ea Vide tee Be asitedeint PUR er Pere we anh ta oh cbt iA Ade haa 2 Ala Wella ah a feath 3 Ly Ae seaboadag Vit Malo issih wy, ie rane ‘ Agta et AED Lagat spin gt : : fa tat eA ORE LUM tAe vy ntae et ¥, ¢ Pea a Hee ah ee byt ie ene Cer cat, aint fatacus epee yswca tat Waa ye! otal ta dey P56 ~! : ST PHYTOLOGIA An international journal to expedite plant systematic, phytogeographical and ecological publication OCT 07 2004 Vol. 86 August 2004 No. 2 R. P. ADAMS, Juniperus deltoides, a new species, and nomenclatural notes on J. polycarpos and J. turcomanica (Cupressaceae) ..........sccscccssssssecssssessecessesesseee 47 B. S. CROZIER, Subfamilies of Cactaceae Juss. including Blossfeldioideae PpRMnmpEE tes ole. FES Co Sh Vc maw Mt TON NN ude ge 52 D. B. WARD and D. W. HALL, Keys to the flora of Florida -- 10, Galactia i PRR MRNI SE os) ee A Am a 65 B. L. TURNER, Two new species of Wedelia (Asteraceae: Heliantheae), with Beouspectus of the genus for MEXICO -.........c.c.sesccbesesdsceseeleccssoceccestcseetll o.....:. 75 M. H. MACROBERTS, B. R. MACROBERTS and L. S. JACKSON, Observations on Parnassia grandifolia DC. (Saxifragaceae) in the west Gulf LED 4 ESTs 2 is ae pe ER ei PCa a MMR A io acne aL On Neten E ORai 98 E. L. KEITH, Tagetes erecta (Asteraceae), a new escape to the flora of EL nt IE a pane eae 3b IAP elle NSM AANGR ee COMMA 104 J. R. SINGHURST and W. C. HOLMES, Prenanthes barbata (Asteraceae) and Parentucellia viscosa (Scrophulariaceae) new to Oklahoma........................... 107 ). J. ROSEN and B. J. CHRISTOFFERSEN, Rediscovery of Cyperus ipeaimpiius (Cypctagedc) 1 TEXAS oo ee 110 _ .L. TURNER, A new combination in Texas Croton (Euphorbiaceae)........ 113 | ee ea IMERLOLTNAOT 3/5005 iL) 5) coat ae aN mech ke ory, 114 ARENA ALSE 2 25)08 2 F351) Fh I one eT 8 114 _PHYTOLOGIA (ISSN 00319430) Phytologia, a journal of plant systematics, phytogeography and vegetation ecology, is published three times a year by the Warner Herbarium, a research unit of the Department of Biological Sciences, Sam Houston State University. Editor Justin K. Williams e-mail: bio_jkw@shsu.edu PHONE: (936) 294-1552 FAX (936) 294-3940 Subscription Phytologia Information & P.O. Box 2012 Scholarly Exchange Sam Houston State University Huntsville, TX 77341-2012 e-mail: bio_jkw@shsu.edu Copyright 2004 by the Warner Herbarium, Sam Houston State University. Printed by the Sam Houston Press. Phytologia (Aug 2004) 86(2) 47 JUNIPERUS DELTOIDES, A NEW SPECIES, AND NOMENCLATURAL NOTES ON J. POLYCARPOS AND J. TURCOMANICA (CUPRESSACEAE) Robert P. Adams Department of Biology, Baylor University, Waco, TX 76798-7388, U.S.A. ABSTRACT A new species from the Mediterranean, Juniperus deltoides R. P. Adams is recognized based on published data on DNA (RAPDS, nrDNA) sequence, morphology and terpenoids. Juniperus turcomanica is recognized as a variety of J. polycarpos, based on RAPD data, terpenoids and morphology. KEY WORDS: Juniperus, Cupressaceae, nomenclature. Adams et al. (2003) examined the relationships among Juniperus drupacea Labill., J. macrocarpa Sibth. & Sm., J. navicularis Grand., J. oxycedrus L. (Greece and Spain), J. oxycedrus var. badia H. Gay comparing nrDNA (ITS) sequences, Random Amplified Polymorphic DNAs (RAPDs), Inter-Sequence Simple Repeats (ISSR) and terpenoids. Although the purpose of that work was to evaluate different kinds of data, we found that the two populations of J. oxycedrus (Greece, Spain) were as different in the these four data sets as the other recognized species in the study. Comparing the specimens of J. oxycedrus (Spain) with the type for J. oxycedrus (LINN!) revealed that they are the same. Thus, the plants from Greece are appropriately recognized as a new species: Juniperus deltoides R. P. Adams sp. nov. TYPE: GREECE. 14 km e. of Archova, 420 m, Lat. 38° 26.720’ N; Long. 22° 41.678’ E, 22 May 2001, R. P Adams 9436 (HOLOTYPE: BAYLU; ISOTYPES: K, NY) 48 Phytologia (Aug 2004) 86(2) Plantas dioicas; frutices vel arbores usque 12 m, saepe coronis pyramidalis. Folia aciculares 9-17 mm longae 1.5-2.4 mm_ latae, base folio fere latiora quam lamina. Folia vittis glaucis duabus in superficiebus adaxialis plerumque non impressis. Strobilus in anno secundo maturescens, globosus_ viridis stramineo-brunneolescens atrorubens in maturitate. Semina plerumque tres. Plants dioecous; shrubs or trees to 12 m, often with pyramidal crowns. Leaves acicular, 9--17 mm long, 1.5--2.4 mm wide, base of the leaf nearly as wide the blade, having two glaucous bands on the adaxial surface, generally not sunken. Cone ripening in second year, globose, dark red when ripe, green, turning to brownish yellow when ripening. Seeds usually 3. Juniperus oxycedrus (sensu stricto) can be distinguished from J. deltoides by having a narrowing of the leaf base (Fig. 5 1, a), whereas the leaf base is almost as wide as the blade (Fig. 1, b) in /. de/toides (hence the name). In addition, the stomatal bands in J. oxycedrus are sunken (Fig. | a), giving the midrib a raised appearance. In SZ SZ contrast, in J. deltoides, most of the leaves a mm | DB have stomatal bands that are not sunken, giving the leaf a flattened surface (actually concave as illustrated in Fig. 1 b). Fig. 1. Leaves J. oxycedrus (a) and J. deltoides (b). Representative Specimens: BULGARIA. s Varna, 7 Dec 1923, B. Gilliat-Smith 373 (K). CYPRUS. Troodos, Prodhromos, 19 Jun 1939, H. Lindberg s.n. (K). above Prodhromos, 15 May 1957, L. F H. Merton 3179 (K). GREECE. Crete, Mt. Spathi, 1760m, 16 May 1967, Barclay 257 (K). 14 km e Arachova, 420 m, 22 May 2001, Adams 9436, 9437, 9438 Phytologia (Aug 2004) 86(2) 49 (BAYLU). 8 km w Lemos, 1010 m, 27 Aug 1999, R. P. Adams, 8787, 8788 (BAYLU). Mt. Parnassus, 1280 m, 3 Aug 1987, R. P Adams 5642, 5643 (BAYLU). 10 km w Crysoritsi, 1200 m, 5 Aug 1987, Adams 5649, 5650 (BAYLU). s side Mt. Parnis, 1070 m, 8 Aug 1987, Adams 5656, 5657 (BAYLU). 7 km w Lemos, 1100 m, 5 Oct 1988, Adams 5988, 5989 (BAYLU). Lake Prespa, 850 m, 3 Jul 1932, Alston & Sandwith 1107 (K). Pyrgos, Athos Peninsula, 1938, Loch 15 (K). 2 km from Mazia, 760 m, 15 Nov 1973, Brickell & Mathew 8133 (K). IRAQ. Sarsank, 16 Jun 1970, S. Omar 37692 (K). 20 km ne Zarkko, 550 m, 4 Aug 1957, Ali Rawa 23166 (K). Zawitah, 2750 m, 30 Aug 1933, Rustam 4818 (K). Bekme Gorge, w end, 440 m, 13 Apr 1955, H. Helbaek 764 (K). near Tabriz, 1927, B. Gilliat-Smith 2117 (K). ITALY. 3 km w Raiano, 520 m, 26 May 2001, R. P Adams 9445, 9446, 9447 (BAYLU). MACEDONIA. near Zelenikovo, 23 Dec 1935, Mrs. Ilic s.n. (K). SLOVENIA. between Parenzo & Rovigno, 16 Aug 1922, W. B. Turrill 1009 (K). sw Trebnje, 18 Apr 1935, Jackson & Turrill 86 (K). TURKEY. 30 km n Eskieshir, 1064 m, 20 May 2001, R.P Adams 9430, 9431, 9432 (BAYLU). 8 km from Belen towards Antakya, 600 m, 6 May 1965, M J. E. Coode & B. M. G. Jones 524 (K). Macka, 300 m, 2 Apr 1960, Stainton 8152 (K). Elmali, 1800 m, 19 Apr 1936, T. A. Tengwall, 400 (K). Artvin, 700 m, 27 Jun 1957, Davis & Hedge 30095 (K). above Ankara, 400 m, 5 Jul 1974, P H. Davis 13062 (K). UKRAINE. Yalta, near Nikita, 150 m, 29 May 1959, Davis 33086 (K). Juniperus polycarpos K. Koch from central Asia is a very polymorphic species that is often included as a variety of J. excelsa M.- Bieb. (Farjon, 2001). Adams (2001) examined populations referable to J. polycarpos sensu stricto (Armenia), J. seravschanica Kom. (Kazakhstan, Pakistan), and J. turcomanica B. Fedtsch. (Turkmenistan). Both terpenoids and RAPDs showed J. excelsa to be quite resolved from the J. polycarpos complex (three aforementioned taxa). The leaf terpenoids showed that J. turcomanica to be somewhat different from J. polycarpos of Armenia and J. seravschanica from Kazakhstan and Pakistan. The RAPDs data suggest the recognition of J. polycarpos K. 50 Phytologia (Aug 2004) 86(2) Koch. var. seravschanica (Kom.) Kitamua and J. turcomanica as valid infraspecific taxa. Re-examination of the morphology of J. polycarpos and J. turcomanica suggests to the author that it would be prudent to recognize J. turcomanica as a variety of J. polycarpos. Juniperus polycarpos K. Koch var. turcomanica (B. Fedtsch.) R. P. Adams, comb. nov. BASIONYM: Juniperus turcomanica B. Fedtsch. in Fedtschenko & al., Fl. Turkmenii 1: 14. 1932. TYPE: lost or destroyed (Imkhaniskaya, 1990). (LECTOTYPE: D. P. Gedevanov & D. A. Dranitsyn 148, 3 v 1912, Turkmenia, Kopet Dag, Dschalilu, (chosen by Imkhaniskaya [1990] LE!) Distribution: Elburz and Kopet Mts. of Iran and Turkmenistan. The following key may be used to distinguish the recognized varieties of J. polycarpos: 1. Foliage slender, 0.7- 0.8 mm in cross section (on ultimate branchlets), seed cones 7-9(-10) mm long; scale leaves tightly appressed, giving a smooth branchlet, (1-)2-3(-4) seeds/cone.................ee0000: var. turcomanica 1. Foliage stout, 0.9-1.0 mm in cross section (on ultimate branchlets), seed cones 9-11 mm long or more; scale leaves with a beak or keel so branchlet appears as a string of beads, 3-6 seeds/cone 2. Seed cones 9-11 mm long, at least some scale leaves with very narrow, elongated, brown glands, MOP MU PCUIREM <5: cccss cess secede Sesser ee var. polycarpos 2. Seed cones 8-10 mm long, scale leaves with clear, ellipsoid glands, often ruptured, with a clear exudates: 2222560052. fsue aos noses te peeeeee eee ee var. seravschanica Phytologia (Aug 2004) 86(2) 51 ACKNOWLEDGEMENTS Thanks to Paul Fryxell for assistance in providing the Latin description of J. deltoides. LITERATURE CITED Adams, R. P. 2001. Geographic variation in leaf essential oils and RAPDs of Juniperus polycarpos K. Koch in central Asia. Biochem. Syst. Ecol. 29: 609 - 619. Adams, R. P., A. E. Schwarzbach and R. N. Pandey. 2003. The concordance of terpenoid, ISSR and RAPD markers, and ITS sequence data sets among genotypes: an example from Juniperus. Biochem. Syst. Ecol. 31: 375-387. Farjon, A. 2001. World checklist and bibliography of conifers. 2nd ed. Royal Botanical Gardens, Kew, London. Imkhanitskaya, N. N. 1990. Taksonomicheskaya zametka 0 Juniperus excelsa (Cupressaceae). Bot Zurn. 75(3): 402-409. 52. Phytologia (Aug 2004) 86(2) SUBFAMILIES OF CACTACEAE JUSS., INCLUDING BLOSSFELDIOIDEAE SUBFAM. NOV. Bonnie S. Crozier Section of Integrative Biology, School of Biological Sciences University of Texas, Austin, Texas 78713, U.S.A. ABSTRACT New data from chloroplast DNA comparisons reveal Blossfeldia Werd. (Cactaceae) as an isolated lineage between a basal grade of subfamilies Opuntioideae-Pereskioideae-Maihuenioideae and a more derived sister pair of clades, making recognition of a new subfamily warranted. Based on parsimony and Bayesian cladistic analyses of chloroplast DNA sequences reported elsewhere, as well as morphological distinctiveness, Blossfeldioideae Crozier subfam. nov. is erected and Rhipsalidoideae Burnett is resurrected. An abbreviated history of suprageneric classification in the family and a key to the six subfamilies of Cactaceae recognized here is provided. KEY WORDS: Blossfeldia, Blossfeldioideae, Maihuenioideae, Opuntioideae, Pereskioideae, Rhipsalidoideae, Cactaceae. As in many other families of flowering plants, there has been little unanimity in the suprageneric classification of the Cactaceae Juss. but slowly the discovery of new taxa, careful morphological observation, and other contributions to phylogenetic knowledge have led to refinements. The Cactaceae (Cacti) of Jussieu (1789) encompassed all the known cacti of the time under the single genus Cactus L., but also included Ribes L. (Grossulariaceae). Apparently aware of studies by de Candolle (1828) and Lindley (1830) that excluded Grossulariaceae from Cactaceae, Eaton (1836) nonetheless chose to divide Jusssieu’s Cactaceae into two subfamilies, distinguishing Cactoideae (Cacteae) from Grossularieae. The Grossulariaceae were not included in Burnett’s (1835) concept Phytologia (Aug 2004) 86(2) 53 of Cactaceae (Nopalaceae) divided into subfamilies Rhipsalidoideae (Rhipsalidae) including only the genus Rhipsalis Gaertner, and Opuntioideae (Opuntidae) including the genera Mammillaria Haw., Melocactus Link & Otto, Echinocactus Link & Otto, Cereus Mill., Opuntia Mill., and Pereskia Mill. Engelmann’s (1876) creation of subfamily Pereskioideae (Peireskieae) and division of the family into three subfamilies for the Botany of California begins the modern era in cactus classification. Treating only a few genera Engelmann heralded the classification in three subfamilies years ahead of Schumann (1890,1898) whose Cactoideae (Cereoideae), Pereskioideae (Peireskioideae) and Opuntioideae have been included in most subsequent taxonomic studies (Berger 1926,1929; Backeberg 1958,1966; Buxbaum 1958, Barthlott and Hunt 1993: but see also Britton and Rose 1919-1923; Hunt 1967; Benson 1982 for recognition of these same groups at tribal rank). Cactoideae Eaton, Opuntioideae Burnett, and Pereskioideae Engelm. appear to be validly published, and under the International Code of Botanical Nomenclature (Greuter et al. 2000) these names take priority over the subfamilial names authored by Karl Schumann. The Opuntioideae and Pereskioideae have been clearly circumscribed and almost uniformly recognized in modern times, discounting the uncertain placement of Maihuenia Schum., a genus of only two species from the southern Andes and Patagonia. First associated with the caespitose opuntias, Maihuenia was soon reassigned to the Pereskioideae by Schumann (1898) based on spine, flower, and seed characters. Gibson’s (1977) interpretation of stem tissues and pollen features supported this placement, however Bailey (1968) excluded Maihuenia from the Pereskioideae based on stem and vascular anatomy, and was also unwilling to place it with Opuntioideae based on similar terete leaves. Bailey noted similarities of pollen and highly modified wood that to him suggested a possible relationship with the Cactoideae (Cereoideae). Later, the genus was raised to subfamilial rank by Fearn (1996) who may have been spurred by provisional molecular evidence (see Leuenberger 1997 p.58 and references within, Doweld 1999 p.25). With the exclusion of Maihuenia the monophyly of the Pereskioideae, including only Pereskia Mill., has never been questioned on morphological grounds. 54 Phytologia (Aug 2004) 86(2) i100 Caotoideae Phipsalidcideae Blo ssteldicide at Maihutnicids at Opuntioide a Pereskicide at Portulaca oeat Figure 1. Relationships of Cactaceae subfamilies. Congruent topologies resulted from maximum parsimony and Bayesian phylogenetic analyses of 16,620 base pair chloroplast data set comparing 98 cactus species and 3 outgroup taxa (simplified from Crozier et al. 2004 in prep.). Bootstrap values shown above branches; Bayesian probabilities shown below. For one of the 52 most parsimonious trees the Consistency Index excluding uninformative characters = 0.48, Retention Index = 0.78. However, the molecular study of Nyfeller(2002) was unableto support the monophyly of Pereskia. A broader sample of six Pereskia species in the study by Crozier et al. (2004 in prep.) does form a monophyletic clade with moderate bootstrap support and significant Bayesian probability (see Fig. 1). Together the Opuntioideae, Pereskioideae and Maihuenioideae represent less than 15% of the species of the family. The rest of the family, a morphologically diverse group of more than 1100 species (Hunt 1999), has traditionally been lumped into the single subfamily Cactoideae based on the absence of synapomorphies defining the Pereskioideae and Opuntioideae. This diversity has usually been subdivided into 8 or 9 tribes (see Barthlott 1988 for a review; Barthlott and Hunt 1993). However, comparative analyses of chloroplast DNA Phytologia (Aug 2004) 86(2) 55 sequence data now provide statistically well-supported evidence of two distinct major lineages. The name Rhipsalidoideae Burnett can be used to recognize the clade containing most columnar cacti, epiphytes, and globular cacti of South America (see Fig. 1). Molecular studies are rapidly increasing our recognition of monophyletic lineages inthe Cactaceae facilitating improved classification that reflects evolutionary relationships. Results of parsimony analysis of more than 6,000 base pairs of chloroplast DNA sampled from 120 representative taxa across the family first revealed Blossfeldia as a monophyletic lineage sister to the Cactoideae-Rhipsalidoideae clade with strong bootstrap support (Crozier and Jansen, 2001). Nyfeller (2002) independently inferred the same position of Blossfeldia rejecting the possibility that this might be a taxon-sampling artifact (long-branch attraction) in parsimony and maximum likelihood combined analysis of tmnK-matK and trnL-trnF data. Crozier et al. (2004 in prep.) compared 98 species of cacti, and outgroups from the Portulacaceae and Didieraceae for 16,620 base pairs of chloroplast data using parsimony analysis that yielded strong bootstrap support for the Blossfeldioideae as well (see Fig. 1.). Furthermore, statistical support for this relationship of Blossfeldia was 100% probable in a Bayesian analysis run for 4 million generations of that combined data set representing 13 functional regions of the chloroplast, including genes, introns and intergenic spacer regions. In addition, Blossfeldia shares with the Pereskioideae, Opuntioideae and Maihuenioideae unique chloroplast DNA motifs in multiple markers (Crozier et al., 2004 in prep.). For this reason and its distinctive morphology I am proposing to place Blossfeldia in its own subfamily. Blossfeldioideae Crozier, subfam. nov. Type: Blossfeldia Werd., Kakteenkunde 11:162 (1937). Monotypic (1 species). Type species: B. liliputana Werd. Caudex crassus, caulis simplex dein proliferans, depresso-globosis vel disciformibus 2.5 cm diametro vel parvioribus, neque costatis neque tuberculatis vertice depressioribus lanoso, sine hypodermata epidermata una tabulato sine epicuticulo ceracea, parietibus cellularum 56 Phytologia (Aug 2004) 86(2) epidermis externus vix incrassatis, stomata perpauci 1 per mm’, stomata fovea areolari restrictis. Semina parva globosa strophiolo fere quam grandiore quam semina gerentibus. Perennial herb from a fleshy taproot, succulent, poikylohydric, body swelling immediately after rainfall. Stem solitary or caespitose, individual stems obpyriform when hydrated or flattened, disc-shaped with conspicuous central cuplike depression when dessicated, lacking ribs or pronounced tubercles, 1-1.5 (2.5) cm in diameter. Stomata restricted to areolar pits, overall density much less than 1 per mm’. Pericarpel sculptured with podaria tipped by small lanceolate to triangular scales, or with only a few scales and essentially glabrous on the lower part, bearing whitish to gray wooly hairs in the axils.. Pollen subspherical, tricolpate, with smooth exine. Fruit a juicy berry, spherical to ovoid or pyriform, about 0.5 cm across, with podaria bearing large scales, and axillary hair in small bundles, without bristles, side splitting when ripe then disintegrating over time to release the seeds. Seeds globose, small, 0.5 mm in diameter, testa minutely papillose, shiny red-brown, with large ivory hilum. Blossfeldieae Crozier, trib. nov. Type: Blossfeldia Werd., Kakteenkunde 11:162 (1937). Monotypic (1 species). Type species: B. liliputana Werd. Caudex crassus, caulis simplex dein proliferans, depresso-globosis vel disciformibus 2.5 cm diametro vel parvioribus, neque costatis neque tuberculatis vertice depressioribus lanoso, sine hypodermata epidermata una tabulato sine epicuticulo ceracea, parietibus cellularum epidermis externus vix incrassatis, stomata perpauci 1 per mm’, stomata fovea areolari restrictis. Plantae aphyllae, areolis inermibus coactis. Flores solitarii axillaribus turbinatis vel infundibuliformibus 0.6-1.5 cm longis hermaphroditis. Fructis globosis vel pyriformibus bruneis dehiscentibus dein fatiscentibus. Semina parva globosa strophiolo fere quam grandiore quam semina gerentibus. Phytologia (Aug 2004) 86(2) 57 Perennial herb from a fleshy taproot, succulent, poikylohydric, body swelling immediately after rainfall. Stem solitary, caespitose with age dividing dichotomously, laterally, or sprouting from rootstock exposed to light, individual stems obpyriform when hydrated or flattened disc shaped with conspicuous central cuplike depression when dessicated, lacking ribs or pronounced tubercles, 1-1.5 (2.5) cm in diameter, grey- green to dark green depending on environmental conditions. Stems with a single epidermal layer, external cell walls barely thickened, lacking epicuticular wax coating. Stomata restricted to areolar pits, overall density much less than 1 per mm’. Areoles spirally arranged, circular in face view, tomentose, unarmed. Flowers solitary, subapical arising from the depressed felted crown, usually hermaphroditic, sometimes dichogamous or even unisexual (trimonoecism), opening only in full sun. Flowers turbinate to funnelform, 0.6-1.5 cm long, sometimes with nectary glands. Pericarpel sculptured with podaria tipped by small lanceolate to triangular scales, or with only a few scales and essentially glabrous on the lower part, bearing whitish to gray wooly hairs in the axils. Perianth segments sequentially intergrading in form and color, reflexed. Outermost tepals acuminate, olive-brown. Inner tepals narrowly ovate, rounded apically, white to pale yellowish-white. Androecium in more than two series, equivalent in length. Stamen sometimes wanting, filaments whitish, anthers yellow to gold-yellow. Pollen subspherical, tricolpate, with smooth exine. Stigma and style sometimes wanting, whitish. Stigmatic branches lanceolate, tapering distally, papillose only on the inner surfaces and margins. Fruit a juicy berry, spherical to ovoid or pyriform, about 0.5 cm, wall irregularly sculptured with podaria bearing large scales, and axillary hair in small bundles, without bristles, brown, side splitting when ripe then disintegrating over time to release the seed. Seeds globose, small, 0.5 mm in diameter, testa minutely papillose, shiny red-brown, with large ivory hilum. Chromosome number: n = 33 (Ross, 1981). Blossfeldia \acks xeromorphic stem features of other globular cacti (Barthlott and Porembski 1996) and its globose ornamented and arillate seed is distinctive in the family. Equally distinctive is the restriction of stomata to areolar crypts and extremely low density of stomata on the 58 Phytologia (Aug 2004) 86(2) stem. Based on well supported molecular analyses this smallest member of the Cactaceae represents an isolated lineage, and appears to be the only extant transitional form between the basal grade of subfamilies Pereskioideae-Opuntioideae-Maihuenioideae and a strongly supported clade of more derived cacti (Fig. 1.). At present no other member of the Blossfeldioideae has been identified. Whereas distinct morphologies separate subfamilies Opuntioidae, Pereskioideae, Maihuenioideae and Blossfeldioideae, the remainder of species are so morphologically diverse that phyletic subdivision of the group has been difficult because of parallel and convergent evolution in vegetative and floral morphology (Buxbaum 1958; Barthlott and Hunt 1993). Classifications have been confusing and unstable, and circumscription of suprageneric taxa continues to be modified to try to meet modern expectations of monophyly (Buxbaum 1974, Gibson and Nobel 1986; I.0.S. 1986,1990; Barthlott 1988, Barthlott and Hunt 1993). In light of recent molecular studies a review of the entire suprageneric classification of the family, at least at the subfamilial level, seems in order. The two lineages comprising the clade sister to Blossfeldioideae are quite distinct (see Fig. 1.) and well supported by high bootstrap values and Bayesian probabilities based on the chloroplast DNA studies of Crozier et al. (2004 in prep.). These results show that the two groups are much more distantly related than are the groups of genera within each of them. It is appropriate to recognize these sister clades at equal rank. In so doing the information content of the classification is increased, and the adoption of six subfamilies is not so numerous as to negate its usefulness. Therefore, I am proposing that the proper application of the autonym Cactoideae belongs to the clade of North American globular cacti that includes Mammillaria mammillaris Haw., the conserved type species of the family. This clade corresponds to tribe Cacteae sensu Barthlott and Hunt (1993), though Backeberg (1966) may have been the first to recognize this monophyletic lineage with his subtribe Boreocactinae, a nomen nudum. The morphologically isolated position of this group from other tribes was pointed out specifically by Barthlott (1988) who Phytologia (Aug 2004) 86(2) 59 noted “Zu allen tibrigen Triben kénnen keine Beziehungen erkannt werden.” The earliest valid subfamilial name applicable to the sister clade of columnar, epiphytic and South American globular cacti appears to be Rhipsalidoideae Burnett. The Rhipsalidoideae as recognized here includes all members of tribes Rhipsalideae DC., Echinocereae (Br. & Rose) Buxb., Hylocereae (Br. & Rose) Buxb., Cereae Salm-Dyck, Pachycereae Buxb., Trichocereae Buxb., Browningieae Buxb. and, with the exclusion of Blossfeldia, Notocacteae Buxb. The Cactaceae has been notorious for parallel evolution in morphology that thwarts phylogenetic classification. Parallel reduction in shoot, leaf, flower and seed development in multiple phylogenetic lineages was described by Buxbaum (1951, 1956 and 1965) following phylogenetic ideas of Berger (1926). Although abundant molecular synapomorphies distinguish the Cactoideae and Rhipsalidoideae, unique morphological synapomorphies uniting each clade are difficult to identify. A key to the six subfamilies recognized on the basis of morphological discontinuities and DNA evidence is provided below. KEY TO THE SUBFAMILIES OF CACTACEAE la. Areoles bearing glochids, seeds large, alveolate; bonyzarileoverimne, the entire’Seed>......enceccseeeeaeces ee Opuntioideae 1b. Areoles without glochids, seeds small, usually exarillate or only the hilum covered by a strophiole or corky strophiolar pad. 2a. Plants with persistent photosynthetic leaves on stems. 3a. Plants tree-like or shrubs with laminar leaves...Pereskioideae 3b. Plants low caespitose shrubs with terete leaves.................55 EE OO ce iets SPH AE a EEE EEA RE oe Maihuenioideae 2b. Plants without persistent photosynthetic leaves on stems. 60 Phytologia (Aug 2004) 86(2) 4a. Stems virtually lacking stomata except in sunken crypts; stems lacking thickened cuticle with epidermis lacking thickened outer cell walls, lacking thickened hypodermal layer, stem always flattened globular or disciform less than 25mm diameter; round seeds with strophiole nearly equal in size to the rest of the See 0 Le ci Re are AP PORN Wi oo fe MOE Ge Ss Blossfeldioideae 4b. Stems with stomata or if few then not restricted to cylindrical crypts, usually with thickened cuticle, an epidermal layer with outer cell walls thickened and a hypodermal layer; stems variously flat, triangular globular or columnar; seeds not round, seeds usually exarillate (except in Parodia, Strombocactus, and Aztekium). 5a. Flowers with naked pericarpels, bearing neither areoles nor scales. 6a. Plants globular or short cylindrical, never tall columnar, never with a cephalium, never epiphytic, restricted to North America or the Caribbean; seeds usually with disjunct hilum and micropyle ........Cactoideae 6b. Plants with flat (Schlumbergera), triangular, columnar (Pachycereus, Pilosocereus, Espostoopsis) stems or epiphytic (Rhipsalis, Hatiora, Lepismium), if globular then bearing a cephalium (Melocactus) or restricted to South America (Uebelmannia, some species of Matucana), seeds with conjunct hilum-micropylar region 22). 625-2 seseeeee Rhipsalidoideae 5b. Flowers with pericarpels bearing scales and/or areoles. 7a. Areoles on pericarpel felted, or with hairs, bristles or SpinéSs....e2e0 eee eee Rhipsalidoideae Phytologia (Aug 2004) 86(2) 61 7b. Areoles on pericarpel naked. 8a. Stems columnar or epiphytic, seeds always with conjunct hilum-micropylar region, Amencanion@ anlbbeatsceuseasc-teeeeeaeees Rhipsalidoideae 8b. Stems globular or short cylindrical (Astrophytum, Echinocactus, Sclerocactus papyracantha) or barrel shaped (Ferocactus, Echinocactus), never epiphytic, seeds usually with disjunct hilum and micropyle, restricted toyNorth-AmeriCaseensd senesced Cactoideae ACKNOWLEDGMENTS Sincere thanks are extended to James P. Reveal for pointing out early valid names of subfamilies, to Paul Fryxell for correcting the Latin descriptions, and to Miles Anderson for supplying living material of Blossfeldia liliputana used in molecular studies. Many thanks go to Jose L. Panero for helpful discussions and improvements to this contribution, and to Jim Henrickson for helpful comments in review. LITERATURE CITED Backeberg, C. 1958. Die Cactaceae Vol.1. Gustav Fischer, Jena. 638p. Backeberg, C. 1966. Cactus Lexicon. Blandford Press, Dorset. 828p. Baily, I.W. 1968. Comparative anatomy of the leaf-bearing Cactaceae XVII. Preliminary observations on the problem of transitions from broad to terete leaves. J. Arnold Arbor. 49: 370-376. Barthlott, W. 1988. Uber die systematischen Gliederungen der Cactaceae. Beitr. Biol. Pflanz. 63:17-40. 62 Phytologia (Aug 2004) 86(2) Barthlott, W. and Hunt, D.R. 1993. Cactaceae. Pp 161-196 in The Families and Genera of Vascular Plants II, Flowering Plants, Dicotyledons. K. Kubitzki, J.G. Rohwer, and V. Bittrich eds. Springer-Verlag, London. Barthlott, W. and Porembski, S. 1996. Ecology and morphology of Blossfeldia liliputana (Cactaceae): a poikylohydric and almost astomate succulent. Bot. Acta. 109: 161-166. Benson, L. 1982. The Cacti of the United States and Canada. Stanford Univ. Press. Stanford. 1044p. Berger, A. 1926. Die Entwicklungslinien der Kakteen. Gustav Fischer, Jena. 103 p. Berger, A. 1929. Kakteen. Eugen Ulmer, Stuttgart. 346p. Britton, N.L. and Rose, J.N. 1920-1923. The Cactaceae Carnegie Institution of Washington Publ 248. 4 vols. Burnett, G.T. Nopalaceae. Pp. 741-745, 1130 in Outlines of Botany vol. 2. John Churchill, London.1190p. Buxbaum, F. 1958. The phylogenetic division of the subfamily Cereoideae, Cactaceae. Madrono 14(6): 177-216. Buxbaum, F. 1974. Cactaceae. In Endler, J. and Buxbaum, F. Die Pflanzenfamilie der Kakteen 3 ed. A. Philler Minden. Crozier, B.S. and Jansen, R.K. 2001. Historical relationships in the American desert plant family Cactaceae. I.0.B.P. Symposium: Origins and Biology of Desert Flora. Albuqueque, New Mexico. Crozier, B.S., Panero, J.P, Jansen, R.K. and Davila Aranda, P.S. 2004. Chloroplast phylogeny and evolution of the new world family Cactaceae (in prep). Phytologia (Aug 2004) 86(2) 63 de Candolle, A.P. 1828. Cacteae. Pp 457-476 in Prodromus systematis naturalis regni vegetabilis III. London. Doweld, A.P. 1999. Tribal taxonomy of Pereskioideae and Opuntioideae (Cactaceae). Succulents 1: 25. Eaton, A. 1836. Botanical Grammar & Dictionary, modernized down to 1836. 4" ed. Engelmann, G. 1876. Cactaceae. Pp. 242-250 in W.H. Brewer and S. Watson, Botany of California Vol 1. Polypetalae. Geological Surver of California Cambridge, Mass. Fearn, P.J. 1996. A Review of the Origins of the Cactus Family and the Search for a System of Classification. Wimborne, Dorset.: P.J. Fearn. Unpaged. Gibson, A.C. 1977. Vegetative anatomy of Maihuenia (Cactaceae) with some theoretical discussions of ontogenetic changes in xylem cell types. Bull. Torr. Bot. Club 104: 35-48. Gibson, A.C. and Nobel, P.S. 1986. The Cactus Primer. Harvard, Cambridge. 286p. Greuter, W., McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Filgueiras, T.S., Nicholson, D.H., Silva, P.C., Skog, J.E., Trehane, P., Turland, N.J., and Hawksworth, D.L. 2000. International Code of Botanical Nomenclature (Saint Louis Code). Koeltz Scientific Books. K6nigstein. 474p. Hunt, D.R. 1967. The genera of the Cactaceae. Pp. 427-467 in Hutchinson, J. The genera of flowering plants vol. 2. Oxford. 117p. Hunt, D.R. 1999. CITES Cactaceae Checklist. 2™ ed. Royal Botanic Gardens Kew and I.0.S. London. 315p. 64 Phytologia (Aug 2004) 86(2) I.0.S. (International Organization for Succulent Plant Study). 1986. The genera of the Cactaceae: towards a new consensus. Bradleya 4: 65-78. 1.0.8. (International Organization for Succulent Plant Study). 1990. The genera of the Cactaceae: progress towards consensus. Bradleya 8: 85-107. Jussieu, A. L.1789. Cacti, Les Cactes. Pp 310-312 in Genera plantarum secundum ordines naturals disposita, juxta methodum in horto regio pariesiensi exaratam, anno M. DCC. LXXIV. Apud Viduam Herissant et Theophilum Barrois. Paris. 498p. Leuenberger, B. 1997. Maihuenia — monograph of a Patagonian genus of Cactaceae. Bot. Jahrb. Syst. 119: 1-92. Lindley, J. 1830. Cacteae. Pp. 54-56 in Introduction to the natural system of Botany. Longman, Rees, Orme, Brown, and Green. London. 374p. Nyfeller, R. 2002. Phylogenetic relationships in the cactus family (Cactaceae) based on evidence from trnK/matK and trnL-trnF sequences. Am. J. Bot. 89:312-326. Ross, R. 1981. Chromosome counts, cytology, and reproduction in the Cactaceae. Amer. J. Bot. 68:463-470. Schumann, K. 1890. Cactaceae. Pp. 187-194 in Martius,K. Flora Brasiliensis vol. 4. Monachii et Lipsiae. Schumann, K. 1898. Gesamtbeschreibung der Kakteen, (Monographia Cactacearum). J. Neumann, Neudamm. 832p. Phytologia (Aug 2004) 86(2) 65 KEYS TO THE FLORA OF FLORIDA -- 10, GALACTIA (LEGUMINOSAE) Daniel B. Ward and David W. Hall Department of Botany, University of Florida Gainesville, Florida 32611, U.S.A. ABSTRACT Galactia (Leguminosae) is represented in Florida by 11 species, one of which consists of three varieties. The application of G. regularis and G. volubilis is reversed from that of other studies. Galactia volubilis var. baltzelliana is described as new, and G. volubilis var. fasciculata is recognized as a new combination. Galactia microphylla is corrected to G. minor. Dioclea multiflora is rejected as a transfer into Galactia. An amplified key is given to the Florida taxa. KEY WORDS: Galactia, Leguminosae, Florida flora. The genus Galactia (Leguminosae) is a familiar yet difficult component of the southeastern flora. Though the taxa of which it is constituted are few, and though some are sharply distinct, others show variations so subtle and intergrading that no two investigators are in agreement as to the number of species or the characteristics by which they may be separated. Further reducing understanding are several recent nomenclatural reinterpretations which impose unfamiliar names or even reverse the application of two well-known names from their accustomed usage. The first comprehensive synopsis of North American Galactia was that of Anna M. Vail (Bull. Torrey Bot. Club 22:500-511. 1895), a study of surprising quality considering the little field information then available. With access to vastly greater herbarium materials and opportunity for field experience, H. J. Rogers (Ph.D. diss., Duke Univ. 1949), under 66 Phytologia (Aug 2004) 86(2) the direction of H. L. Blomquist, prepared a fully detailed systematic treatment of Galactia of the United States; this thesis, regrettably never published, has been underutilized by later authors. Perhaps put off by Vail’s sometimes trichotomous or even quadrichotomous key and lacking ready access to Rogers’s thesis, recent authors have formulated independent treatments to Florida species: A. Herndon, Rhodora 83:471-472. 1981; A. F. Clewell, Guide to the Vasc. Plants of the Florida Panhandle. 1985; R. P. Wunderlin, Guide to the Vasc. Plants of Florida. 1998. Though their summaries are valuable, these authors were either limited in their geographic coverage or placed in synonymy certain taxa recognized by previous authors that now seem worthy of recognition. For decades the acknowledged leader in taxonomic investigation of the American legumes has been Duane Isely of Iowa State University. Dr. Isely, who over a period of years spent many weeks working alongside or in association with the present authors, incorporated many of their observations of Galactia into his treatment of the southeastern species (Vasc. Flora of the Southeastern U. S. 3(2):147-152. 1990). Yet the breadth of Isely’s task (to address all species of Leguminosae throughout the Southeast), together with several significant reinterpretations of current nomenclature, and the value of formalizing a varietal taxon discussed by Isely but not named or fully described -- as well as the differences in interpretation held by authors more recent than Isely -- has stimulated the present authors to submit a further treatment of Florida Galactia. A major nomenclatural complication has been the analysis by W. H. Duncan (Sida 8:170-180. 1979) that the Linnaean epithets applied to two widespread species of Galactia have long been misinterpreted and, moreover, have been used one for the other; that is, the names are valid, but their correct application is the reverse of earlier usage. By Duncan’s assessment, G. regularis (L.) BSP. is the small-flowered (7-9 mm. long), vigorously twining plant previously called G. volubilis, while true G. volubilis (L.) Britt. is the large-flowered (10-14 mm. long), scarcely- twining taxon known elsewhere as G. macreei or G. regularis (Small Phytologia (Aug 2004) 86(2) 67 in part, 1933; Fernald, 1950; Gleason, 1952; Gleason and Cronquist, 1963; Wilbur, 1963; Wilbur in Radford et al., 1964; Clewell, 1985; Isely, 1990; Wunderlin, 1998). In each case, by reference to the actual specimens underlying the pre-Linnaean publications referred to by Linnaeus, Duncan argued these specimens and other materials available to Linnaeus supported this new assessment. Duncan stands nearly alone in this interpretation; it is disappointing that subsequent authors have neither followed Duncan nor chosen to examine or attempt refutation of the factual basis for his nomenclatural judgments. The present authors felt obligated to independently assess the basis for these names. The argument, as reported by Duncan, was perhaps too briefly stated. The basionym of Galactia regularis is Dolichos regularis L. (1753:726). Linnaeus had no specimen directly supporting this name, but gave it the diagnosis: DOLICHOS foliis ovatis obtusis, pedunculis multifloris, petalis aequalis magnitudinis figuraeque, and referred to Gronovius (1739:82). Linnaeus had taken that language from the phrase- name of Gronovius (who further noted it to be racemosis). Gronovius also had cited Clayt. n. 121. John Clayton’s specimen (photo, BM) bears the text, “A trifoliated slender twining plant with small reddish flowers, growing in spikes from the wings [stipules?] of the leaves. Dr. Clayton ex Virginia num. 121.” Clayton’s specimen corresponds to the plant here (and by Duncan) called G. regularis; the “small” flowers are definitive. It thus seems inescapable that Galactia regularis (L.) BSP. is based on the small-flowered plant. [Aside from the specimens themselves, the text accompanying a second Clayton specimen, his 213 (photo, BM), confirms that Clayton recognized both a small-flowered and a large-flowered species. On this second sheet Clayton had written, “A slender twining trifoliated plant with a large purple papilionaceus flower, and a thin compressed pod. This grows in shady woods, an[d] flowers in July. Dr. Clayton ex Virg. n. 213.” This second specimen corresponds to the plant here (and by Duncan) called G. volubilis.] 68 Phytologia (Aug 2004) 86(2) The basionym of Galactia volubilis is Hedysarum volubile L. (1753:750). Again, Linnaeus had no specimen directly supporting this name, but cited three references: Dillenius (1732), Linnaeus’s own Hortus Cliffortianus (1737), and Royen (1740). Only one of these authors appears to have relied directly upon a specimen. Royen merely referred to Hortus Cliffortianus, while Hortus Cliffortianus relegated the plant to its Appendix, an indication that there was no underlying specimen, then cited Dillenius. Fortunately the Dillenius specimen is extant (photo, OXF); Duncan found it to be the large-flowered plant. Thus Linnaeus, by inclusion of the Dillenius reference and, indirectly, Dillenius’s specimen in his circumscription, caused Galactia volubilis (L.) Britt. to be based on the large-flowered plant. The exceptionally larger leaflets and flowers of plants of Galactia volubilis from populations in west-central peninsular Florida, brought to the attention of the present authors by Leland and Lucy Baltzell, Lakeland, has suggested the need for formal recognition. Galactia volubilis (L.) Britt. var. baltzelliana D. B. Ward & D. W. Hall, var. nov. Similis G. volubilis s.s. (sensu. Duncan, 1979), sed floribus grandioribus (a 21 mm. longis), foliis subcoriaceis, et caulibus quibus fortiter scandentibus. TYPE: United States. Florida, Lake Co.: 1505 Moss Ave., 1 mi. n. of Leesburg, 25 Sep 1975, D. W. Hall 413, with L. M. Baltzell (HOLOTYPE, FLAS 151164; ISOTYPES, FLAS 151162, 151163). PARATYPES: Lake Co.: Forest Service road 71, Ocala National Forest, 1.4 mi. e. of Fla. 19, S39, T15S, R27E, 2 Aug 1962, D. B. Ward 3036, with A. A. Will (FLAS); Leesburg, 19 Sep 1967, L. M. Baltzell 153 (FLAS); 0.5 mi. n. of Leesburg, L. M. Baltzell 5327, 8 Aug 1973 (FLAS); Youth Camp, 3 mi. n. of Leesburg, 31 Aug 1980, L. M. Baltzell 11190 (FLAS). Levy Co.: 3 km. n.e. of Cedar Key, 28 Aug 1982, K. A. Kron 1029 (FLAS). Phytologia (Aug 2004) 86(2) 69 The several collections here placed in Galactia volubilis var. baltzelliana were examined by Isely, who agreed (1990:151) they represent a “spectrum of characters” that may deserve formal nomenclatural status. In the field they seem even more distinct than the diagnostic characters indicate, for the stems of var. baltzelliana climb robustly through overlying vegetation, while var. vol/ubilis remains either prostrate or laxly climbing. An 1895 collection from Tampa, western coastal Florida, named Galactia fasciculata, has been difficult to assign. Though accepted as a species by Vail (1895), its original author, and by Rogers (1949), it has been dismissed by later workers as a synonym of G. regularis (here treated as G. volubilis). Yet its fascicled inflorescences and orbicular, deeply emarginate leaves (illustrated by Rogers) do not well correspond to that taxon. It is rare, if not wholly obliterated by the growth of urban Tampa. Rogers cites only a single additional collection (Volusia Co.: Enterprise, 24 Aug 1903, Britton and Wilson s.n. (NY)), and no equivalent plants have been encountered in the present study. That this distinctive entity not be lost, it is here given appropriate varietal rank. Galactia volubilis (L.) Britt. var. fasciculata (Vail) D. B. Ward & D. W. Hall, comb. et stat. nov. Basionym: Galactia fasciculata A. M. Vail, Bull. Torrey Bot. Club 22:505. 1895. Galactia brachypoda Torr. & Gray (1838) is a puzzling entity, based upon two A. W. Chapman specimens (NY) from “Middle Florida” (1.e., central panhandle Florida), thought by Isely (Brittonia 38:352-359. 1986) to represent either a “rare hybrid” or a “freak form” of Galactia erecta. Rogers (1949) cited an additional Chapman specimen (MO) from Wewahitchka, Gulf Co., possibly also the source of the “Middle Florida” specimens. Since this location is a center of endemism on the Gulf coast, these collections may represent an endemic species, long- uncollected or perhaps extinct. [A specimen cited by Rogers from Dade Co. (Hood in 1912 - FLAS) is apparently atypical G. volubilis.] This entity has been held as distinct by most early authors; Rogers provided a suite of characters by which the two may be separated. But for the 70 Phytologia (Aug 2004) 86(2) present, G. brachypodais placed in synonymy of the clearly related, if not conspecific, G. erecta, awaiting further collections and understanding. The transferring authors of the combination Galactia microphylla have been given by Isely (Brittonia 38:352-359. 1986; Vasc. FI. Southeastern U. S. 3(2):150. 1990) as “Rogers ex Hall & Ward.” Though in 1974 D. W. Hall and D. B. Ward had indeed prepared a manuscript, progenitor of the present document, transferring A. W. Chapman’s variety microphylla to specific rank, and had acknowledged H. J. Rogers who had proposed the same transfer in his unpublished 1949 thesis, the Hall- Ward paper was only in draft form when Isely in 1986 called attention to the desirability of specific rank for the taxon. The author credited with this transfer should thus be Isely. The transfer was unnecessary, however, since the taxon is apparently synonymous with G. minor W. H. Duncan (Phytologia 37:59-61. 1977), which has priority at specific rank. The name Galactia parvifolia A. Rich. in Sagra has long been used for a plant of south peninsular Florida, but application of the name and the plant’s relations to other Florida species was unclear. Access to an isotype (photo, P) confirmed that the name, as applied to Florida plants, is correct. A recent proposal that Dioclea multiflora (Torr. & Gray) Mohr be incorporated into Galactia (R. H. Maxwell, Castanea 44:241-246. 1979) is not accepted. Though a case was made that this species is incongruous in its classic assignment to Dioclea HBK., no argument was presented that it would not be even more aberrant within Galactia as historically delimited. The authors are grateful to A. R. Vickery, British Museum, for photostats and suggestions regarding typification of Galactia regularis, to A. Lourteig, Museum National d’Histoire Naturelle, Paris, for help with typification of G. parvifolia, to Walter S. Judd and Kent D. Perkins for constructively reviewing the manuscript, and especially to Duane Isely for his stimulating enthusiasm for all things leguminous. Phytologia (Aug 2004) 86(2) 71 Galactia P. Br. Milk Peas! 1. Leaves pinnately compound, leaflets 5-9; flowers ivory-white; vining perennial herb. Moist to dry thickets, open woodlands, roadsides. Peninsula, throughout (excl. Dade, Monroe Cos.); common. Spring- summer-fall. ELLIOTT’S MILK PEA. Galactia elliottii Nutt. 1. Leaves trifoliolate; flowers pink to red or purple. 2. Stems erect or decumbent, bending alternately (zigzag) at the nodes; flowers pale purple to white; perennial herb. Moist to dry open sandy pinelands. Panhandle (east to Leon, Suwannee Cos.); infrequent. Spring-summer. [?G. brachypoda Torr. & Gray] Galactia erecta (Walt.) Vail 2. Stems prostrate or climbing and twining, not bending alternately at the nodes. 3. Stems prostrate, trailing, or scrambling on low vegetation (or, if vigorously climbing, flowers >13 mm. long). 4. Petioles usually less than 1.5 cm. long; flowers mostly 1-3 per panicle; trailing perennial herb. Dry sandy pinelands. West and central panhandle (east to Liberty, Gadsden Cos.); infrequent. Summer. [G. floridana Torr. & Gray var. microphylla Chapm.; G. microphylla (Chapm.) Rogers ex Isely] Galactia minor Duncan 4. Petioles usually greater than 2 cm. long; flowers 5 or more per panicle. 1. The “amplified key” format employed here is designed to present in compact form the basic morphological framework of a conventional dichotomous key, as well as data on habitat, range, and frequency. This paper is a continuation of a series begun in the 1970s (vide Phytologia 35:404-413. 1977). 72 Phytologia (Aug 2004) 86(2) 5. Leaflets narrowly oblong (or a few oval); calyx 8-9 mm. long; trailing perennial herb. Rocky pinelands. Endemic to south peninsula (Dade Co.: Everglade Keys); rare. All year. Galactia pinetorum Small 5. Leaflets ovate, occasionally narrowly so; calyx usually less than 8 mm. long (if longer, then leaflets broadly ovate). 6. Stems and leaves puberulent to glabrate; trailing perennial herb. Pinelands, hammock margins. Endemic to south peninsula (Dade Co.: Everglade Keys); rare. Spring. [G. prostrata Small] ENDANGERED (Federal, State listings). Galactia smallii Rogers ex A. Herndon 6. Stems and leaves with prominent, spreading pubescence. 7. Calyx 6-8 mm. long; flowers (keel) 11-14 mm. long; trailing perennial herb. Low pinelands. Endemic; panhandle, south to mid-peninsula (Hillsborough, Polk Cos.); infrequent. Summer. [G. regu/aris, misapplied] Galactia floridana Torr. & Gray 7. Calyx 9-12 mm. long; flowers (keel) 11-21 mm. long; trailing or climbing perennial herb. Dry sandy woodlands, thickets. Summer-fall. Galactia volubilis (L.) Britt. 7a. Leaflets near-orbicular, deeply emarginate; inflorescences fascicled. Hillsborough Co.: Tampa; rare, perhaps no longer extant. [G. fasciculata Vail] var. fasciculata (Vail) D. B. Ward & D. W. Hall 7a. Leaflets broadly ovate, obtuse or rounded; inflorescences usually single. Phytologia (Aug 2004) 86(2) 73 7b. Flowers (keel) 11-15 mm. long; leaves thin or slightly coriaceous; stems prostrate or laxly climbing. Throughout; common. [G. glabella Michx.; G. macreei M. A. Curtis; G. regularis, misapplied] var. volubilis 7b. Flowers (keel) 13-21 mm. long; leaves somewhat coriaceous; stems vigorously climbing (but usually without twining). West-central peninsula (Lake, Levy Cos.); rare. var. baltzelliana D. B. Ward & D. W. Hall 3. Stems climbing, twining; flowers (keel) 7-14 mm. long. 8. Mature fruits 5.5-9 mm. broad, clearly falcate; stems woody, high climbing; flowers red-violet; twining perennial herb. Brushy coastal thickets, pinelands. South peninsula (north along west coast, to Lee County); locally frequent. All year. [G. spiciformis Torr. & Gray] Galactia striata (Jacq.) Urban 8. Mature fruits less than 6 mm. broad, straight or slightly falcate; stems not woody, not very high climbing. 9. Leaflets with long soft sometimes matted hairs on both surfaces; calyx long-pubescent; petals dark red to rose-purple, drying dusky rose; ovary and pod densely pubescent with long soft hairs; twining perennial herb. Dry sands of high pinelands. North Florida, from mid-panhandle (Franklin Co.), to east coast (Duval Co.); infrequent. Spring-summer. Galactia mollis Michx. 9. Leaflets with stiff appressed or spreading short hairs, or glabrous above; calyx appressed to spreading short-pubescent; petals pink to purple, drying white to variously purple (but not dusky rose); ovary and pod with stiff appressed to spreading short hairs. 74 Phytologia (Aug 2004) 86(2) 10. Leaflets elliptic to narrowly ovate; pubescence spreading; twining perennial herb. Moist to dry thickets, open hammocks, roadsides. Throughout; common. Spring- summer-fall. [G. mississippiensis (Vail) Rydb.; G. volubilis, misapplied] Galactia regularis (L.) BSP. 10. Leaflets linear to narrowly elliptic; pubescence appressed; twining perennial herb. Pinelands. South peninsula (Dade Co.: Everglade Keys; Monroe Co.: Florida Keys); infrequent. Fall-winter-spring. Galactia parvifolia A. Rich. in Sagra Phytologia (Aug 2004) 86(2) 75 TWO NEW SPECIES OF WEDELIA (ASTERACEAE: HELIANTHEAE), WITH A CONSPECTUS OF THE GENUS FOR MEXICO B. L. Turner Plant Resources Center University of Texas Austin, TX, 78712-0471, USA ABSTRACT Two new species of Wedelia are described from Mexico: W. elottiana from the state of Jalisco, and W. juxtlahuacana from the state of Oaxaca. Additionally, a new combination, Wedelia xylopoda (Greenm.) B. L. Turner is proposed. A conspectus of the 27 species currently recognized for Mexico is presented, along with a key to the taxa. KEY WORDS: Asteraceae, Wedelia, Mexico, Jalisco, Oaxaca. As noted below, Strother (1991) provided a taxonomic account of Wedelia for North America. Turner (1992a,b) subsequently provided an assessment of the Mexican species, this in connection with his forthcoming treatment of the Heliantheae of Mexico (cf. Comps of Mexico 5: Eupatorieae, Phytologia Memoirs 11. 1997). The following account of Wedelia will acquaint the interested reader with this formidable enterprise. Both of the new taxa are described and keyed in this treatment, and will appear as such in the final product, except for a shortening of their descriptive accounts. Additionally, all of the taxa will be mapped, much as for taxa of the tribe Eupatorieae, as alluded to in the foregoing. 76 Phytologia (Aug 2004) 86(2) WEDELIA Jacq. Aspilia Thouars Complaya Strother Thelechitonia Cuatrecasas Suffruticose perennial herbs, shrublets or shrubs 1-3 m high. Stems erect, procumbent or prostrate, arising from branched ligneous roots or woody crowns, rarely rhizomatous. Leaves mostly opposite, rarely alternate above, simple or 3-lobed, 3-nervate or rarely pinnately nerved. Heads mostly radiate, 1-several, terminal or rarely axillary, mostly 1-4 cm across the extended rays. Involucres 2-3 seriate, subequal or rarely graduate, the outer series often green and somewhat foliate. Receptacle plane to convex, paleate. Ray florets mostly 5-21 (rarely absent), pistillate or neuter, fertile or sterile, the ligules yellow, white or rarely purple. Disk florets 15-150, the corollas yellow. Achenes, those of the ray, either 2- or 3-sided, when 3-sided usually somewhat tangentially compressed and winged, those of the disk radially compressed, with or without apical wings; pappus a sessile or raised narrow crown of minute scales arising from the center of a clavate body, the borders of which may or may not possess 1-3 rather delicate bristles. Base chromosome numbers, x = 10, 11, 12, 14 and 15. Type species, Wedelia fruticosa Jacq. REFERENCES McVaugh, R. 1984. Wedelia, in Flora Novo-Galiciana 12: 1080-1092. Nash, D. 1976. Wedelia, in Flora Guatemala, Fieldiana, Bot. 24: 349- 354. Pruski, J. F. 1996. Compositae of the Guayana Highland-XI... Novon 6: 404-418. Robinson, H. 1984. Studies in the Heliantheae (Asteraceae) XXXIIL... Phytologia 55: 415-423. ---------- and J. Cuatrecasas. 1992. Thelechitonia Cuatrecasas, an older name for Complaya Strother (Ecliptinae-Heliantheae-Asteraceae). Phytologia 72: 141-143. Strother, J. 1991. Taxonomy of Complaya, Elaphandra, logeton, Jefea, Wamalchitamia, Wedelia, Zexmenia, and Zyzyxia (Compositae: Heliantheae: Ecliptinae).Syst. Bot. Monographs 33: 1-111. Phytologia (Aug 2004) 86(2) 77 Strother, J. 1999. Wedelia, in Flora Chiapas 5: 149-151. Turner, B. L. 1988. New species, names and combinations in Wedelia (Asteraceae). Phytologia 65: 348-357. ---------- . 1992a. New Species of Wedelia (Asteraceae, Heliantheae) from Mexico and critical assessment of previously described taxa. Phytologia 72: 115-126. ---------- . 1992b. New names and combinations in New World Wedelia (Asteraceae, Heliantheae). Phytologia 72: 389-395. A large complex genus of perhaps 200 or more species, widely distributed throughout the tropical world, but largely centered in South America. It is closely related to Zexmenia but the latter can be distinguished by its thin-winged, flatter, disk achenes which mostly possess rather persistent awns at maturity, these usually broadened at the base and arising from the corner of the achenes (as opposed to a cup-like boss at the apices). I follow Mc Vaugh (1984) and Turner (1988, 1992a,b) in treating Aspilia within the broad fabric of Wedelia. Strother (1991) revised Wedelia for North America; in this he excluded W. trilobata, placing it in a newly erected genus, Complaya (=Sphagneticola, if accepted, the earliest name; cf. Pruski 1996), which is largely distinguished by its style branches and indistinct carpopodium, among other characters. Since the phyletic position of W trilobata is moot, I have retained this in Wedelia, pending further elucidation. Robinson (1992: Phytologia 72: 144) recently agreed that Aspilia is synonymous with Wedelia, although his earlier work (1984) suggested otherwise. KEY TO SPECIES 1. Ray florets without style branches (neuter), or rays absent.......... (2) 1. Ray florets with style branches (pistillate and fertile)............... (16) DARAY HOLES: PLCSCN beactye eerie aes et Ne er ere ia Sekt sia (4) PARA SOTEESNA DSM tts an aeeerey ee atl ogy Meat ae emia een: (3) 3. Leaves markedly lacerate; vestiture of stem softly EIESISLS AVI 7 ieee cetera mm ees es aae im eerste aan aie ae W. keilti 78 Phytologia (Aug 2004) 86(2) 3. Leaves remotely serrulate to nearly entire; vestiture roughly hispid or hispidulous; Jal........................06- W. aggregata A Rays whiteior yellow. 2. ve.5)Sa.i cee ene ele ce So Ee (6) AURays tedior purplish’. .c12e2eeseeceeaccteewe ace hee eee eee (5) 5. Leaves mostly elliptical to oblanceolate, widest above the middle7Gues@axa@psis-. ue eereec nena etree W. purpurea 5. Leaves mostly lanceolate (to rarely lanceolate- elliptic), widest below the middle; Sin, Dur, Nay.......... W. xylopoda 6(4). Lower surfaces of leaves minutely appressed- strigose; leaves linear-lanceolate, 5-10 cm long, with only a single well- defined mid-vein; Son, Chi, Sin.......... W. greenmanii 6. Lower surfaces of leaves variously pubescent, but not as described in the above, the leaves various, 3-nervate if linear-lanceolate, then pilose or scabrous beneath with coarse haitss.2...c2.asscohseeceece cach fee Se eReee eee (7) 7. Leaves spatulate, 1.0-1.5 cm wide; rays white; Nay, Jal AiG 25 ee renee oes a Bacteria we Lane eee W. grayi 7. Leaves ovate to ovate-lanceolate; rays yellow, salmon-colored or pinkish (white in W. mexicand)...........-.++0+++ (8) 8. Well-developed leaves on primary branches with blades mostly 35-100 mm long; stems erect or suberect.................. (11) 8. Well-developed leaves on primary branches with blades mostly 15-30 mm long:2.25: 2-422..32= 5s eee (9) 9. Heads aggregate, sessile or nearly so, if solitary then peduncles 1-8 mm long; vicinity of El Tuito, Jal............ W. talpana 9. Heads solitary, not aggregate, the peduncles mostly 10-40mm long 3333) 222.2250 5.2822 eee (10a) Phytologia (Aug 2004) 86(2) 79 10a. Ray corollas salmon-colored, pinkish or white; Mex, CUTS a AN ern ROR Ta MG es W. hintoniorum 10a. Ray corollas yellow or yellow-orange; Sin, Dur, INVA are hens eis ei cle orate eet areata ea /auid ck sateen ON ante (10b) 10b. Rays yellow; leaves widest at or above the middle; Opec ee eet ae oso Se ee ee a ea eextlahuacana 10b. Rays white; leaves widest above the middle; STMT YIN as. bees st eS eae See eno A nexiCana 11(8). Leaves broadly ovate, 1.5-2.5 times as long as wide; Stcmsilax Onrecumbent-Chizvenes nee ee W. chihuahuana 11. Leaves ovate-lanceolate, elliptic-ovate to lanceolate, B-OmlMes ASHONGaSsWIdE sss saece see eee ee echt eran: (12) 12. Leaves markedly lacerate, the lower surface bestowed with few to numerous minute uncinate MaRS OHI a) caste soonceanclaanoecmameniee anette eminent W. pimana 12. Leaves remotely serrulate to entire, the lower surfaces without uncinate hairs; Sin, Dur, Nay, Jal.................. (13) 13. Primary leaves lanceolate-ovate to elliptic-ovate, those at mid-stem mostly (6)10-20 mm wide; involucres PIOSUVAS 1 MIMIC. hacer atrs eden maccise socio os Sone elec Cen (15) 13. Primary leaves linear-lanceolate, those at mid-stem mostly 3-10 mm wide, the margins enrolled; involucres MoOstlyiG-Ssamimynt Shure alee Aceemseee cence ea eee (14) 14. Divaricately branched low shrublets 10-30 cm high; leaves and stems with erect or ascending hairs; SADIE sect sek See OR oR RRR ey ene W. gonzaleziarum 80 Phytologia (Aug 2004) 86(2) 14. Stiffly erect shrublets 30-100 cm high; leaves and stems with appressed hairs, or if erect then numerous and very short, superficially strigose; Jal...... W. cronquistii 15(13). Hairs on achenes mostly 0.1-0.2 mm long; Col Mic eG IGS, See EEA oii SO ea W. strigosa 15. Hairs on achenes mostly 0.2-0.5 mm long; Nays Jali ZacyA ou dcmosike neha cae hee eae aoe ae ee W. rosei 16(1). Stems prostrate or trailing, rooting at the nodes, forming loose, but copious ground-cover; leaves, at leastisome‘onthems neatly 3-lobedasee 2a: atag eee ete W. trilobata 16. Stems erect, not normally rooting at the nodes: leaves not neatly 3-lobedin.c2..2---secec cece eee eee (17) 17. Erect to recumbent herbs or shrublets, not normally rhizomatous; style branches gradually ACUIMMTMALS 2.5. ccds Sa steke ako csec os cee ako ssood sled Sisians aati seiote Verse latei ae eens Glo 17. Stiffly erect, rhizomatous, coarsely hispid herbs to | m high, forming clumps in wet meadows; style branches abruptly apiculate; Nay, Jal....-.....2-.-.. as-.sss ener (18) 18. Leaves sessile, the blades cordate or subcordate Att DAS Sa Leases caso ctoce Cecloc cise ucta vote Sepak Saat ae a W. cordiformis 18. Leaves with winged petioles, the blades otherwise BO OCS on cani de Sona ere tno aaeriotadoohcbcasa W. simsioides 19(17). Leaves with hairs predominently alike, if of two kinds then the finer hairs not uncinatess2 222+ -.cosec eee eee (23) 19. Leaves with two kinds of hairs on the undersurface, a longer group of straight hairs and below these at least a few uncinate or recurved hairs............ (20) 20. Stems suffruticose, stiffly erect, to 1.5 m high, sparsely branched; achenes of the ray and disk similar, unwinged, or rarely the ray achenes winged; Nue, Tam, San, (Gulf Slopes)................. W. ayerscottiana Phytologia (Aug 2004) 86(2) 81 20. Stems variously procumbent to erect, the plants often divaricately-branched, shrubby, and up to 2 m high; achenes often heteromorphic, those of the ray and disk winged or not; Pacific slopes and Geitinal WVESOULNW ARGUS #24 erhrcets creel ee ee mane nen er eet ean ee (21) 21. Leaves elliptic to lance-linear, sessile or nearly so; Racitierslopess Nayadalo caaceatcoscw scene se cen eek ce W. vexata 2laeaves ovate to lanceolate; clearly petiolates.23. 2us.- esac: (22a) 22a. Perennials; ray corollas bright yellow, the ligules mostly 5 mm long or more; widespread................ W. scabra 22a. Annuals with slender tap-roots; ray corollas pale yellow to white, the ligules 1-5 mm long; Jal, Cps.................. (22b) 22b. Leaves lanceolate; disk florets mostly sterile; Jal......W. elottiana 22b. Leaves oval to ovate; disk florets mostly fertile; Cps.....W. iners 23(19). Erect, coarse, suffruticose, shrubs or shrublets mostly 1-2 m high; Ver, Oax, Cps (Isthmus of sliehwantepee) Aner sie ease eee eee W. tehuantepecana 23. Perennial herbs or suffruticose shrublets 0.2-0.9 m high Sontoltidimorth wards ieenscees ee sece teeta arene (24) 24. Stems stiffly erect, suffruticose, becoming a bushy shrublet; leaves commonly lacerately- lobed at the base; widespread, e Chi, Coa, Nue, AParmigtoy Mie xuote io tateis ebb) ote hd dina rat Ai Ret Se Oe W. hispida 24. Stems procumbent, recumbent, or erect suffruticose herbs; leaves mostly ovate and evenly serrate; SontweChinRaciticslopes: stone ease oe ae eae ee (25) 25. Leaves mostly 2.5-6.0(8.0) cm long; herbs 30-60 cm highs wiChivandadjacent:Somm.asee-ce sana W. chihuahuana 25. Leaves mostly 6-12 cm long; shrubs or shrublets to PS nubiohsbaciic slopes alitoiGueeat rec. see oe W. strigosa 82 Phytologia (Aug 2004) 86(2) WEDELIA AGGREGATA (Greenm.) B. L. Turner, Phytologia 65: 348. 1988. Aspilia aggregata Greenm. Known only by the type, Jal, between Bolanos and Guadalajara; Sep. Superficially resembling W. rosea but the heads rayless, sessile and the involucres narrowly campanulate with graduate purple bracts. Turner (1988), not having examined the type, treated this taxon as encompassing W. rosea. Thanks to Strother’s study of the group (1991) in which the two taxa are treated as distinct species, Turner (1992a,b) reevaluated the group and agreed with Strother’s assessment. WEDELIA AYERSCOTTIANA B. L. Turner, Phytologia 60: 125. 1986. Wedelia acapulcensis var. averscottiana (B. L. Turner) Strother Nue, Tam, San, mostly Gulf slopes, pine-oak forests, 400-2000 m; Jun-Oct. Stiffly erect, simple-stemmed, suffruticose herbs to | m high; much resembling W. scabra but the fruits of both ray and disk florets similar, maculate, wingless and the leaves often rather lacerate and sparsely pubescent beneath. This species has the leaf-vestiture of W. scabra but otherwise looks like W. hispida; the involucres are quite narrow with striate bracts, resembling those of Aldama dentata. It usually occurs in low cloud forests with the cycad genus Dioon along the eastern flanks of the Sierra Madre Oriental. WEDELIA CHIHUAHUANA B. L. Turner, Phytologia 65: 348. 1988. w Chi canyons in pine-oak forests, 1700-2100 m; Jun-Sep. Trailing or weakly ascending perennial herbs to 60 cm high; leaves mostly 2-8 cm long, 1-5 cm wide; petioles 1-4 mm long; blades ovate, 3-nervate, moderately to sparsely pubescent beneath with straight hispid hairs, the margins serrate to somewhat irregularly lacerate; heads campanulate, radiate, 1-2 cm across the extended rays, the ultimate Phytologia (Aug 2004) 86(2) 83 peduncles 2-8 cm long; involucres 6-10 mm high, 2-3 seriate, the bracts subequal or the outer somewhat longer. Ray florets 8-13, the ligules 5- 10 mm long. Disk florets 30-50, the corollas yellow, 4.5-5.0 mm long. Achenes unwinged, the pappus a crown of scales ca 0.5 mm high and 1 or 2 lateral bristles, 1.0-1.5 mm long. Much-resembling the more eastern W. hispida but the leaves evenly serrate, pubescent principally along the veins beneath, the heads borne 2-3 in lax corymbs. It might also be mistaken for W. strigosa, but the latter is a shrub or shrublet confined to the Pacific slopes from 0-300 m. WEDELIA CORDIFORMIS McVaugh, Contr. Univ. Michigan Herb. 9: 460. 1972. Known only from the type (Jal, ca 12 km wnw Tototlan towards Zapotlanejo), poorly drained meadows, ca 1800 m; Aug. Perennial herbs, growing in clumps to | m high; stems stiffly erect, very coarsely hispid, 4-5 mm thick at mid-stem; leaves opposite, 6-10 cm long, 2-4 cm wide, sessile or nearly so; blades lance-ovate, somewhat clasping, pinnately-veined, roughly hispid, the margins serrulate; heads radiate, 4-5 cm across the extended rays, the ultimate peduncles 5-15 cm long; involucres 13-17 mm high, the bracts subequal; ray florets 13-18, the ligules 15-20 mm long; achenes pubescent, the pappus of a single awn ca 1.5 high and a crown of scales ca 0.7 mm high. A very distinct species, not easily confused with another. Mc Vaugh (1984) provided an excellent illustration of the taxon. WEDELIA CRONQUISTII B. L. Turner, Phytologia 65: 350. 1988. Known only from Jal (ca 23 km min of Guadalajara), oak woodlands, 1500-1600 m; Nov. Suffruticose perennial herbs to 1 m high; leaves opposite throughout, 3-6 cm long, 2-6 mm wide; petioles 0-1 mm long; blades linear to linear- 84 Phytologia (Aug 2004) 86(2) lanceolate, pinnately nervate, densely strigose beneath with coarse hispid hairs; heads radiate on ultimate peduncles 3-10 cm long; involucres campanulate, 3-4 seriate, 10-12 mm high, the bracts subgraduate to nearly equal; ray florets 5-11, the ligules yellow, 6-9 mm long; disk florets 30-50; achenes clavate 4.0-4.5 mm long. Resembling W. rosei but the leaves markedly appressed-strigose beneath and the heads larger (involucres 10-12 mm high vs 6-10 mm). Strother (1991) placed W. cronquistii in synonymy under his concept of W. greenmanii, a view with which Turner (1992a,b) disagreed, the characters given in his key being compelling, not to mention their geographic disparities. WEDELIA ELOTTIANA B. L. Turner, sp. nov. Similis Wedeliae iners (S. F. Blake) Strother sed foliis tenioribus, integris vel paene integris, habentibus capitula fructifera parviora (4-6 mm alta vice capitulorum 7-10 mm altorum) et flores discorum steriles pro parte maxima. Stiffly erect tap-rooted annual herbs to 50 cm high. Stems moderately pilose with an array of short to long hairs 0.2-1.0 mm long. Leaves alternate throughout; petioles 1-3 cm long; larger leaves with blades broadly lanceolate to ovate, thin, pinnately veined, 5-7 cm long, 1.5-2.0 cm wide, pubescent below and above with scattered coarse hairs to 1 mm long, below these a minute array of mostly uncinate hairs; margins entire or nearly so. Heads borne on ultimate peduncles 3-7 cm long; involucres composed of ca 8 subequal bracts in 2-3 series. Receptacles small, plane, the bracts lanceolate, ca 5 mm long. Ray florets 5-6, pistillate, fertile; tubes ca | mm long; ligules ovate, ca 5 mm long, 3.5 mm wide, weakly 2-nervate; achenes ovoid, markedly winged, ca 5 mm long, 4 mm wide, the body of the achene enclosed within broad corky wings, the latter 0.6-1.0 mm wide. Disk florets yellow, 3-4 mm long; tubes 1.0- 1.5 mm long; lobes 5, deltoid, ca 0.5 mm long, mostly sterile (the style branches seemingly fused ventrally, but a few of the outermost florets apparently producing clavate wingless fertile achenes 3-4 mm long, 0.6-1.0 mm wide). Phytologia (Aug 2004) 86(2) 85 TYPE: MEXICO. JALISCO: Mpio. La Huerta, Rancho Cuixmala, Gargollo farm, on E side of Cerro de la Alborada. “Tropical deciduous Forest on slopes of Cerro de la Alborada, following the water main which reaches main Cuixmala entrance road. Uncommon perennial, apparently flowering first year from seed; uncommon on rocky slope along pipeline trail in partial shade. Flowers yellow.” Elevation “below 50 m.” 4 Nov 1991, Emily J. Lott 4135 (HOLOTYPE: TEX; ISOTYPES: UCR). This taxon is apparently closely related to the more southern W. iners, with which it is compared to in the above diagnosis, and which I identified it as, upon receipt of type material. While Emily thought the species to be a perennial in its first year of growth, the specimen appears to be an annual, much as she noted. At least I know of no other perennials of a similar nature from the area concerned. It is a pleasure to name this striking plant for my friend of many years; she has a long history of field work in both Texas and Mexico. Emily currently resides in Austin, Texas with her family. WEDELIA GENTRYI B. L. Turner, Phytologia 66: 500. 1989. Chi, and n Sin, pine-oak woodlands, 1800-2300 m; Jul-Sep. Much-resembling W. greenmanii but readily distinguished by its roughly hispid hairs on the lower leaf surfaces, stems and involucres, the hairs spreading or ascending (vs minutely strigose and closely appressed throughout in W. greenmanii). Strother (1991) submerged this taxon in his concept of W_greenmanii, this refuted by Turner (1992a,b). WEDELIA GONZALEZIARUM B. L. Turner, Phytologia 65: 349. 1988. Known only from se Dur (Mpio. Mezquital), pine-oak forests, ca 2400 m; Oct-Nov. 86 Phytologia (Aug 2004) 86(2) Much-resembling W. rosei but the stems mostly short and ascending to procumbent (10-25 cm high vs 100-150 cm), the leaves coarsely hispid beneath (vs softly silky pilose), the heads single at the apices of branches (vs 2-4 on short peduncles (mostly 1-2 cm long vs mostly 2-8 cm). The specific name was originally spelled, in error, W. gonzalezorum, as noted by Strother (1991). WEDELIA GRAYI McVaugh, Contr. Univ. Michigan Herb. 9: 462. 1972: Aspila albiflora A. Gray, not Wedelia albiflora Hiern. Nay, Jal, and s Zac, pine-oak woodlands, grassy areas, 1500-2500 m; Aug-Oct. Much-resembling W. mexicana but the leaves longer, the blades mostly elliptic to oblanceolate, stems ligneous and branched so as to form low mats; those of W. mexicana, while decumbent, appear to arise from a well-defined, woody, corm-like, caudex or rootstock; in addition, the rays of W. mexicana are yellow and the heads are smaller. WEDELIA GREENMANIIB. L. Turner, Phytologia 65: 352. 1988. Aspilia stenophylla Greenm., not Wedelia stenophylla Merrill Son, w Chi and ne Sin, pine-oak woodland, 1000-2000 m; Jul-Nov. Suffruticose herbs or shrublets 50-70 cm high; stems brittle, erect, much-branched from the base; leaves opposite throughout, 5-14 cm long, 3-12 mm wide; petioles 0-5 mm long; blades linear to linear-lanceolate, pinnately nervate or weakly 3-nervate below, minutely appressed- hispid throughout, the margins remotely serrulate to nearly entire; heads radiate, 2-5 in slender terminal open corymbs, the ultimate peduncles 3-8 cm long; involucres campanulate, 6-7 mm high, 2-3 seriate, the bracts subequal, the outer series with acute or acuminate apices, the inner series obtuse or rounded; ray florets 5-8, neuter, the ligules yellow, 5-6 mm long; disk florets 20-30, the corollas yellow; achenes broadly clavate, Phytologia (Aug 2004) 86(2) 87 ca 5 mm long, wingless, pubescent, the pappus a short narrow crown of scales ca 0.7 mm high. Strother (1991) recognized specimens from Jal as belonging to this taxon, but these appear to belong to a distinct species, W. cronquistii (Turner 1992a,b). WEDELIA HINTONIORUM B. L. Turner, Phytologia 65: 352. 1988. sw Mex and e Gue, pine-oak forests, 2000-2300 m; Jun-Aug. Perennial herbs to 25 cm high; stems suffruticose erect or recumbent from a woody, corm-like, root or crown; leaves 1.3-3.0 cm long, 3-7 mm wide; petioles absent or nearly so (0-1 mm long); blades narrowly ovate to oblanceolate, weakly 3-nervate, hispidulous above and below, the margins somewhat serrulate apically; heads single on peduncles 3-12 mm long; involucres 8-12 mm high, 2-3 seriate, the bracts subequal, the outermost 3-5, mostly linear-oblanceolate, leaf-like; receptacular bracts linear-lanceolate in outline, 3-fid, about as long as the florets; ray florets 8, the ligules reportedly salmon-colored but drying white, 6-9 mm long; disk florets ca 30, ca 5 mm long, glabrous. This species superficially resembles W. grayi and W. mexicana; it differs from both in possessing nearly sessile, hispidulous leaves. WEDELIA HISPIDA H.B.K., Nov. Gen. & Sp. 4: 216. 1820. Aspilia potosina Brandegee Lipochaeta texana Torrey & Gray Seruneum hispidum (H.B.K.) Kuntze Viguiera longipes J. Coulter Viguiera pauciflora Brandegee Wedelia acapulcensis var. hispida (H.B.K.) Strother Wirtgenia texana (Torrey & Gray) Sch.-Bip. Zexmenia hispida (H.B.K.) A. Gray Zexmenia texana (Torey. & Gray) A. Gray 88 Phytologia (Aug 2004) 86(2) eChi, Coa, Nue, Tam, San, Que, Hid, Mex and adjacent U.S.A., dry calcareous hills, and rocky slopes 300-2000 m; Apr-Sep. Much resembling W. scabra but the plants more branched from the base and the leaves mostly smaller, narrower and lacking the minute uncinate hairs on the undersurface which distinguish that species; the heads are also borne singly on very elongate peduncles (mostly 15-30 cm long); chromosome number, n = ca 26 pairs. Strother (1991) considered this taxon to be a variety of his W. acapulcensis (= W. scabra in the present account) but it appears to be amply distinct, at least I discern no intergradation of the characters which distinguish between these. Wedelia hispida superficially resembles W. chihuahuana of w Chi but the former is readily distinguished by its coarsely and evenly pubescent leaves and larger solitary heads. WEDELIA INERS (Blake) Strother, Syst. Bot. 33: 72. 1991. Zexmenia iners Blake Cps, e Oax?, and Guatemala southwards, tropical forests, often weedy along streams and wet thickets, 50-900 m; all seasons. Erect annuals to 60 cm high, the stems arising from rather delicate well-defined tap-roots; leaves mostly 3-11 cm long, 1-4 cm wide; petioles 0.5-1.5 cm long; blades broadly ovate to lanceolate; ray florets 5, pistillate, fertile, the ligules white to pale yellow, mostly 1-4 mm long; chromosome numbers, n= 11, ca 22 pairs (Strother, 1991). A very distinct taxon, having the vestiture of W. scabra but readily distinguished by its annual habit and heads with small ray florets. WEDELIA JUXTLAHUACANA B. L. Turner, sp. nov. Similis Wedeliae hintoniorum B. L. Turner, sed radii lutei sunt (vice alborum aut salmoneorum), et folia ovalia sunt (latissima prope medium vice oblanceolatorum vel linearium-oblanceolatorum, latissimorum super medium). Phytologia (Aug 2004) 86(2) 89 Prostrate rhizomatous perennials forming clumps 20-50 cm across (label data). Stems purplish, bearing up-turned appressed hairs. Leaves opposite; blades narrowly to broadly oval, 1.5-2.2 cm long, 0.6-1.0 cm wide, 3-nervate from near the base, pubescent above and below like the stems; margins weakly serrulate. Heads terminal, single, ca 3.5 cm across the extended rays. Peduncles 3-5 cm long. Involucres 8-9 mm high, ca 10 mm wide (pressed); outermost bracts 3-5, lanceolate, eciliate; inner bracts 5-8, oblanceolate, ca as long as the outer bracts, their apices scarious, rounded, ciliate. Receptacular bracts lanceolate, somewhat longer than the subtended florets. Ray florets 7-8, neuter; ligules yellow, ca 1.5 cm long, 0.3-0.5 cm wide. Disk florets ca 20, fertile; corollas yellow, glabrous, 6-7 mm long. Anthers black, their appendages eglandular, linear-lanceolate, ca 1 mm log. Achenes (immature) ca 3 mm long, appressed-pubescent, the pappus a short erose crown. TYPE: MEXICO. OAXACA: SANTIAGO JUXTLAHUACA. Mpio. San Martin Peras, “Km 18 de la desviacion a Canada de Lobos sobre el nuevo camino a campos de fresas.” 2000 m, 24 Oct 1994, J. L. Panero 5274 (HOLOTYPE: MEXU; ISOTYPES: MSU, TEX!). This novelty is closely related to W. hintoniorum, having most of the attributes of that species. When the latter was first described I took its ligules to be yellow, although, as noted in the description, this was not clear in the holotype. Subsequent collections prove the corollas to be white (Panero 7352, 5 Oct 1997, TEX), this also noted by Strother (1991). The species is named for Santiago Juxtlajuaca, where first collected. WEDELIA KEILI B. L. Turner, Phytologia 65: 353. 1988. Known only from the type (ca 25 km s of Uruapan, Mic), grassy hillsides, ca 1000 m; Aug. Perennial herbs to 25 cm high; much-resembling W. mexicana but the leaves with markedly lacerate margins and the plants densely soft- pilose throughout; chromosome number, n = 22 pairs. 90 Phytologia (Aug 2004) 86(2) This species superficially resembles the widespread W. scabra but differs in its smaller, herbaceous, habit, softly pubescent foliage, lacerate leaves, etc. WEDELIA MEXICANA (Sch.-Bip.) Mc Vaugh, Contr. Univ. Michigan Herb. 9: 462. 1972. Aspilia mexicana (Sch.-Bip.) Hemsl. Wedelia tegetis Strother (in part) Wirtgenia mexicana Sch.-Bip. Sin and adjacent Dur, pine-oak woodland along hiway 40, talus slopes, 1500-2500; Oct-Nov. Perennial herbs 15-30 cm high; stems weakly ascending to procumbent, hirsutulous; leaves opposite throughout, mostly 2-3 cm long, 8-12 mm wide; petioles 5-10 mm long; blades ovate, weakly 3- nervate, roughly hispid above and below, the hairs recurved but not uncinate; the margins serrulate to nearly entire; heads radiate, single and terminal, the peduncles 1-3 cm long; involucres 5-6 mm high, 2-3 seriate, the bracts subequal; ray florets 5-8, the ligules 5-8 mm long; achenes 3.0-3.5 mm long, pubescent, the pappus a crown of scales ca 0.5 mm high; chromosome number n = 12 pairs (Mayfield 1061 TEX). McVaugh (1984) knew this taxon only from the type collection; several additional collections have been examined from the Sin-Dur border (TEX) along highway 40, west of Durango City. Strother (1991) applied the name W. tegefis to all of this material, except for the type of W. mexicana, Which he retained as distinct, although its type was probably collected in the same region as his W. tegetis (Turner 1992a,b). Strother also included in his concept of W. tegetis, white-rayed specimens with relatively large heads from n Nay. I include these within my concept of W. grayi. WEDELIA PIMANA B. L. Turner, Phytologia 72: 116. 1992. Chi, waterfall at Nabogame, grassy areas, 1800 m; Aug. Phytologia (Aug 2004) 86(2) 91 Much resembling W. scabra but the ray florets neuter and the leaves markedly lacerate. Wedelia pimana might also be compared with W. chihuahuana but the latter has recumbent or sprawling stems and a very different, merely hispid vestiture. WEDELIA PURPUREA (Greenm.) B. L. Turner, Phytologia 65: 354. 1988. Aspilia purpurea Greenm. Aspilia scabrida Brandegee Gue, Oax and Cps, pine-oak forets, 900-2100 m; Jun-Nov. Perennial, often suffruticose, herbs 15-50 cm high; stems prostrate to recumbent, arising from an enlarged woody crown; leaves opposite throughout, 2-4 cm long, 0.5-2.0 cm wide; petioles 1-4 mm long; blades ovate-elliptic to elliptic-oblanceolate, 3-nervate, sparsely to moderately hispidulous beneath; heads radiate, single, terminal, the peduncles mostly 4-20 cm long; involucres campanulate, 2-3 seriate, 8-10 mm high, the bracts subequal; rays 8-11, the ligules deep purple or maroon; disk florets 20-40; n=ca 12 or 13 pairs. Because of its purple rays, this species is readily distinguished from most of the Wedelias in southern Mexico. WEDELIA ROSEI (Greenm.) McVaugh, Contr. Univ. Michigan Herb. 9: 462. 1972. Aspilia angusta Blake Aspilia angustifolia A. Gray (not A. angustifolia Oliv. & Hiern.) Aspilia rosei Greenm. Aspilia xylopoda Greenm. Sin, Dur, Nay, Jal, s Zac and Agu, pine-oak woodlands, 800-2500 m; Aug-Nov. 92 Phytologia (Aug 2004) 86(2) Suffruticose herbs or shrublets 1.0-1.5 m high; stems coarsely hispidulous, stiffly erect, arising from a woody root-crown from which arise several thick, seemingly fibrous, roots; leaves opposite throughout, mostly 8-12 cm long, 0.6 -2.5 cm wide; petioles 1-10 mm long; blades narrowly lanceolate to elliptic-lanceolate, coarsely hispid above, rather softly silky-pilose beneath, pinnately nervate or weakly 3-nervate below, the margins serrulate to entire; heads mostly 2-4 in strict terminal cymes, the ultimate peduncles mostly 2-15 cm long; involucres campanulate, 6-10 mm long, the bracts subgraduate to subequal, strigose, loosely arranged and often recurved at the apices; ray florets 8-11, the ligules yellow or rarely white to purplish, 7-16 mm long; disk florets 20-60, yellow; achenes 3.5-5.0 mm long, pubescent, the pappus a lacerate crown 0.6-1.0 mm high. As noted by McVaugh (1984), this is a highly variable species. Turner (1988) accepted McVaugh’s suggestion that the type of Aspilia aggregata is an aberrant individual of W. rosei with nearly sessile heads; since the name concerned predates W. rosei, priority required taking up the former name. Strother (1991) accepted W. rosei as distinct from W. aggregata, the latter known only from the type and differing markedly from W. rosei in having rayless, sessile heads. Turner (1992a,b) subsequently examined the types concerned and agreed with Strother’s assessment. WEDELIA SCABRA (Cav.) B. L. Turner, Phytologia 65: 354. 1988. Buphtalmum scabrum Cav. Seruneum acapulcense (H.B.K.) Kuntze Wedelia acapulcensis H.B.K. Wedelia acapulcensis var. cintalapana Strother Wedelia acapulcensis var. parviceps (Blake) Strother Wedelia acapulcensis var. ramosissima (Greenm.) Strother Wedelia hispida var. ramosissima (Greenm.) K. Becker Wedelia parviceps Blake Zexmenia epapposa M. E. Jones Zexemia hispida var ramosissima (Greenm.) Greenm. Phytologia (Aug 2004) 86(2) 93 Sin, Nay, Jal, Mic, Mex, Ver, Gue, Oax, Tab, Cps, Yuc, Cam, Qui and Guatemala southwards, 0-1300 m; all seasons. Perennial suffruticose herbs or shrublets 0.5-3.0 m high; stems stiffly erect and divaricate to arching or sprawling; leaves opposite, 3-10 cm long, 1-4 cm wide; petioles 0-6 mm long; blades ovate to elliptic- lanceolate, 3-nervate, the undersurface with stout, erect or ascending hairs and below these another layer of rather short, weak, hairs with recurved apices; heads radiate, terminal, 1-several on ultimate peduncles 1-15 cm long; involucres 5-15 mm high, 2-3 seriate, the outer series shorter or longer than the inner; ray florets 5-11, the ligules, 5-14 mm long; disk florets 10-40; chromosome numbers, n= ca 11, 12 and 23 pairs. A widespread highly variable species, presumably divisible into several infraspecific geographical varieties. The larger-headed, longer- peduncled, more prominently shrubby or divaricately-branched plants from Jal to Cps are perhaps properly called var. scabra, regardless of the disposition of yet other taxa, although Strother (1991) adopted the name W. acapulcensis for the entire complex, feeling that the earlier name, W. scabra, was inadequately typified for his nomenclatural purposes. I can not agree with Strother’s widely inclusive treatment, especially his submergence of the well-known, long-established, W. hispida, as a varietal element of W. acapulcensis. He also recognized the varieties parviceps and ramosissima, which appear to be reasonably organized populational phases, having mostly smaller heads on shorter peduncles. Strother cited and mapped two collections of W. s. var. ramosissima as occurring in Tam and San. I include both of these in my concept of W. ayerscottiana. WEDELIA SIMSIOIDES McVaugh, Contr. Univ. Michigan Herb. 9: 462. 1972. Known only by collections from s Nay, marshy meadows, oak zone, 800-900 m; Aug-Sep. 94 Phytologia (Aug 2004) 86(2) Perennial rhizomatous herbs to | m high; leaves opposite throughout, 6-10 cm long, 3-7 cm wide; petioles markedly winged, 1-2 cm long; blades ovate to deltoid, 3-nervate, roughly hispid, the margins irregularly serrate, lacerate or lobe. A very distinct species, presumably closest to W. cordiformis. McVaugh (1984) provides an excellent illustration of the taxon. WEDELIA STRIGOSA Hook. & Arn., Bot. Beechy Voy. 435. 1841. Aspilia strigosa (Hook. & Arn.) Benth. ex Hemsl. Jal, Col, Mic, and Gue, lower Pacific slopes in tropical deciduous forests, 1-300 m; Jun-Jan. Much resembling W. scabra but the vestiture of the leaves without an understory of short uncinate hairs, and the ray florets neuter (vs pistillate). Strother (1991) recognized this taxon, correctly noting its relationship to W. acapulcensis (= W. scabra of the present treatment), the latter a widespread, mostly higher montane element in Mexico; W. strigosa is mostly confined to the lower montane slopes from near sea to ca 300 m, as noted by Strother. In the vicinity of Acapulco plants seemingly intermediate to W. strigosa and W. scabra occur, possessing uncinate hairs, but these not part of a smaller understory of smaller hairs as occurs in W. scabra. WEDELIA TALPANA B. L. Turner, Phytologia 72: 117. 1992. Jal, vicinity of El] Tuito (Mpio. Talpa), 900-1800 m; Nov-Mar. Brittle-stemmed shrubs or shrublets 30-50 cm high; resembling W. rosei but having smaller leaves (mostly 2-3 cm long vs 3-8 cm) and heads sessile or nearly so. Phytologia (Aug 2004) 86(2) 95 Strother (1991) called attention to some of the plants which comprise this taxon, noting that these might be deserving of specific recognition. Additional collections from the same region, in my opinion, has confirmed this observation. WEDELIA TEHUANTEPECANA B. L. Turner, Phytologia 65: 355. 1988. Wedelia acapulcensis var. tehuantepecana (B. L. Turner) Strother e Oax, s Ver, and Cps, tropical deciduous forests, 50-700 m; Jul- Nov. Much-branched erect or procumbent shrubs or shrublets 0.5-1.5 m high, leaves opposite, 4-10 cm long, 1.5-4.0 cm wide; petioles 0.5-1.5 cm long, the upper portion winged; blades ovate to subdeltoid, 3-nervate, coarsely hispid, uncinate hairs absent, the margins serrulate to coarsely and irregularly dentate; heads radiate, mostly 1-3 in terminal cymes, the ultimate peduncles mostly 2-8 cm long; involucres 10-12 mm high, narrowly campanulate, 2-3 seriate, the bracts subequal, the outer series greener and somewhat longer than the inner; ray florets 5-11, sterile (rarely fertile), the ligules 5-10 mm long; achenes, those of the ray, with a raised crown bounded by 3 bristles, those of the disk 5-6 mm long with a raised crown and usually 2 or 3 bristles 3-6 mm long, the margins markedly winged at maturity; n= ca 26 pairs. Closely related to W. scabra but lacking the fine, uncinate, indument on the under-surface of the leaves of that species and possessing mostly larger heads with longer achenes with raised crowns having bristles 3-6 mm long. WEDELIA TRILOBATA (L.) Hitche., Rep. Missouri Bot. Gard. 4: 99. 1893. Complaya trilobata (L.) Strother Sphagneticola trilobata (L.) Pruski Thelechitonia trilobata (L.) H. Rob. & Cuatrecasas 96 Phytologia (Aug 2004) 86(2) Ver, Oax, Tab, Cps, Yuc, Cam, Qui and Guatemala southwards, widely cultivated in Mexico and elsewhere as a ground cover and often escaping, 0-200 m; all seasons. Easily recognized by its prostrate habit and long-trailing stems which root at the nodes; leaves often 3-lobed; relatively large, axillary or terminal, yellow-orange heads; chromosome numbers, n = 20-30 pairs according to Strother (1991); Koyama (1985: Acta Phytotax. Geobot. 36:171) reports a count of 2n = 56, or 23 pairs. This species was treated as belonging to the genus Sphagneticola by Strother (who used the generic name Complaya, a synonym of Thelechitonia, as noted by Robinson and Cuatrecasas, 1992, the latter proving to be synonymous with Sphageticola, cf. Pruski 1996). WEDELIA VEXATA Strother, Syst. Bot. 33: 67. 1991. Nay and Jal, Pacific slopes, 0-100 m; Jul-Aug. Much-resembling W. scabra but distinguished by its linear-elliptic to lance-elliptic, sessile leaves, with nearly clasping, blades. Strother (1991) was clearly vexed by this taxon, hence its name. It has the pistillate ray florets and vestiture of W. scabra, but differs markedly in having more elongate, ellipitcal or lance-elliptic, nearly sessile leaves. Wedelia vexata might also be compared with W. strigosa, the latter possessing neuter ray florets and, in the region where it apporoaches W. vexata (Jal), non-uncinate hairs. WEDELIA XYLOPODA (Greenm.) B. L. Turner, comb. nov. Based upon Aspilia xylopoda Greenm., Proc. Amer. Acad. Arts 40: 40. 1904. Nay, s Dur and closely adjacent Sin; pine-oak woodlands, ca 2000 m; Aug-Nov. Phytologia (Aug 2004) 86(2) 97 Much resembling W. rosea but semiprostrate shrublets having heads with lavender or purple rays and involucres 2-seriate, the outer series as long as or longer than the inner series, not at all graduate. McVaugh (1984) noted that this taxon “may be merely a purple- flowered form of Wedelia rosei,” although he retained it as Aspilia xvlopoda. | know the species only by the type and several additional collections (LL, TEX) from along hiway 40 at the Sin-Dur border. Strother (1991) included this species within his broad concept of W. rosea. 98 Phytologia (Aug 2004) 86(2) OBSERVATIONS ON PARNASSIA GRANDIFOLIA DC. (SAXIFRAGACEAE) IN THE WEST GULF COASTAL PLAIN Michael H. MacRoberts and Barbara R. MacRoberts Bog Research Shreveport LA 71104 and Herbarium, Museum of Life Sciences Louisiana State University in Shreveport Shreveport LA 71115 Lynn Stacey Jackson U.S.D.A. Forest Service Kisatchie National Forest Winn Ranger District Winnfield LA 71483 ABSTRACT We surveyed for and monitored Parnassia grandifolia, a West Gulf Coastal Plain rare and imperiled species with a disjunct distribution. KEY WORDS: Parnassia grandifolia, pitcher plant bog, Louisiana, floristics. INTRODUCTION Parnassia grandifolia, a southeastern obligate wetland endemic, is of interest because it is globally rare and a disjunct species raising the biogeographical question as to whether extant populations are relicts from a once continuous distribution or are recent long distance colonizers, or recolonizers. Phytologia (Aug 2004) 86(2) 99 In a previous paper, we reviewed the status of Parnassia in the West Gulf Coastal Plain (MacRoberts etal. 1997; see also McDaniel etal. 1997). In this paper, we describe our subsequent observations. The purpose of our recent work was threefold: first, search for additional populations of P. grandifolia in Louisiana, second, monitor those populations, and third, compare the ecology and morphology of Louisiana populations to populations elsewhere in the region. We, therefore, surveyed baygalls and bogs in promising habitat in and around our original discovery site. We traveled to Mississippi and Oklahoma to examine Parnassia habitat in those places and examined herbarium specimens. Botanical names follow Kartesz and Meacham (1999) and authorities can be read in that work. LOUISIANA SURVEYS We concentrated --- but did not confine --- our surveys to the vicinity of the site of our first discovery of the species on the Winn Ranger District, Kisatchie National Forest, in Natchitoches Parish. The original population (Site A), found on 21 March 1997, is described in MacRoberts et al. 1997. After this discovery, we found two additional sites, both close to the original site. Site B, found on 26 July 1999, is about 100 meters west of Site A and about 15 meters east of a pitcher plant bog on a hillside just a few meters above a baygall. This population consisted of three small clumps that varied from 40 to 50 leaves in a 1 meter square area between 1999 and 2003. Associated species included Acer rubrum, Athyrium filix-femina, Callicarpa americana, Cornus florida, Dichanthelium sphaerocarpon, Magnolia virginiana, Mitchella repens, Morella caroliniensis, Nyssa sylvatica, Persea palustris, Pinus palustris, Melanthium virginicum, Osmunda_ cinnamomea, O. regalis, Rhododendron oblongifolium, Sassafras albidum, Smilax rotundifolia, Woodwardia areolata, and Viburnum nudum. 100 Phytologia (Aug 2004) 86(2) Site 1997 1998 1999 2000 2001 2002 2003 A 32 72 96 6 24 tS 30 B - - 1 1 2 0 3 C - - 0 0 0 0 0 Table 1. Number of flowering scapes produced by three “populations” of P. grandifolia in the Kisatchie National Forest. Site C, found on 26 March 1999 in a pitcher plant bog, (see MacRoberts et al. 2002 for a description of this bog) consisted of only 22 leaves in a | meter square area. This population was about 350 meters west of Site A. In May 2000, it had 29 leaves, but by September 2000, after a severe drought, the site was dry and all above ground vegetation had disappeared. In November 2001, one leaf appeared; it was not found at this location in 2002 or 2003. Associated vegetation included Acer rubrum, Aletris aurea, Alnus serrulata, Dichanthelium spp., Eriocaulon decangulare, Eupatorium fistulosum, Gelsemium sempervirens, Helianthus angustifolius, Liatris pycnostachya, Magnolia virginiana, Melanthium virginicum, Méitreola_sessilifolia, Morella caroliniensis, Nyssa sylvatica, Osmunda_ regalis, Persea palustris, Ptilimnium sp., Rhododendron oblongifolium, Sarracenia alata, Scleria ciliata, Smilax laurifolia, Toxicodendron vernix, Viburnum nudum, and Viola primulifolia. All three sites occur on Betis mildly acidic (4.8-5.4 pH) low-nutrient loamy fine sands on north and west-facing 5-20 degree slopes (Martin et al. 1990). All three sites have between 90% to 100% shrub/canopy cover with filtered light. We monitored flowering --- which occurs in late October and early November --- between 1997 and 2003 (Table 1). A severe drought in 2000 dried out all sites resulting in the collapse of flowering at Site A. The extent of vegetation, however, was not affected except at Site C as described above. Phytologia (Aug 2004) 86(2) 101 MISSISSIPPI AND OKLAHOMA POPULATIONS In order to get a better picture of habitat for this species, we made trips to southern Mississippi on 4 May 1999 and to southern Oklahoma on 26 September 2000 to visit known populations. In Mississippi, P. grandifolia grows in three types of pitcher plant bog habitat: seepage bogs, springhead seepages, and muck or quaking bogs (see Eakes 1989; Folkerts 1991). Associates include Arnoglossum ovatum, Chaptalia tomentosa, Cyrilla racemiflora, Drosera spp., Eriocaulon compressum, Ilex coriacea, Lycopodiella alopecuriodes, Magnolia virginiana, Morella caroliniensis, Rhynchospora macra, Sabatia macrophylla, Sarracenia alata, S. psittacina, Smilax laurifolia, Sphagnum sp., and Toxicodendron vernix. Parnassia grandifolia grows in both open areas and in various degrees of shade. Some of the populations we observed were growing in total or near-total sunlight. There is only one known P. grandifolia site in Oklahoma. The plants were first found by Taylor and Taylor (1978) in an “upland marsh” in northern Choctaw Co. We found them growing in full sun at the edge of the marsh in permanently wet mucky soils surrounded by dense marsh vegetation. Associates included: Boehmeria cylindrica, Conoclinium coelestinum, Eupatorium perfoliatum, Helianthus angustifolius, H. grosseserratus, H. mollis, Helenium autumnale, Juncus sp., Melanthium virginicum, Rudbeckia fulgida, R. hirta, Saccharum giganteum, Salix sp., Solidago sp., Sorghastrum nutans, Typha sp., and Woodwardia areolata. What was interesting about the Oklahoma population was that not only were they growing in full sunlight, but they were much more robust than other populations we had seen and were in flower in mid-September, fully two months earlier than the Kisatchie National Forest P. grandifolia populations. This early blooming was also the case in 1976 when the species was first collected at this site by John Taylor. We measured scape height for comparison with Natchitoches Parish populations. Eight scapes for Oklahoma plants ranged from 35 cm to 60 cm and averaged 102 Phytologia (Aug 2004) 86(2) 49 cm. Fifteen scapes for Kisatchie National Forest plants ranged from 26 cm to 50 cm and averaged 39 cm. DISCUSSION Being a disjunct and scattered species in the West Gulf Coastal Plain, P. grandifolia presents an interesting subject for biogeographic studies. Do these scattered populations represent remnants of once continuous populations clinging on after the Ice Age or are they recent long distance colonizers or re-colonizers (see map in MacRoberts et al. 1997)? While no one has assessed genetic and morphological variation within and between West Gulf Coastal Plain populations, we noted several striking variations among populations including flowering period, habitat preference, and size of plants. These variations may indicate significant genetic differences evolved in isolation. Parnassia grandifolia is not the only member of this genus to have disjunct populations in our region: P. asarifolia is also disjunct (McDaniel et al. 1997; MacRoberts et al. 1997). Thus, the genus might serve as an interesting subject for genetic study, with the ultimate aim of testing different biogeographic hypotheses as Watson et al. (2002) has recently done for Eriocaulon koernickianum. The results of these studies, in turn, would have implications for conservation priorities and for understanding the development of the West Gulf Coastal Plain flora. ACKNOWLEDGMENTS The research was supported by National Forest Foundation Cost- share Agreement No. 08-99-06-CCS-008. Steve Leonard and Julia Moore, The Nature Conservancy, Camp Shelby, showed us Mississippi sites. Connie Taylor aided with relocating the Oklahoma site. Jason Singhurst and Eric Keith commented on an earlier version of the paper. LITERATURE CITED Eakes, C.B. 1989. Studies on the life history and habitat of the Bogstar (Parnassia grandifolia DC.) in south Mississippi. MS Thesis, University Southern Mississippi, Hattiesburg, Mississippi. Phytologia (Aug 2004) 86(2) 103 Folkerts, G.W. 1991. A preliminary classification of pitcher plant habitats in the southeastern United States. J. Alabama Acad. Sci. 62:199- 224. Kartesz, J.T. and C.A. Meacham. 1999. Synthesis of North American Flora. Version 1.0. North Carolina Botanical Garden, Chapel Hill, North Carolina. MacRoberts, B.R., M.H. MacRoberts, L.M. Stacey, and D.C. Moore. 1997. The status of Parnassia (Saxifragaceae) in the West Gulf Coastal Plain. Phytologia 83:53-57. MacRoberts, B.R., M.H. MacRoberts, and L.S. Jackson. 2002. Floristics and management of pitcher plant bogs in northern Natchitoches and Winn parishes, Louisiana. Proc. Louisiana Acad. Sci. 64:14-21. Martin, P.G., C.L. Butler, E. Scott, J.E. Lyles, M. Marino, J. Ragus, P. Mason, and L. Schoelerman. 1990. Soil Survey of Natchitoches Parish, Louisiana. U.S.D.A. Soil Service, Washington, D.C. McDaniel, D.L., D.L. Marsh, and W.D. Crank. 1997. Rediscovery of Parnassia asarifolia (Parnassiaceae) in Arkansas. J. Arkansas Acad. of Sci. 51:208-209. Taylor, R.J. and C.E. Taylor. 1978. Additions to the vascular flora of Oklahoma. Sida 7:361-368. Watson, L.E., A.B. Kornkven, C.R. Miller, J.R. Allison, N.B. McCarty, and M.M. Unwin. 2002. Morphometric and genetic variation in Eriocaulon koernickianum Van Heurck & Muller-Argoviensis (Eriocaulaceae): a disjunct plant species of the southeastern United States. Castanea 67:416-626. 104 Phytologia (Aug 2004) 86(2) TAGETES ERECTA (ASTERACEAE), A NEW ESCAPE TO THE FLORA OF TEXAS Eric L. Keith Raven Environmental Services, P.O. Box 6482, Huntsville, Texas 77342, U.S.A. eric.keith@excite.com ABSTRACT Tagetes erecta, known under several common names including common or Aztec marigold, is reported as escaped from cultivation in Texas for the first time. KEY WORDS: Tagetes, Asteraceae. Tagetes erecta L., common or Aztec marigold, has been reported as escaped from cultivation in several states to the east and west of Texas including the adjacent states Oklahoma, Arkansas, and Louisiana (Cronquist 1980, Gandhi and Thomas 1989, Nature Serve 2004, Radford et. al. 1968, USDA 2004). Until now, the species has never been reported escaped in Texas (Correll and Johnston 1970, Hatch et. al. 1990, Jones et. al. 1997, Kartesz and Mecham 1999, Nesom and Brown 1998, Slona- Nelson 1996, Turner et. al. 2002). Approximately 20 plants were found on black clayey soil in a weedy lot recently cleared for construction. The flowers were in shades of both orange and yellow. These plants may be the result of direct seeding or contaminants in grass seeds. However, no other unusual weedy or commonly cultivated species were detected in the area. A key to the commonly escaped species of Tagetes can be found in Cronquist (1980), Gandhi and Thomas (1989), and Radford et. al. (1968). Voucher specimen: Walker Co.: clayey disturbed area ca. 100 m, W of the corner of Hwy 75 and 10" St., 28 May 2004, Keith 701 (SBSC, SHST). Phytologia (Aug 2004) 86(2) 105 ACKNOWLEDGMENTS Thanks to Larry Brown of Houston Community College for verifying the identity of this species and to Larry and Jason Singhurst, Texas Parks and Wildlife, for a critical review of the paper. LITERATURE CITED Correll, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, TX. Cronquist, A. 1980. Vascular flora of the Southeastern United States, Volume I: Asteraceae. The University of North Carolina Press. Chapel Hill, N.C. Gandhi, K. N. and R. D. Thomas. 1989. Asteraceae of Louisiana. Sida Botanical Miscellany No. 4. SMU Herbarium, Dallas. Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1990. Checklist of the vascular plants of Texas, MP-1655. Texas Agric. Exp. Sta., College Station. Jones, S.D., J.K. Wipff, and P.M. Montgomery. 1997. Vascular plants of Texas. Acomprehensive checklist including synonymy, bibliography, and index. University of Texas Press. Austin. Kartesz, J.T. and C.A. Mecham. 1999. Synthesis of the North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill, North Carolina. NatureServe. 2004. NatureServe Explorer: An online encyclopedia of life [web application]. Version 3.1. NatureServe, Arlington, Virginia. Available http://www.natureserve.org/explorer. (Accessed: May 29, 2004 ). 106 Phytologia (Aug 2004) 86(2) Nesom, G. L. and L. E. Brown. 1998. Annotated checklist of the vascular plants of Walker, Montgomery, and San Jacinto Counties, East Texas. Phytologia 84: 107-153. Radford, A. E., H. E. Ahles, C. R. Bell. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press. Chapel Hill, N.C. Slona-Nelson, K. B. 1996. Floristic study of Walker County, Texas: Asteraceae. Masters of Arts, Sam Houston State University. Turner, B.L., H. Nicholls, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Vol. 1. Sida, Botanical Miscellany 24. Botanical Research Institute of Texas. USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (http://plants. usda.gov). National Plant Data Center, Baton Rouge, LA 70874- 4490 USA. Phytologia (Aug 2004) 86(2) 107 PRENANTHES BARBATA (ASTERACEAE) AND PARENTUCELLIA VISCOSA (SCROPHULARIACEAE) NEW TO OKLAHOMA Jason R. Singhurst Wildlife Diversity Program, Texas Parks and Wildlife Department, Austin, Texas 78704 U.S.A. Jason.Singhurst@tpwd.state.tx.us Walter C. Holmes Department of Biology, Baylor University Waco, Texas 76798-7388 U.S.A. Walter Holmes@Baylor.edu ABSTRACT Prenanthes barbata and Parentucellia viscosa are reported as new to the flora of Oklahoma. KEYWORDS: Prenanthes,Parentucellia,Asteraceae, Scrophulariaceae, Oklahoma. The following taxa are not reported from Oklahoma by Taylor and Taylor (1994) or the PLANTS database (USCS, USDA. 2004), thus are considered as new to the state. All locations cited are in the West Gulf Coast Plain vegetational region (Taylor and Taylor 1994). Prenanthes barbata (Torr. & A. Gray) M. L. Milstead ex A. Cronquist (Asteraceae). Four populations of P barbata were located on a stream terrace along Mt. Fork Creek below Beaver Bend Lake and Mt. Fork River within Beaver Bend State Park in Mc Curtain County. The Mt. Fork Creek plants consisted of two populations, one with about 25 flowering individuals and another with about 15 flowering individuals. Two Mt. Fork River populations consisted of about ten flowering individuals at one site and five flowering individuals at the other 108 Phytologia (Aug 2004) 86(2) site. The dominant tree species at the sites included: Acer rubrum, A. saccharum, Liquidambar styraciflua, Quercus alba, QO. muhlenbergii, Q. shumardii and Vaccinium elliottii. Common herbaceous plants included Chasmanthium sessilifolium, Solidago caesia, and Carex spp. These populations are about 200 km west of the nearest known occurrence in Arkansas in Grant Co. (USDA, NRCS. 2004) and approximately 100 km northeast of the nearest known Texas occurrence in Lamar Co. (Singhurst et al., 2004). Voucher specimens: OKLAHOMA. Me Curtain Co.: Beaver Bend State Park, Mt. Fork Creek trail below Beaver Bend Lake, 0.7 mi. S of spillway, 6 Oct 2000, Singhurst 10118 (BAYLU); Beaver Bend State Park, park dam trail, 0.4 mi. S. of dam, 6 Oct 2000, Singhurst 10115 (BAYLU); Beaver Bend State Park, W side of Mt. Fork River below dam, 20 Oct 2002, Singhurst & Singhurst 11343 (BAYLU); Beaver Bend State Park, W side of Mt. Fork River below Broken Bow Reservoir dam, 19 Oct 2003, Singhurst & Singhurst 12355 (BAYLU); Beaver Bend State Park, W side of parking lot turn around at trail entrance of park dam, 19 Oct 2003, Singhurst & Singhurst 12534 (BAYLU). Two other species of Prenanthes occur in Oklahoma, P. altissima L. and P. alata (Hook.) D. Dietr. (Milstead 1964). Prenanthes altissima is distinguished by its yellow to green-yellow florets numbering about five or six per capitulum. Prenanthes alata, which has a similar flower color and floret number to P. barbata, is characterized by its triangular to occasionally elliptic basal leaves that are slightly hastate at the base. Prenanthes barbata has oblanceolate to elliptic basal leaves that are never hastate. Prenanthes barbata is considered to be rare in Texas (Correll and Johnston 1970), and since it is now known from only four populations at Beaver Bend State Park, it should be considered as a candidate for sensitive species status by the Oklahoma Natural Heritage Inventory. Parentucellia viscosa (L.) Caruel (Scrophulariaceae). This Old World native has been naturalized on the west coast (California, Oregon, and Washington) and in the south central (Arkansas, Louisiana, Mississippi, Phytologia (Aug 2004) 86(2) 109 and Texas) United States (USDA, NRCS. 2004). The species is rather widespread in east Texas and is quite common in adjacent Bowie and Red River counties (pers. observ.), bordering the Red River. The species is conspicuous because of its bright yellow flowers that have earned it the common name Yellow Glandweed. This species typically occurs in mesic roadsides in association with Vicia spp., Rumex spp., Lolium perenne, Casteleja spp., Eleocharis spp., etc. Voucher specimen: OKLAHOMA. Me Curtain Co.: 11.2 miles north of Junction of Red River and OK Hwy 37, May 1999, Singhurst 10252 (BAYLU). ACKNOWLEDGEMENTS We wish to thank Oklahoma State Parks for permitting access to Beaver Bend State Park. We are also grateful to John Pruski of the Missouri Botanical Garden and Eric Keith of Raven Environmental Services, Huntsville, Texas, for review of the manuscript. LITERATURE CITED Correll, D. S. and M. C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. Milstead, W. L. 1964. A revision of the North American species of Prenanthes. Unpub. dissertation, Purdue University. Singhurst, J. R., R. J. O’Kennon, and W. C. Holmes. 2004. The genus Prenanthes (Asteraceae: Lactuceae) in Texas. Sida 21: 181-191. Taylor R. J. and C. E. Taylor 1994. An annotated list of the ferns, fern allies, gymnosperms and flowering plants of Oklahoma. Southeastern Oklahoma State University, Durant. USDA, NRCS. 2004. The PLANTS database (MACROBUTTON HtmlResAnchor http://plants.usda.gov/plants). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. 110 Phytologia (Aug 2004) 86(2) REDISCOVERY OF CYPERUS CEPHALANTHUS (CYPERACEAE) IN TEXAS David J. Rosen and Bradley J. Christoffersen U.S. Fish and Wildlife Service 17629 El Camino Real, Suite 211 Houston, TX 77058-3051, U.S.A. Brownand Elsik (2002) reported collections of Cyperus cephalanthus Torr. & Hook. from a small coastal prairie remnant in Harris County, apparently the first record in Texas since the type specimen was collected near Galveston Bay (Torrey 1836; Carter and McInnis 1993). Following their report, the site was excavated and formed into a detention basin. Since the type location has never been relocated and presumed destroyed, the only known extant population of C. cephalanthus in Texas at the time was extirpated. Recently, during floristic surveys of the Nash Prairie in Brazoria County, a flourishing population of C. cephalanthus comprising over 50 individuals was discovered. Plants were growing in saturated, fine sandy loam soils in the nearly level fringes of a deep prairie depression. The Nash Prairie has been managed for decades by routine mowing, haying, and burning, preventing the encroachment of Sapium sebiferum (L.) Roxb. into these seasonal wetlands, the preferred habitat for C. cephalanthus. Tucker et al. (2003) suggested that Cyperus cephalanthus is naturalized in the United States, largely based on its uncertain relationship with C. Jaetus Presl., a similar South American species. However, Carter and McInnis (1993) provide detailed ecological, biological, and historical information suggesting C. cephalanthus is native to the coastal prairies of Texas and Louisiana. Although its nativity may remain unknown, Cyperus cephalanthus, like C. drummondii Torr. & Hook., is a reliable indicator of relatively undisturbed coastal prairie remnants in Texas and Louisiana (Carter and McInnis 1993; Carter et al. 1999; Grace et al. 2000; Rosen 2004). Phytologia (Aug 2004) 86(2) 111 Voucher specimen: U.S.A. TEXAS. Brazoria Co.: Nash Ranch, 275-acre hay meadow W of CR 25, about 8.7 miles N of its intersection with Hwy. 35 in West Columbia, locally common in seasonally wet fringes of deep prairie depression with Amsonia repens Shinners, Boltonia diffusa Ell., Carex festucacea Schkuhr ex Willd., C. longii Mack., C. triangularis J. B6ck., Cyperus drummondii Torr. & Hook., C. haspan L., C. pseudovegetus Steud. var. pseudovegetus, Diodia virginiana L., Eleocharis palustris (L.) Réem. & Schult., Hydrolea ovata Nutt. ex Choisy, Hymenocallis liriosme (Raf.) Shinners, Juncus coriaceus Mack., J. nodatus Coville, Leersia hexandra Sw., Ludwigia glandulosa Walter, Panicum hians Ell., Panicum subg. Dichanthelium spp., P. virgatum L., Paspalum plicatulum Michx. var. plicatulum, Physostegia intermedia (Nutt.) Engelm. & A. Gray, Polygonum hydropiperoides Michx., Rhynchospora caduca Elliott, R. corniculata (Lam.) A. Gray, Rudbeckia nitida Nutt. var. texana Perdue, Schizachyrium scoparium (Michx.) Nash var. scoparium, Tridens strictus (Nutt.) Nash, and Vernonia missurica Raf., 21 May 2004, Rosen and Christoffersen 2950 (BRIT, MICH, VSC, TEX). ACKNOWLEDGMENTS We are grateful to Peter Conaty and John Phillips for granting access to the Nash Prairie. We thank Richard Carter and Charles Bryson for critical review of this manuscript. LITERATURE CITED Brown, L. E. and I. S. Elsik. 2002. Notes on the flora of Texas with additions and other significant records. II. Sida 20:437-444. Carter, R. and N. McInnis. 1993. Status report on Cyperus cephalanthus. Unpublished report submitted to U.S. Fish & Wildlife Service, Jackson, Mississippi. 140 pp. Carter, R., D. K. Alexander, C. T. Bryson, and A. Lazari. 1999. The taxonomy of Cyperus virens and Cyperus drummondii in the Southeastern United States. Sida 18:1049-1063. 112 Phytologia (Aug 2004) 86(2) Grace, J. B., L. Allain and C. Allen. 2000. Vegetation associations in a rare community type - coastal tallgrass prairie. Plant Ecology 147:105-115. Rosen, D. J. 2004. Noteworthy collections of Cyperus drummondii (Cyperaceae) from Texas. Sida 21:495-497. Torrey, J. 1836. Monograph of North American Cyperaceae. Ann. Lyceum Nat. Hist. New York 3:238-448. Tucker, G. C., B. G. Marcks, and J. R. Carter. 2003. Cyperus. In: Flora of North America Editorial Committee, eds. Flora of North America North of Mexico, Vol. 23. Oxford Univ. Press, New York. Pp. 141-191. Phytologia (Aug 2004) 86(2) 113 A NEW COMBINATION IN TEXAS CROTON (EUPHORBIACEAE) B. L. Turner Plant Resources Center University of Texas Austin, TX, 78712-0471, USA Croton thermophilus (M.C. Johnst.) B. L. Turner, comb. nov. Based on C. corymbulosus Engelm.var. thermophilus M. C. Johnst., Southw. Naturalist 3: 187. 1959. Jackie Poole (Botanist, Tx. Parks and Wildlife, pers. comm.) called to my attention that in the Atlas of the Vascular Plants of Texas (Turner et al. 2003; Sida, Bot. Misc. 24) I treated the above taxon as a species, but neglected to make the appropriate format combination in the text itself. This is corrected here. Johnston originally described this taxon as a variety of C. corymbulosus Engelm., but latter transferred this to the earlier name C. pottsii (K1.) Muell. Arg., a widespread, relatively variable, weedy species of the Southwestern U.S.A. and Northern Mexico. As well noted by Poole (Guide to the Rare Plants of Texas, in prep.), C. thermophilus is readily distinguished by its usually markedly fractiflex stems, mostly smaller leaf blades with obtuse apices, and yet other characters. While both taxa may occur in relatively close proximity on southeastern Brewster County, I have not observed intergrades between these, either in the field or in the herbarium (LL, SRSC, TEX), nor have such been noted by yet others 114 Phytologia (Aug 2004) 86(2) SUBSCRIPTION INFORMATION PHYTOLOGIA is published three times a year, P.O. Box 2012, Sam Houston State University, Huntsville, TX 77341-2012. Annual domestic individual subscription (3 issues): $30.00. Annual domestic institutional subscription (3 issues): $90.00. Foreign and/or airmail postage extra. 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. Authors should arrange for two workers in the appropriate field to review the manuscript before submission. Copies of reviews should be forwarded to the editor with the manuscript. Additional review may occur for articles proposing significant nomenclatural changes and or revisions to established taxa. Effort will be made to locate a reviewer for author(s) if he/she desires. Articles should be submitted digitally, either by disk, CD or email. Language of manuscripts may be either English or Spanish. Format of articles should follow past issues of PHYTOLOGIA (preferably single spaced and Times New Roman in 10 point font.) Cost of publication is currently $13.00 US per page for publication without reprints. PDF files of reprints will be made available to authors upon request. Page charges are due with manuscripts. No extra charges are made for line drawings or grey scale photos. Extra fees will be charged for color photos, the charge will be determined on an individual basis. Phytologia (Aug 2004) 86(2) 115 INDEX TO NEW NAMES AND COMBINATIONS IN PHYTOLOGIA 86(2) Taxa Page Number Blossfeldieae Crozier, trib. nov. 56 Blossfeldioideae Crozier subfam. nov. 55 Croton thermophilus (M.C. Johnst.) B. L. Turner, comb. nov. 113 Galactia volubilis (L.) Britt. var. baltzelliana D. B. Ward & D. W. Hall, var. nov. 68 var. fasciculata (Vail) D. B. Ward & D. W. Hall, comb. nov. 69 Juniperus deltoides R. P. Adams, sp. nov. 47 Juniperus polycarpos K. Koch var. turcomanica (B. Fedtsch) R. P. Adams, comb. nov. 50 Wedelia elottiana B. L. Turner, sp. nov. 84 Wedelia juxtlahuacana B. L. Turner, sp. nov. 88 Wedelia xylopoda (Greenm.) B. L. Turner, comb. nov. 96 hk ra MITHSONIAN INSTITUTION LIBRARIES wii 88 0148 Pata eyy eee Vr ees Pa mobi athe bjkek ed et rere