bot. Jahrb. Syst. 21:329. 1895. Asplundi- anthus stuebelu (Hieron.) King & H. Robins. Phytologia 30:226. 1975. TYPE: ECUADOR. “Campamento Utanag, Valle del Rio Chambo,” 3045 m, Nov 1872, A. Stuebel 272 (Lectotype: B, destroyed ?; Photolec- totype: GH!, MO!, NY!). Two collections were cited in the protologue, Stuebel 164 {from Colombia] and Stuebel 272 [from Ecuador]; by annota- tion on the latter, Hieronymous clearly denoted the type intended and this was formalized by King & Robinson in their transfer of the species to Asplundianthus, where they lectotypified E. stuebelii with Stuebel 272). Asplundianthus pseudostuebelii King & H. Robins. Phytologia 30:225. 1975. TYPE: COLOMBIA. Cundinamarca: ca. 15 km NNW Fa- catativa, ca. 2330 m, 14 Jul 1965, R.M. King, et al. 5923 (HOLO- TYPE: US; Isotype: NY!). Asplundianthus sagasteguii King & H. Robins. Phytologia 39:137. 1978. TYPE: PERU. Piura: Canchaque-Minas Turmalina, 2250 m, 23 Jul 1975, Sagastegui, et al. 8273 (HOLOTYPE: US; Isotypes: F!, MO!, NY!). 132 PHY TOL 0O:G:LA volume 70(2):126-134 February 1991 Shrub or clambering vine. Stems densely tawny puberulous or sordid to- mentulose. Leaves mostly 6-15 cm long, 2.5-6.0 cm wide; petioles 1-3 cm long; blades broadly ovate, trinervate from or somewhat above the base, rarely pen- ninervate, the upper surfaces rugose and glabrate to moderately pubescent with appressed hairs, the undersurfaces pubescent with appressed hairs or densely puberulent to pilose, the margins finely crenulate to nearly entire. Heads numerous in broad rounded cymose panicles, the latter 10-25 cm across, 5-15 cm high, the ultimate peduncles mostly 0-3 mm long. Involucres 6-7 mm high, the bracts ca. 16, 3-5 seriate, graduate. Florets ca. 10 per head, the corollas glabrous, 5-6 mm long, the limb poorly differentiated, ca. 2.5 mm long. Achenes 2-3 mm long, sparsely hispidulous, atomiferous glandular, or both, rarely glabrous throughout, the pappus of 40-50 white barbellate bristles ca. 5 mm long, the apices slender. DISTRIBUTION (Fig. 1): Colombia, Ecuador, and northern Peru, 2500- 3200 m; June-September. The name Asplundianthus pseudostuebeliit has been applied to forms from Colombia with longer, more slender-tapering blades than is usually found. Asplundianthus sagasteguii is said to differ from A. stuebeli (Hieron.) King & H. Robins. by the “minute dense almost hyphal blackish tomentum of the leaf undersurfaces.,” but this is not apparent in an isotype (F); indeed, I can find no characters to distinguish material from northernmost Peru from that of Ecuador and Colombia. REPRESENTATIVE SPECIMENS: COLOMBIA. Caldas: Salento to “Laguneta,” Old Quindio Trail, 2500-3100 m, 1 Aug 1922, Kilhp & Hazen 9106, 9140 (GH); “Pinares,” above Salento, 2600-2900 m, 2-10 Aug 1922, Pennell 9207, 9208 (GH, NY). Cundinamarca: sabana de Bogota, hills of Chapinero, Jun 1923, Pring 201 (MO). Cauca: “Calaguala,” Conuco, 2500- 2800 m, 14-18 Jun 1922, Pennell 7180 (GH); “San Jose,” San Antonio, 2100- 2300 m, 1 Jul 1922, Pennell 7650 (GH). Valle: Mpio. Tulua, Corr. Santa Lucia, finca San Luis, 7800 m, 22 Sep 1984, Devia A. 716 (MO). ECUADOR. Canar: vicinity of Azogues, 16-17 Sep 1918, Rose 22787 (GH, NY). Chimborazo: near Pimo, 9 Jul 1945, Camp E-4122 (MO, NY). Pichin- cha: “Andes of Quito,” 9000 ft, 1848, Jameson 617 (fragment, GH); 8 km W of Aloag, 3030 m, 22 Jul 1977, Stuessy & Jansen 4888 (TEX). Tungurahua: Cusatagua, near Ambato, Mar 1919, Pachano 195 (GH). PERU. Amazonas: Chachapoyas, 5 Mar 1901, Mathews s.n. (GH). Lam- bayeque: between Huaratara and Colaya, 2000 m, 5 Jul 1986, Quiroz 1980 (F). Piura: Cuella del Indio (road to Huancubamba), 2800 m, 13 Sep 1981, Lopez M. 8874 (¥, MO). Turner: Recension of Asplundianthus group of Eupatorium 133 Eupatorium toroi B.L. Robins. Eupatorium toroi B.L. Robins., Contr. Gray Herb. 104:28. 1934. Asplundi- anthus toroi(B.L. Robins.) King & H. Robins., Phytologia 30:226, 1975. TYPE: COLOMBIA. Antioquia: Titiribi, ca. 50 km from Medellin, 30 Jun 1928, Rafael A. Toro 1201(HOLOTYPE: NY!; Fragment holotype: GH!). Clambering shrubs. Stems densely puberulent to subglabrate. Leaves ovate elliptic to ovate lanceolate, mostly 5-10 cm long, 1.8-4.0 cm wide, pinnately veined, glabrous throughout, somewhat glandular punctate beneath, the mar- gins entire; petioles 3-10 mm long. Heads sessile and numerous in closely packed corymbose panicles. Involucres 5-6 mm high, the bracts ca. 16, 3-4 seriate. Florets 4-6 per head, the corollas glabrous, ca. 4 mm long, reportedly “pale yellow,” the limb weakly differentiated, ca. 2.2 mm long. Achenes ca. 2 mm long, sparsely hispidulous apically, the pappus of 30-40 bristles, the apices somewhat swollen and barbellate. DISTRIBUTION (Fig. 1): north central Colombia, montane forests, ca. 1900 m; flowering May-June. ADDITIONAL SPECIMEN EXAMINED. COLOMBIA: Antioquia: Mpio. Granada, road between San Carlos-Granada at km 24, 1900 m, 21 May 1988, Zarucchi, et al. 6783 (MO). Eupatorium trachyphyllum Hieron. Eupatorium trachyphyllum Hieron., Bot. Jahrb. Syst. 36:467. 1905. Asplun- dianthus trachyphyllus (Hieron.) King & H. Robins., Phytologia 30:226. 1975. TYPE: PERU. Huanuco: Tambillo, Jelski 697 (HOLOTYPE: B, destroyed?; Fragment type: GH!; Photoholotypes: GH!, TEX!; Isotype MO!). Eupatorium scabrifolum B.L. Robins., Contr. Gray Herb., n.s. 77:36. 1926. Asplundianthus scabrifolius (B.L. Robins.) King & H. Robins., Phytologia 30:226. 1975. TYPE: PERU. Huanuco: Mito, 2745 m, 23 Jul-14 Aug 1922, Macbride & Featherstone 1873 (HOLOTYPE: F; Photoholotypes: GH!, NY!, TEX!; Isotype: GH!). Clambering shrubs to 7 m high. Stems much branched above, densely trusty tomentulose at first but soon glabrate. Leaves 6-9 cm long, 1.5-4.0 cm wide; petioles ca. 1 cm long; blades ovate to narrowly lanceolate, subpinnately nervate, hispidulous to scabridulous, the surfaces reticulately nerved and be- tween these many minute atomiferous glands, the margins entire or nearly so. Heads numerous in cymose panicles, the ultimate peduncles mostly 1-3 mm 134 PHYTOL OGIA volume 70(2):126-134 February 1991 long. Involucres 5-6 mm high, the bracts ca. 16, 3-4 seriate, graduate. Disk florets 5-9 per head, the corollas lilac, ca. 5 mm long, glabrous, the limb ca. 2.5 mm long. Achenes ca. 2.5 mm long, sparsely hispidulous, the pappus of 40-50 white barbellate bristles 4-5 mm long, the apices slender. DISTRIBUTION (Fig. 1): Known only from Department of Huanuco, Peru, 1800-2800 m; May-August. Other than the more linear lanceolate leaves of Eupatortzum scabrifolium B.L. Robins., I can find little else to distinguish this from E. trachyphyllum Hieron., both occurring in the same general region. According to label data on the type of the latter, FE. trachyphyllum is a liana up to 7 m long. ACKNOWLEDGMENTS This study is based upon about 100 specimens from the following herbaria: F, GH, LL, MO, TEX. I am grateful to Guy Nesom and Linda Escobar for reviewing the manuscript Phytologia (February 1991) 70(2):135-141. FLORISTICS OF THREE BOGS IN WESTERN LOUISIANA B.R. MacRoberts & M.H. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. ABSTRACT The floristics, species diversity, and soil characteristics of three west Louisiana bogs are described and compared with other bogs in the re- gion. KEY WORDS: Pitcher plant bog, hillside seepage bog, Louisiana, floristics, Sarracenia, Kisatchie National Forest INTRODUCTION In two previous papers we describe the floristic composition of four west Louisiana pitcher plant bogs (a.k.a., hillside seepage bogs) (MacRoberts & MacRoberts 1988, 1990a). In the present paper we describe the floristic com- position and species diversity of one pitcher plant bog and two bogs that lacked pitcher plants. The reason we undertook this study is twofold. First, almost nothing has been published about bogs west of the Mississippi River delta (see literature in MacRoberts & MacRoberts 1988, 1990a; Nixon & Ward 1986; Bridges & Orzell 1989; Frost, et al. 1986). Second, we thought that there were two types of bogs in our study area: ones that lacked pitcher plants and seemed to be drier, and ones that had pitcher plants and appeared to be wetter. In other words, there appeared to be a soil moisture difference. Also, the drier bogs had extensive stands of Ctenitum aromaticum (Walt.) Wood, a grass absent from the four very wet bogs we had studied previously. SLUDY SITES Frog Arrow, 360A, and 360B bogs are located in the Kisatchie Ranger District of the Kisatchie National Forest, Natchitoches Parish, about 9 km east of Lotus at the headwaters of the Bayou L’Ivrogne drainage, at about 100 m above sea level. The three bogs are within a 0.7 km radius of each other. 135 MacRoberts & MacRoberts: Floristics of Louisiana bogs 136 Table 1. Soil Characteristics. | Exchangeable ions (ppm) __| ions PNAS Ded: | 2 900 | Frog Arrow | 5.0 Only one of them (Frog Arrow) has Sarracenia. All three have Clenzum. Frog Arrow measures 2.4 ha; 360A measures 2.2 ha; 360B measures 0.9 ha. Frog Arrow is a relatively flat bog with a slight slope. 360A and 360B have a greater slope, ca. 3-5 degrees. All three are surrounded by upland longleaf pine forest. 360A and 360B abut riparian woodland at their lower edge. Frog Arrow abuts riparian habitat on one side only. All three are open, with a few scattered pines and shrubs (MacRoberts & MacRoberts 1990b). All occur on Anacoco loam (fine, montmorillonitic, thermic Vertic Albaqualfs) with Malbis soil (fine loamy, siliceous, thermic Plinthic Paleudults) upslope (Martin, al. 1990). Sphagnum is present in all the bogs but is not abundant. The climate is described in our previous papers and in Martin, et al. (1990). All three bogs have been variously damaged by logging. All were prescribed burned in the winter (nongrowing season) of 1989-1990. METHODS We visited the bogs at two week intervals from April through October 1990. Voucher specimens for many of the species were collected. Rare or easily identifiable plants were not collected. We follow MacRoberts (1984, 1989) for scientific nomenclature. Soil samples taken from the upper 15 cm of each bog were analyzed by A & L Agricultural Laboratories, Memphis, Tennessee. In order to determine species diversity we established ten one meter square plots and two twenty-five meter square plots in Frog Arrow and 360A bogs and recorded species in them every two weeks. RESULTS Table 1 gives soil information for the three bogs. Table 2 lists the species found at the bogs. “F” indicates presence at Frog Arrow, “A” indicates presence at 360A, and “B” indicates presence at 360B. Absence of a letter indicates presence at all three bogs. 137 PHY TOLO GLA volume 70(2):135-141 February 1991 Table 2. Taxa present at Frog Arrow, 360A, and 360B bogs. DENNSTAEDTIACEAE - Pteridium aquilinum (L.) Kuhn. LYCOPODIACEAE - Lycopodium alopecuroides L., L. appressum (Chapm.) Lloyd & Underw., L. carolinianum L. OSMUNDACEAE - Osmunda cinnamomea L. (F), O. regalis L. (F, B). PINACEAE - Pinus palustris P. Mill., P. taeda L. AMARYLLIDACEAE - Aypozis rigida Chapm. BURMANNIACEAE - Burmannia capitata (Walt.) Mart. (F, B). CYPERACEAE - Carer glaucescens Ell. (F, B), Dichromena latifolia Baldw. ez Ell. (F), Eleocharis tuberculosa (Michx.) Roem. & Schult. (F, B), Fuzrena squarrosa Michx., Rhynchospora chalarocephala Fern. & Gale (F), R. globularis (Chapm.) Small var. globularis, R. glomerata (L.) Vahl. (F), 2. gracilenta A. Gray (A), R. olgantha A. Gray, R. plumosa Ell., R. pusilla Chapm. ez M.A. Curtis (F, A), R. rariflora (Michx.) Ell., Scleria ciliata Michx. (F), S. georgiana Core (B), S. reticularts Michx. ERIOCAULACEAE - Eriocaulon decangulare L., Lachnocaulon anceps (Walt.) Morong. JUNCACEAE - Juncus scirpoides Lam., J. trigonocarpus Steud. (F). LILIACEAE - Aletris aurea Walt., Smilaz laurifolia L., S. rotundifolia L. (A, B). ORCHIDACEAE - Calopogon tuberosus (L.) B.S.P., Platanthera integra (Nutt.) A. Gray ez Beck, P. nivea (Nutt.) Luer. (A), Pogonia ophioglossoides (L.) Juss. (F), Spiranthes cernua (L.) L.C. Rich. (F, A), S. vernals Engelm. & A. Gray. POACEAE - Andropogon gerardii Vitman, Andropogon virginicus L. (F, A), Anthaenantia rufa (Ell.) Schultes, Aristida virgata Trin., Ctentum aromaticum (Walt.) Wood, Dichanthelium acuminatum (Sw.) Gould & Clark, D. dichotomum (L.) Gould (A), D. scoparium (Lam.) Gould, Eragrostis spectabilis (Pursh) Steud., Muhlenbergia erpansa (Poir.) Trin., Panicum virgatum L., Paspalum floridanum Michx. (F), Paspalum laeve Michx., Setaria geniculata (Lam.) Beauv. (A), Tridens ambiguus (Ell.) Schultes. XYRIDACEAE - Xyris ambigua Beyr. ez Kunth, X. baldwinzana Schultes, X. caroliniana Walt., X. difformis Chapm. var. curtissii (Malme) Kral, X. drummondi Malme, X. torta Smith. ACERACEAE - Acer rubrum L. ANACARDIACEAE - Rhus copallina L. (F, A), Tozicodendron radicans (L.) Kuntze (F), T. verniz (L.) Kuntze (F, A). APIACEAE -— Eryngium integrifolium Walt., Ozypolis rigidior (L.) Raf. (F), Ptilimnium capillaceum (Michx.) Raf. (F, B). MacRoberts & MacRoberts: Floristics of Louisiana bogs 138 Table 2 (continued). AQUIFOLIACEAE - Ilez opaca Ait. (F), J. vomitoria Ait. ASCLEPIADACEAE - Asclepias longifolia Michx., A. obovata Elliott (B).* ASTERACEAE - Aster ericoides L. (A, B), Bigelowia nuttallii Anderson (A), Chaptalia tomentosa Vent., Coreopsis hnifoha Nutt., Eupatorium leucolepis (DC.) Torrey & A. Gray, E. rotundifolum L., Helianthus angustifolius L., Heterotheca graminifolia (Michx.) Shinners (F, A), Liatris pycnostachya Michx., Marshallia tenuifolia Raf., Senecio tomentosa Michx. (A, B), Solidago nitida Torrey & A. Gray (A, B). CAMPANULACEAE - Lobelia reverchonu B.L. Turner. CAPRIFOLIACEAE - Viburnum nudum L. (F, A). CLUSIACEAE - Hypericum fasciculatum Lam., H. setosum L. (F), H. stans (Michx.) Adams & Robson. DROSERACEAE - Drosera brevifolia Pursh, D. capillaris Poir. ERICACEAE - Vaccinium corymbosum L. FABACEAE - Tephrosia onobrychoides Nutt. GENTIANACEAE - Bartonia paniculata (Michx.) Muhl. (A), Sabatia gentianoirdes Ell. HAMAMELIDACEAE - Liguidambar styraciflua L. LAMIACEAE - Scutellaria integrifolia L. LAURACEAE - Persea borbonia (L.) Spreng. LENTIBULARIACEAE - Pinguicula pumila Michx., Utricularia cornuta Michx., U. juncea Vahl, U. subulata L. LINACEAE - Linum medium (Planch.) Britt. LOGANIACEAE - Cynoctonum sessilifolium (Walt.) St. Hil. (A), Gelssmzum sempervirens (L.) St. Hil. (A, B). MAGNOLIACEAE - Magnolia virginiana L. MELASTOMATACEAE - Rhezia lutea Walt., R. mariana L. var. mariana, R. petiolata Walt. MYRICACEAE - Myrica cerifera L. M. heterophylla Raf. NYSSACEAE - Nyssa sylvatica Marsh. ONAGRACEAE - Ludwigia hirtella Raf., L. linearis Walt. (A, B). POLYGALACEAE - Polygala cruciata L., P. incarnata L. (A, B), P. mariana P. Mill. (A), P. nana (Michx.) DC. (A), P. ramosa Ell. ROSACEAE - Rubus loutsianus Berger. RUBIACEAE - Hedyotis uniflora (L.) Lam. (F, B). 139 PHYTOLOGIA volume 70(2):135-141 February 1991 Table 2 (continued). SARRACENIACEAE - Sarracenia alata Wood. (F). SCROPHULARIACEAE - Agahnis purpurea (L.) Penn., Gratiola pilosa Michx. (F, A). VIOLACEAE - Viola primulifolia L. (F, A). * Asclepias viridiflora Raf. reported from Strange Road Bog (MacRoberts & MacRoberts 1988) should be A. obovata Elliott. DISCUSSION The soils of these three bogs are similar and are similar to the soils of the bogs we have studied previously except that they are, in general, a little less acidic. We recorded 123 taxa for the three bogs, representing 80 genera and 43 families. Frog Arrow had 104 taxa; 73 genera, and 42 families. 360A had 101 taxa: 70 genera and 39 families. 360B had 92 taxa: 66 genera and 39 families. The average number of taxa for the three bogs is 99 species, which is comparable to bogs we have studied earlier and to bogs in east Texas (Nixon & Ward 1986). The three bogs had 88% of the 48 species listed by Nixon & Ward (1986) for six east Texas bogs and 85% of the 52 most prevalent species listed by Bridges & Orzell (1989) for bogs in southwestern Louisiana and southeastern Texas. Sorensen’s index of similarity (see Nixon & Ward 1986) shows that these bogs are floristically similar: Frog Arrow/360A (83), Frog Arrow/360B (84), 360A/360B (86). They are less like Middle Branch and Strange Road bogs (index of similarity in mid- to low 60’s) than like Fixit (a.k.a. Bog Bayou L’Ivrogne) and Woodcock (index of similarity in mid- to low 70’s) (MacRoberts & MacRoberts 1988, 1990a). Nonetheless, our original premise — that the three bogs in the present study would be substantially floristically different from the bogs we had studied earlier in this area - was not supported. The bogs differed primarily in the presence or absence of a few conspicuous species. The four one meter square plots in Frog Arrow bog had from 16 to 21 species and the six one meter square plots in 360A bog had from 14 to 22 species (not counting Sphagnum). The average of the ten samples was 18.6 species. The two twenty-five meter square plots had 33 species (Frog Arrow) and 32 species (360A). Allen, et al. (1988) found about 20 species per square meter in bogs in Vernon Parish, Louisiana. Our figures are considerably below those given by Walker & Peet (1983) for pine-wiregrass savannas in North Carolina. They found between 22 and 35 MacRoberts & MacRoberts: Floristics of Louisiana bogs 140 species per square meter and between 43 and 57 species per twenty-five square meters, depending on moisture and fire gradient. ACKNOWLEDGMENTS Thanks are due the staff of the Kisatchie National Forest for their coop- eration during the study and to Ella Edwards, Elray Nixon, Edwin Bridges, and Steve Orzell who helped us in various ways. LITERATURE CITED Allen, C.M., C.H. Stagg, & S.D. Parris 1988. Analysis of the vegetation in pitcher plant bogs in two baygalls at Fort Polk in west central Louisiana. Proc. Louisiana Acad. Sci. 50:1-6 Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the west gulf coastal plain. Natural Areas Journal 9:246-262. Frost, C.C., J. Walker, & R.K. Peet. 1986. Fire-dependent savannas and prairies of the southeast. In: Wilderness and Natural Areas in the East- ern United States: A Management Challenge, eds. D.L. Kulhavy & R.W. Conner. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, TX, pp. 348-357. MacRoberts, B.R. & M.H. MacRoberts. 1988. Floristic composition of two west Louisiana pitcher plant bogs. Phytologia 65:184-190. 1990a. Vascular flora of two west Louisiana pitcher plant bogs. Phytologia 68:271-275. MacRoberts, D.T. 1984. The Vascular Plants of Louisiana. Bull. Museum of Life Sciences, No. 6, LSU-Shreveport, LA. —_____. 1989. A Documented Checklist and Atlas of the Vascular Flora of Louisiana. Bull. Museum of Life Sciences, No. 7-9, LSU-Shreveport, LA. MacRoberts M.H. & B.R. MacRoberts. 1990b. Size distribution and density of trees in bogs and pine woodlands in west central Louisiana. Phytologia 68:428-434, Martin, P.G., et al. 1990. Soil Survey of Natchitoches Parish, Louisiana. U.S. Dept. of Agriculture, Soil Conservation Service. 141 PHYTOLOGIA volume 70(2):135-141 February 1991 Nixon, E.S. & J.R. Ward. 1986. Floristic composition and management of east Texas pitcher plant bogs. In: Wilderness and Natural Areas in the Eastern United States: A Management Challenge, eds. D.L. Kulhavy & R.W. Conner. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, TX, pp. 283-287. Walker, J. & R.K. Peet. 1983. Composition and species diversity of pine- wiregrass savannas of the Green Swamp, North Carolina. Vegetatio 55:163-179. Phytologia (February 1991) 70(2):142-144. TWO NEW SPECIES OF VERBESINA (ASTERACEAE) FROM GUERRERO, MEXICO B. L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT Two new species of Verbesina, V. chilapana and V. pseudovir- gata, are described from Guerrero, México. The former is related to V. oaracana DC., but differs in having larger heads with loose oblance- olate bracts and nearly wingless achenes; the latter is closely related to V. virgata Cav., but differs in having larger glabrous leaves, markedly winged stems, and shorter rays. KEY WORDS: Asteraceae, Verbesina, México Routine identification of Mexican Asteraceae has revealed the following novelties in Verbeszna. Verbesina chilapana B. Turner, sp. nov. Verbesinae oazacanae DC. similis sed capitulis majoribus brac- teis involucro laxis incohaerentibus foliaceisque et acheniis ad ma- turitatem sine alis differt. TYPE: MEXICO. Guerrero: km 69 on the Chilpancingo- Chilapa-Tlapa road, rare in a small pocket of oak-pine forest, only a few plants seen, 2100 m, 7 Nov 1990, Jose L. Panero 2037 (HOLO- TYPE: TEX!; Isotypes: MEXU, TENN). Sparsely branched shrubs 0.5-1.0 m high. Stems densely short pubescent, narrowly winged (ca. 1 mm wide). Leaves alternate, 8-12 cm long, 3-5 cm wide; petioles 3-7 mm long; blades elliptic-ovate, widest at or near the middle, pinnately nervate, hispidulous above, densely soft pilose beneath, the margins serrulate. Heads hemispheric, 5-7 to a stem, the ultimate peduncles mostly 2-4 cm long. Involucres 1.5-2.0 cm across, the bracts subequal, triseriate, loose and foliaceous, obovate, ca. 6 mm long, 3-4 mm wide. Receptacles conical, the chaff lanceolate, stiffly acute, exceeding the florets. Ray florets ca. 32, 142 143 PHYTOLOGIA volume 70(2):142-144 February 1991 neuter, sterile, the ligules yellow, 4-8 mm long. Disk florets numerous, the corollas yellow, ca. 5 mm long, the lobes ca. 1 mm long. Achenes ca. 2.5 mm long, glabrous or nearly so, wingless at maturity, or seemingly so; pappus awns delicate, 1-2 mm long. The species is most closely related to Verbesina oaracana DC. and V. auric- ulata DC., both of which possess relatively large heads and alternate leaves. It differs from both in having more numerous rays, dark colored, loose foliaceous involucral bracts and smaller leaves which are softly pubescent beneath. Verbesina pseudovirgata B. Turner, sp. nov. Verbesinae virgatae Cav. sed foliis majoribus glabris ac sub- tiliter reticulatis in paginis infernis et caulibus valde alatis differt. TYPE: MEXICO. Guerrero: km 127 of the Chilpancingo- Chilapa-Tlapa road, along the road in pine-oak forest, 2080 m, 7 Nov 1990, Jose L. Panero 2044 (HOLOTYPE: TEX!; Isotypes: MEXU, TENN). Weak stemmed shrubs 1-2 m high. Stems pubescent with appressed minute hairs, markedly winged, rarely not. Leaves alternate or rarely opposite, 12-20 cm long, 2-5 cm wide; petioles winged throughout, or seemingly absent and the blade clasping the stem; blades obovate to oblanceolate, glabrous or nearly so, the undersurfaces finely reticulate venose, eglandular, the margins entire or nearly so. Heads numerous, arranged in stiffly erect cymose panicles, the ultimate peduncles ascending, mostly 1-4 cm long. Involucres 4-5 mm high, 2-3 seriate, the bracts subgraduate, appressed, greenish yellow. Receptacle subconical, the pales 4-6 mm long, apiculate but not recurved. Ray florets 5-8, pistillate, fertile, the ligules yellow, 2-4 mm long. Disk florets 40-60, the corollas tubular, sparsely pubescent, 3.5-4.0 mm long, the lobes ca. 0.3 mm long. Achenes ca. 2.5 mm long, 1.5 mm wide, the wings membranous, ca. 0.5 mm wide above, gradually tapering below; pappus awns equal, ca. 2 mm long. ADDITIONAL SPECIMENS EXAMINED: MEXICO. Guerrero: Taxco, 12 Oct 1938, Ruth Abbott 485 (GH); along trail from Taxco to Casahuates, mountains west of and above Taxco, 5800-6300 ft, 6 Nov 1949, E. Moore, Jr. 5536 (CU). The species superficially resembles Verbesina virgata Cav. but is readily distinguished by its large, nearly glabrous, finely reticulate, essentially sessile, entire leaves. It is also distinguished by its conspicuous winged stems and small ray florets. The Abbot collection, cited above, is noteworthy in having on the same sheet, two sprigs of the plant, one with opposite and the other with alternate leaves; otherwise, the two shoots are very similar. Turner: New species of Verbesina in México 144 ACKNOWLEDGMENTS I am grateful to Jose Panero for calling these to my attention, to Guy Nesom for the Latin diagnoses, and to him and Linda Escobar for reviewing the manuscript. Information for Authors Articles from botanical systematics and ecology, including biographical sketches, critical reviews and summaries of literature will be considered for publication in PHYTOLOGIA. Manuscripts may be submitted either on computer diskette, or as typescript. Diskettes will be returned to authors after action has been taken on the manuscript. Diskettes may be 5.25 inches or 3.5 inches but must be written in DOS format as flat ASCII files. 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