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SPERLING: C: R. ‘New species and | new combinations in Anredera Juss.
oe (Basellaceae)... eee hoy Stier Cenyey ce teens cwache uss: ae TURNER, B. L., Stevia ealeadana (Asteraceae) a new species from Oaxaca, Bee VIN re ate ei yooe ehedeg esse ears oO TURNER, B. Lo Mensdora gypsophila (Ceaceae), a new oer from near . Galeana, Nuevo Leén, México........:. ere re oer eee Rene nen eo mean lae 8 AXELIUS, B., A new combination i in Physalis (Solanaceae). eeas beer acne 10 TURNER, B oS Taxonomic overview: of Hedyotis nigricans (Rubiaceae) and «closely allied TAM ee Gees cs oer eee pe ee isis pec ene ee 12 HUNT, D.M., M:H- MACROBERTS. & BR. ‘MACROBERTS, The status of Quercus arkansana Sarg. (Fagaceae) in Texas. ......:.......c..se0e ee ees ee 22 HERRERA A., Y., Chromosome numbers report... ............ Age tree Wl eerer 325 MORDEN, C. W.., ‘A new combination i in Muhlenbergia Panceses iene 28 TURNER, B.L., Sedum booleanum (Crassulaceae), a new red- flowered species from Nuevo. Ret MExicOc 2 2 Oe a eee ee YAHARA, T. & A. SOEJIMA, ve new species of Stevia on Miexice oe Giang TURNER, B.L., Paronychia hintonioram (Caryophyliacesc). a new. species _.. from Nuevo Leon and Verdcruz,, MEXICO.) 2 fois i es fe Gare 38 TURNER, B.L., A-new SPECIE of Pittocaulon (Asteraceae, ‘Senecioneae) from - Oaxaca, México. eee ae ne oe gee Ce ee ae ear eat ees: TURNER, B.L., Two new varieties of Hedeoma palmeri A aimaceae) from ~ — northeastern Medes oe 47
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Phytologia (July 1995) 79(1):1-4.
NEW SPECIES AND NEW COMBINATIONS IN ANREDERA JUSS. (BASELLACEAE)
Calvin R. Sperling’
National Germplasm Resources Laboratory, Room 402, Building 003, Barc-West, Beltsville, Maryland 20705 U.S.A.
ABSTRACT
Two new species of Anredera, A. aspera Sperling and A. densiflora Sperling, are described, and four new combinations, A. brachystachys (Mog.) Sperling, A. floribunda (Mogq.) Sperling, A. krapovickasii (Villa) Sperling, and A. tucumanensis (Lillo & Hauman) Sperling, are made. These new species and new combinations are from the unpublished Ph.D. dissertation of Calvin R. Sperling.
KEY WORDS: Anredera, Basellaceae, taxonomy
FORWARD
[J. W. Nowicke, Botany Dept., NHB 166, Smithsonian Institution, Washington D.C. 20560 U.S.A.]
In the course of a palynological study that included Anredera, | discovered that two new species and four new combinations established by the late Calvin Sperling (1987) are heretofore unpublished. The Latin descriptions of the two new species, Anredera densiflora from Ecuador and Peru, and A. aspera from northern Bolivia, and the synonymy of the remaining four new combinations, A. tucumanensis (Lillo & Hauman) Sperling, A. floribunda (Moq.) Sperling, A. krapovickasii (Villa) Sperling, and A. brachystachys (Mogq.) Sperling, have been taken from Spelling’s dissertation. Before his death, he approved publication of these names.
Two more new combinations, Anredera diffusa (Mogq.) Sperling and A. marginaia (H.B.K.) Sperling, were recently published (Brako & Zarucchi 1993, p. 1253). In addition to the twelve species of Anredera, Sperling treated the remaining three genera that comprise Basellaceae, Basella L. (5 species), Tournonia Mog. (1 species), and Ullucus Caldas (1 species). Although his study concentrated on Ullucus tuberosus Caldas, Spelling’s dissertation has keys, descriptions, distribution maps, and
' Deceased 20 May 1995.
p PHYTOLOGIA July 1995 volume 79(1):1-4
discussions for the remaining eighteen species. His revision of Basellaceae as a family is the first since 1849, when it was treated by Moquin-Tandon. Spelling’s discussions of relationships among genera and species provided new information that was integrated with the pollen data (Nowicke, in press). It is unfortunate that his dissertation has not been published in its entirety.
The species are taken up in the same sequence as they are in the dissertation. Abbreviations for authors follow Brummitt & Powell (1992).
Anredera Juss.
ANREDERA FLORIBUNDA (Mog.) Sperling, comb. nov. BASIONYM: Boussingaultia floribunda Mog. in DC., Prodr. 13(2):229. 1849. TYPE: COLOMBIA. Ibaque, Goudot s.n. (HOLOTYPE: P, F-fragment!; Photo: GH!).
ANREDERA DENSIFLORA Sperling, spec. nov. TYPE: PERU. Lima, San Buenaventura, 2700-2800 m, 17 June 1925, Pennell 14508 (HOLOTYPE: F'!; Isotypes: GH!, NY!).
Folia ovata vel depresso-ovata, 4.0-7.7 cm longa, 2.0-9.5 cm lata, base cordata vel reniformia, apice acuta (foliis ovatis) vel rotundata (foliis depresso- ovatis). Inflorescentia terminales aut laterales, fasciculato-racemosae, inflorescentibus axillanbus pedunculo plerumque robusto portatis. Bracteae subter pedicellum tnangulares, 1.1-1.8 mm longae, 0.5-0.8 mm_latae, persistentes; bracteae pedicellorum depresso-ovatae vel perdepresso-ovatae, 1.0-1.2 mm longae; 0.5-0.8 mm latae, persistentes. Sepala late ovata vel latissime ovata, alburnea vel alba, siccitate atrobrunnea. Petala obovata vel elliptica, 1.9-2.6 mm long, 1.0-1.3 mm lata, alburnea vel alba, siccitate atrobrunnea, petaliis interioribus tribus tenuioribus quam petaliis extenoribus duobus, fructu ad matunitatem includentibus. Ovarium globosum; stylus singularis, 0.8-1.2 mm longus; stigma obscure trilobatum.
Distribution. Southern Ecuador to southern Peru. 2100-2800 (3900) m.
ADDITIONAL SPECIMENS EXAMINED. ECUADOR. Azuay: Between Molleturo and Toreador, 2590-3900 m, 14 June 1943, Steyermark 53002 (NY). Loja: Loja, 2200 m, 15 April 1946, Espinosa 137 (NY).
PERU. Lambayeque: Prov. Lambayeque, Abra de Porculla, 45 km E of Olmos on the road to Pucara, 1920 m, 13 July 1986, Plowman et al. 14290 (F). Cajamarca: Prov. Cajabamba, Nunubabamba[?], 2600 m, 13 Aug. 1985, Mostacero & Guerra 0059 (F). Huanuco: San Rafael, 8500 ft., 4 Apnl 1923, Macbride 3143 (F); Acomayo, 2100 m, 24 April 1946, Woytkowski 34245 (F,G,MO,UC,USM). Junin: Paucartambo, 2800 m, 23 July 1969, Woytkowski 6719 (GH,MO).
Anredera densiflora can be recognized by the dense inflorescence and flowers in which the sepals and outer two petals spread in fruit. It is similar to A. baselloides Baill. but differs by the ovate leaves, flared petiole, dense inflorescence, smaller flowers, broader sepals, and tnlobed stigma (not divided). This species lacks
Sperling: New species and combinations in Anredera 3
mammiillose cells at the sepal base but does form a very low keel due to contraction of the sepal during drying as in A. baselloides.
ANREDERA TUCUMANENSIS (Lillo & Hauman) Sperling, comb. nov. BASIONYM: Boussingaultia tucumanensis Lillo & Hauman, Anales Mus. Nac. Buenos Aires 33:353. 1925. (Hauman & Ingoyen, Anales Mus. Nac. Buenos Aires 32:159, 449. 1923, nom. nud.). LECTOTYPE (here chosen): BOLIVIA: Prov. of Larecaja, Sorata, between Cochipata and Milipaya along the Ulcumanini River,: 3200 m, March-May 1858, Mandon 1028 (LECTOTYPE: K!: Isolectotypes: BM!, F!, G!, GH!, K!, NY!, P; Photos: F!, GH!, MO! of G). SYNTYPE: ARGENTINA: Prov. of Tucuman, Sierra de Garabatal, 2000 m, 22 March 1922, Schreiter s.n. (LIL).
ANREDERA KRAPOVICKASII (Villa) Sperling, comb. nov. BASIONYM: Boussingaultia krapovickasii Villa, Lilloa 32:305, fig. p. 306. 1966. ‘TYPE: ARGENTINA: Salta, km 28, road between Salta and Jujuy, 31 Jan. 1947, C.A. O’Donell 4723 (HOLOTYPE: LIL).
ANREDERA BRACHYSTACHYS (Mog.) Sperling, comb. nov. BASIONYM: Tandonia brachystachys Mogq. in DC.,. Prodr. 13(2):227. 1849. LECTOTYPE (here chosen): COLOMBIA. Bogota, Goudot I (P-Herb. Mogq., det. by Mog.; Photo: GH!). SYNTYPE: ECUADOR. west side of Pichincha, 8500 ft., [without collector] (K!, P-fragment ex. Herb. Hook.; photo GH!).
ANREDERA ASPERA Sperling, spec. nov. TYPE: BOLIVIA. Prov. La Paz, Dept. Larecaja, Sorata, 68° 40’ W 15° 45’ S, 2530 m, 8 Dec. 1981, Sperling & King 5412 (HOLOTYPE: GH!; Isotype: LPB!, others not distributed).
Planta scandens vix volubilis succulenta mucliaginaque. Caules rubelli asperl praesertim ad nodos. Folia obovata, 2.1-4.2 cm longae, 1.4-2.0 cm latae, base cuneata vel acuta, apice obtusa vel rotundata. Inflorescentia laterales racemosae simplices aut base unifurcatae, pedicellis minutis, 0.5-0.7 mm longis. Bracteae subter pedicellum deltatae, 0.9-1.0 mm longae?, 0.8 mm latae?, adnatae decursivaeque; bracteae pedicellorum rhombicae _ vel perdepresse trullatae, 0.7-0.9 mm longae?, 0.8-1.0 mm latae, apice acutae, base truncatae, lobis lateralibus sagittiformibus, adnatae decursivaeque. Sepala perdepresse-ovata, 2.0-2.4 mm longa, 2.3-2.4 mm lata, apice acuta, viridulo-alba, erecta et per anthesin patentia. Petala obovata, 3 mm longa, 1.5- 1.6 mm lata, alba, erecta et per anthesin urceolata. Ovarium globosum vel obovoideum; stylus singulans, 1 mm longus, super basin ad stigma expansus; stigma obscure trilobatum capitatum. Fructus adhuc ignoti.
Distribution. Known only from the type collection in northern Bolivia.
Anredera aspera can readily be distinguished by its asperous stem and greatly flared style. The flowers are nearly sessile but upon close inspection the very short pedicel is evident. The flowers are erect at anthesis and not spreading like many species of Anredera. The pedicellar bracts are decurrent down the pedicel and continuous with it, forming a cuplet on which the flower is borne.
4 PHYTOLOGIA July 1995 volume 79(1):1-4
This species is similar to Anredera marginata, from which it differs in having always obovate leaves (even in the flowering portion of the stem), nearly sessile flowers that are slightly larger than A. marginata, and flared styles.
Unlike most species of Anredera this plant is scarcely twining, a character which is constant when the plant is cultivated in the greenhouse. In the greenhouse it is more difficult to propagate, being much slower in forming adventitious roots from cuttings than any other species of Anredera.
The species was collected growing alongside Anredera ramosa (Mogq.) Eliasson and in the same general area where the type specimen of A. fucumanensis was collected by Mandon. The type collection was observed being visited by flies, which are the presumed pollinators.
One collection from Bolivia may be this species: BOLIVIA. near La Paz, 10,000 ft., Oct. 1885, Rusby 2570 (NY two sheets, F). It has a similar pedicel and pedicellar bracts, but the leaves are lacking. Fruits are present in this specimen (enclosed in the nigrescent perianth); because the type collection lacks fruits a comparison can not be made. The petals of the Rusby collection are slightly smaller, and most of the flowers on the sheets are in poor condition.
ACKNOWLEDGMENTS
I thank David Lellinger for editing the two Latin descriptions and reviewing the paper, and Dan Nicolson for his review and suggestions.
LITERATURE CITED
Brako, L. & J.L. Zarucchi. 1993. Catalogue of the Flowering Plants and Gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1-xl, 1- 1286.
Brummitt, R.K. & C.E. Powell (Eds.). 1992. Authors of Plant Names. Royal Botanic Gardens, Kew, Great Britain.
Mogquin-Tandon, C.H.B.A. 1849. Basellaceae in Alph. de Candolle, Prodromus Systematis Naturalis Regni Vegetabilis 13(2): 220-230.
Nowicke, J.W. In press. Pollen morphology, exine structure and the relationships of Basellaceae and Didiereaceae to Portulacaceae. Syst. Bot.
Sperling, C.R. 1987. Systematics of the Basellaceae. Ph.D. Dissertation, Harvard University. UMI Dissertation Information Service, Ann Arbor, Michigan.
Phytologia (July 1995) 79(1):5-7.
STEVIA CALZADANA (ASTERACEAE) A NEW SPECIES FROM OAXACA, MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Stevia calzadana B.L. Turner, spec. nov., is described and illustrated from Oaxaca, (Mpio. Coicoyan de las Flores), México. It belongs to the series Corymbosae of Stevia where it relates to S. jorullensis, distinguished from the latter by its linear-lanceolate, glandular-punctate leaves and achenes with aristate pappus scales.
KEY WORDS: Asteraceae, Eupatorieae, systematics, Stevia, México
Routine identification of Mexican composites (Asteraceae) has revealed the following novelty.
STEVIA CALZADANA B.L. Turner, spec. nov. Figure 1. TYPE: MEXICO. Oaxaca: Mpio. Coicoyan de las Flores, Distr. Santiago Juxtlahuaca, E] Arenal, 4 km de Coicoyan de las Flores, carretera a San Martin Peras - Santiago Juxtlahuaca
(17° 17'N x 98° 15’ W), 1775-1890 m, 20 Nov 1994, J.J. Calzada 19539 (HOLOTYPE: TEX):
S. jorullensis H.B.K. similis sed foliis linearibus-oblanceolatis (vs. ovatis), glanduliferis-punctatis (vs. non glanduliferis-punctatis); et pappis acheniorum aristatis (vs. coroniformibis).
6 PHY TOLOGIA July 1995 volume 79(1):5-7
Fig. 1 Stevia calzadana, from holotype.
Tumer: New Stevia from México oe
Stiffly erect, mostly unbranched, perennial herbs ca. 1 m high. Stems reddish, vestiture puberulent with minute upcurved hairs ca. 0.2 mm high. Leaves opposite throughout, gradually reduced upwards; petioles 1-3 mm long. Midstem leaves linear- oblanceolate, 5.0-6.5 cm long, 5-6 mm wide, glabrous throughout, abundantly glandular-punctate, especially beneath, 1-nervate or weakly 3-nervate, the margins entire. Heads arranged in both terminal and axillary, mostly congested, flat-topped cymules 4-10 cm across, the ultimate peduncles mostly 1-5 mm long. Involucres cylindric, 5-6 mm long, the bracts sparsely puberulent to nearly glabrate, their apices acute. Corollas (dried) deep rose-colored, 6-9 mm long, the tube and throat indistinct, glabrous or nearly so, the lobes 1-2 mm long, glabrous without. Achenes with body 3.0-3.5 mm long, minutely hispidulous, the pappus of 3 linear artistate scales ca. 5 mm long, the upper portion barbellate for 1-2 mm, below these a crown of 3 or more united scales ca. 0.5 mm high.
This species is distinguished by its linear-oblanceolote leaves which are essentially glabrous, and 3-aristate achenes. In Grashoff's unpublished doctoral thesis (Univ. of Texas, Austin, 1972) the species will key to or near Stevia jorullensis H.B.K., but it differs from the latter in both leaf shape (linear-oblanceolate vs. ovate) texture (densely glandular-punctate beneath vs. not so), and pappus aristate (vs. coroniform, without aristae).
It is a pleasure to name this distinctive Stevia for J.1. Calzada, extraordinary collector of Mexican plants, now associated with UNAM on the flora of the Mixteca Alta region of Oaxaca.
ACKNOWLEDGMENTS
I am grateful to Gayle Turner for the Latin diagnosis, and Piero Delprete and Mark Mayfield for reviewing the manuscript. The illustration was drawn by Ms. Mana Thompson.
Phytologia (July 1995) 79(1):8-9.
MENODORA GYPSOPHILA (OLEACEAE), A NEW SPECIES FROM NEAR GALEANA, NUEVO LEON, MEXICO.
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A new species, Menodora gypsophila B.L. Turner, is described from gypseous soils near Galeana, Nuevo Le6én, México. It is closely related to the widespread M. coulteri but differs by a number of characters, including leaf- shape, venation, vestiture, and substrate preference.
KEY WORDS: Oleaceae, Menodora, systematics, México
Routine identification of plants from northeastern México has revealed the following novelty.
MENODORA GYPSOPHILA B.L. Turner, spec. nov. TYPE: MEXICO. Nuevo Leén: Mpio. Galeana, Santa Rosa, arid hillside, 1610 m, 6 Oct 1995, Hinton et al. 25643 (HOLOTYPE: TEX).
M. coulteri A. Gray similis differt foliis crassis, pro parte maxima triplinerviis et apiculatis, et caulibus moderate pubescentibus et hirsutis, pilis 0.2-0.3 mm longis (vs. pilis deorsum curvatis et 0.1-0.2 mm longis).
Low much-branched shrublets 10-20 cm high. Stems terete, moderately pubescent with widely spreading hairs mostly 0.2-0.3 mm long. Leaves opposite throughout, gradually reduced upwards, those at midstem lanceolate-elliptic, markedly thickened, bearing 3 raised nerves on the lower surface, pubescent like the stem, entire, the apices apiculate, the blades mostly 5-15 mm long, 4-6 mm wide. Flowers terminal, the pedicels reflexed in fruit. Calices 3-6 mm long; lobes 8-13, 2-4 mm long, linear- lanceolate, pubescent with spreading hairs. Corollas bright yellow; tubes 2-4 mm long; lobes 5-10 mm long, 3-6 mm wide. Anthers yellow, exserted 2-4 mm from the tube. Style exserted 3-5 mm from the tube. Paired capsules ovoid, reflexed, each ca. 5 mm across; seeds obovoid, ca. 4 mm long, 2 mm across, the outer surface spongy and irregularly patterned.
ADDITIONAL COLLECTIONS EXAMINED: MEXICO. Nuevo Leén: Mpio. Galeana, 5 km from Galeana, along the road to Rayones, 1600 m, 27 Jun 1994,
8
Turner: New Menodora from México 9
Hinton et al. 24474 (TEX); 3 km N of Galeana on rather bare gypseous-calcareous ('?) soils, 26 Jul 1993, Turner 93-158 (TEX).
Collections of this species were unknown to me at the time of my treatment of Menodora for North America (Phytologia 71:340-356. 1991.). As indicated by the specimens cited above, this taxon was first collected by myself in 1993 (along with several close-up photographs). The two subsequent collections were made by Jaime and George Hinton in about the same area, apparently also in gypseous soils. I have selected Hinton 25643 as the type of this species because the collections concerned possess relatively large well-developed leaves and bountiful flowers. The other two collections are not as lush and possess leaves about half the size of the type, with decidedly smaller flowers, especially Turner 93-158 which has very small calyces (3-4 mm long) with only ca. 8 lobes (vs. ca. 13 in the type). In most other details, however, the paratypes are like those of the holotype. When originally collected | thought that M. gypsophila might be an aberrant specimen of M. coulteri, the latter having thinner, largely enervate leaves and a finer, down-curved stem-pubescence. The additional Hinton collections have convinced me that the populations concerned deserve a name. I am especially grateful to George Hinton’s perceptive eye who sent me the most recent collection with the observation (pers. letter) that “the leaves have clear venations that are unlike any in our collections, and I couldn't match it to any in your revision of the genus”, which is so, hence the description here.
ACKNOWLEDGMENTS
I am grateful to Gayle Turner for the Latin diagnosis, and to Piero Delprete and Mark Mayfield for reviewing the paper.
Phytologia (July 1995) 79(1):10-11.
A NEW COMBINATION IN PHYSALIS (SOLANACEAE)
Barbro Axelius
Botaniska Institutionen, Stockholms Universitet, S-106 91 Stockholm, SWEDEN
ABSTRACT
Margaranthus solanaceous is transferred to the genus Physalis and hence the monotypic genus Margaranthus becomes a synonym of Physalis.
KEY WORDS: Physalis, Margaranthus, Solanaceae, systematics
Margaranthus Schidl. (Solanaceae) is a monotypic genus from México and the southwestern United States. It was described by Schlechtendal 1838. It has always been regarded as very closely related to the genus Physalis L., differing in form and colour of the corolla and insertion of filaments. The annual M. solanaceous Schlechtendal has an urceolate, violet/greenish corolla with adnate filaments while in Physalis the corollas are campanulate to nearly rotate, yellow or whitish and the filaments are free.
In his monograph, Rydberg (1896) considered Margaranthus as very closely related to Physalis but kept it as a separate genus. In a karyological report, Menzel (1950) noted the great similarities between Margaranthus and Physalis and, based on S/T ratio data placed Margaranthus between the annual sections Angulatae and Pubescentes of Physalis. She did not, however, make any formal transference. Waterfall (1958) in his survey of Physalis in North Amenca commented on_ the similarity of Margaranthus to Physalis and noted that if not in flower, Margaranthus could not be distinguished from small-fruited species of Physalis. He stated that possibly Margaranthus should be included in Physalis, but that further studies including critical species of Chamaesaracha (C. grandiflora (Hook.) Fern., C. nana (A. Gray) A. Gray, both now in Leucophysalis) were needed before formal transference. He thus kept Margaranthus as it was originally described, as a genus of its own. On the other hand he included Quincula lobata, another related, monotypic genus in Physalis, even though Quincula differs from Physalis in several characters besides colour of corolla. This was probably due to Quincula originally being described as a Physalis.
My recent cladistic analyses of the physaloid group, including among others Margaranthus, Quincula, Chamaesaracha, and Leucophysalis (Axelius 1995) has
10
Axelius: New combination in Physalis 11
shown that Margaranthus is well nested within the Physalis clade (including P. pubescens L., P. angulata L., and P. peruviana L.), close to P. pubescens (Axelius 1995, fig. 1). The species of Chamaesaracha, Quincula, and Leucophysalis are more distantly related and found clearly outside the Physalis clade. The species of Chamaesaracha group together and form a very strongly supported sister-relation with Quincula. There is thus a rather strong support for the hypothesis that Margaranthus has originated from an ancestor within the core Physalis. This view is also in accordance with analysis based on molecular data (Martinez 1993). Hence Margaranthus cannot be kept separated from Physalis without splitting the core of this genus into smaller monophyletic entities. Physalis is a large genus which lacks a modern revision and it's circumscription might be questioned in many ways but to keep Margaranthus separated under these circumstances, can not longer be justified.
PHYSALIS SOLANACEOUS (Schlechtendal) Axelius, comb. nov. BASIONYM: Margaranthus solanaceous Schlechtendal, Index Sem. Hort. Hal. 1838 Coll. 8. 1838. TYPE: Cult. in Horto Botanico Halensis 1838, “e seminis in Mexico locis calidioribus coll. C. Ehrenberg”. D.F.L. Schlechitendal s.n. (HOLOTYPE: HAL).
LITERATURE CITED
Axelius, B. 1995. The phylogenetic relationships of the physaloid genera (Solanaceae) based on morphology. Amer. J. Bot. In press.
Martinez, M. 1993. Systematics of Physalis (Solanaceae) section Epeteiorhiza. Ph.D. Dissertation, University of Texas at Austin. Austin, Texas.
Menzel, M. 1950. Cytotaxonomic observations on some genera of the Solaneae: Margaranthus, Saracha and Quincula. Amer. J. Bot.. 37:25-30.
Rydberg, P.A. 1896. The North American species of Physalis and related genera. Mem. Torrey Bot. Club 4:297-374.
Schlechtendal, D.F.L. 1838. Index Seminum in Horto Academico Halensi 1838 Collectorum. Halle, Germany.
Waterfall, U.T. 1958. A taxonomic study of the genus Physalis in North America north of Mexico. Rhodora 60: 107-114; 128-142; 152-173.
Phytologia (July 1995) 79(1):12-21.
TAXONOMIC OVERVIEW OF HEDYOTIS NIGRICANS (RUBIACEAE) AND CLOSELY ALLIED TAXA
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A taxonomic study of Hedyotis nigricans is rendered in which a widespread var. nigricans is recognized, along with five regional or localized, allopatric varieties: var. floridana (southern Florida); var. pulvinata (northeastern Florida); var. austrotexana B.L. Turner, var. nov. (southern Texas); var. gypsophila B.L. Turner, var. nov. (montane regions of Nuevo Leén, México and closely adjacent states); and var. papillacea B.L. Turner, var. nov. (northern panhandle and trans-Pecos, Texas, and closely adjacent New Mexico). A key to these taxa is provided along with maps showing their distributions. Additionally, these taxa are compared with the closely related species H. angulata and H. butterwickiae, and maps showing their distribution are also provided.
KEY WORDS: Rubiaceae, Hedyotis, Houstonia, systematics, Mexico, Texas
Hedyotis nigricans (Lam.) Fosberg (=Houstonia nigricans [Lam.] Fern.) has been variously treated as belonging to the genus Hedyotis or Houstonia (Shinners 1949; Terrell 1986, 1991), some workers preferring an inclusive Hedyotis (including Houstonia), others preferring a more restricted Hedyotis (excluding Houstonia, cf. Terrell 1991). Most current workers accept Hedyotis nigricans as belonging to Hedyotis, including Terrell (1991), albeit tentatively. Terrell (1986) provided a taxonomic overview of H. nigricans for the U.S.A., especially Florida, but did not treat in detail collections from Texas, New Mexico, and México.
The present contribution is based upon the detailed examination of over 800 sheets of Hedyotis nigricans on file at LL, TEX, and SRSC. Key to Texas populations of Hedyotis nigricans and closely related taxa 1. Leaves mostly basal, very numerous and forming pulvinate mats, the stiffly erect
rather naked stems having markedly appressed, stiff-lanceolate leaves; fruits mostly orbicular; southeastern most Brewster Co. ................4. H. butterwickiae
12
~
Turner: Overview of Hedyotis 13
1. Leaves otherwise, mostly cauline and spreading; fruits mostly ovoid (except for Gulf Coastal populations); widespread. «.........is5.¢cessencvsssnensisorsserersecenvass (2)
2. Midstem leaves thick and short, ovate-linear to lanceolate, 1 cm long or less, the margins never enrolled; capsules orbicular at maturity; calyx lobes 1 mm
long or less; rock or cliff-dwelling species of eastern trans-Pecos, Texas and Closely adjacent | MEXICO. sccct.cssderienesseenteswusesannscates H. angulata Fosberg
2. Midstem leaves not as described in the above, the margins to some extent enrolling with dessication; capsules ovoid at maturity; calyx lobes mostly 1 mm
or more long, if shorter then the leaves decidedly linear to linear-oblanceolate; mostly not rock or bare-bluff species, widespread (H. nigricans). ............ (3)
3. Calyx, and/or upper stems and leaves to some extent papillose with extended epidermal cells, these superficially resembling hairs, or else the calyx to some extent beset with callose hair-like emations................cccccccecceceeceeeeseeeneeens (4)
3. Calyx, upper stems and leaves glabrous or merely ciliate along the leaf margins and CAL Kel ODES; 15th ad tanga ntuaarasseabnouswas Meader udesuoxGugensdumdysteey var. nigricans
4. Plants mostly sprawling, low bushy herbs 5-15 cm high; panhandle and trans- BCCOS MIEN AS .3 3) te auetet Neestee Meng beeen meee nt einceene ean cms var. papillacea
4. Plants mostly simple-stemmed, non bushy herbs 20-40 cm high; southern MORAG se eeadon cars cece en Sata iansera tenn (year Senet ... Var. austrolexana
Key to Mexican populations of Hedyotis nigricans
1. Primary leaves at midstem mostly 1-3 mm wide, 3-12 times as long as wide; calyx usually glabrous, or with but a few ciliate hairs; mostly calcareous soils, WW EGGS CO Go ccs seeps beac otubuns aes es ue eee nies conde hSe isin oven dnaeuea segs var. nigricans 1. Primary leaves at midstem mostly 3-6 mm wide, 2.5-3.5 times as long as wide; calyx usually markedly setose with thickened hairs; mostly gypseous soils of southernmost Coahuila, Nuevo Leén, and very closely adjacent Zacatecas and probally Tamaulipas: ts:ci.cisasie rea eeaicabe, soccteesncaev eerie cee sass var. gypsophila
HEDYOTIS BUTTERWICKIAE (Terrell) Nesom, Syst. Bot. 13:434. 1988. Houstonia butterwickiae Terrell
This species, first described by Terrell in 1979, was retained by both Nesom (1988) and Terrell (1991). It is known only by collections from along the ridgetop of the Bullis Range on the Bullis Gap Ranch, in Brewster Co., which is about 20 mi S of Sanderson (Terrell Co.). The taxon is obviously closely related to Hedyotis nigricans but can be immediately recognized by its very narrow, linear-lanceolate, relatively thickened stiffly ascending stem leaves, the basal leaves forming a dense pulvinate array of shorter, somewhat broader elliptic-lanceolate leaves. Additionally, Terrell, in his original description, notes that the taxon has nearly globose capsules and a relatively diffuse inflorescence.
HEDYOTIS NIGRICANS (Lam.) Fosberg, Lloydia 4:287. 1941.
Terrell (1986) treated this taxon as belonging to the genus Houstonia but subsequently (1991) positioned it in Hedyotis. He considered H. nigricans to be a “polymorphic species”, but nevertheless recognized three varicties in the complex, a
14 PHYTOLOGIA July 1995 volume 79(1):12-21
widespread highly variable var. nigricans, and two very localized varieties in Florida, both coastal (Figure 1).
I have examined numerous sheets of this species from over a broad region and recognize three additional varietal taxa, as described below. All of these are largely allopatric with var. nigricans but appear to intergrade into the latter in regions of near contact.
Even with the removal of these several newly described elements, var. nigricans remains quite variable, ranging from rather spindly plants with linear to linear- oblanceolate leaves in eastern and central Texas, to shorter plants with linear-lanceolate to linear-elliptic leaves in west-central Texas and westwards. In México the variety, while quite variable, is less complex, as shown in Figure 2.
HEDYOTIS NIGRICANS var. NIGRICANS
Terrell (1986, 1991) accounted for most of the Texas names involved in the synonymy of this variety, and these are briefly touched upon here.
Houstonia salina A.A. Heller -This name is typified by material from coastal areas of southern Texas (Nueces Co.). I agree with Terrell (1986) that the plants concerned differ but little from typical elements of var. nigricans.
Houstonia tenuis Small -This name is typified by material from central Texas (San Saba Co.) and appears to be the same as var. nigricans, as noted by Terrell (1991).
Houstonia angustifolia Michx. var. rigidiuscula A Gray, Syn. Fl. N. Amer. 1(2):27. 1884. Shinners (1949) transferred this variety into Hedyotis nigricans without comment, merely noting it to be typified by plants collected in “S. and W. Texas, Palmer, Havard, & c. Coast of E. Florida, Rugel. (Mex)”. Unfortunately, to my knowledge, no one has lectotypified the name concerned, but my own evaluations of this apellation are that it was meant to apply to plants having a low nigid stature, mainly occurring in the southwestern U.S.A. (western Texas, New Mexico, and Arizona) and México. Those who might wish to apply this vanetal name to such plants over this region might do so, but I view the variation between such _habital forms as relatively trivial, there being gradual intergradation between such populational forms over a broad region of central Texas and northern México. In short, there seems little merit in attempting to define what the habital limits of var. rigidiuscula might be.
Houstonia angustifolia Michx. var. scabra S. Wats., Proc. Amer. Acad. Arts 18:97. 1883. (TYPE: MEXICO. Coahuila: Carocol Mts, 19-20 Aug 1880, F. Palmer 410; Isotype: LL!). -This name is unaccounted for by Terrell (1986, 1991) but examination of type material shows this to belong to var. nigricans. The Carocol Mts are said to be located about 24 mi southeast of Monclova, México (McVaugh 1956), an area well within the distribution of var. nigricans as defined in the present treatment.
Tumer: Overview of Hedyolis 15
HEDYOTIS NIGRICANS (Lam.) Fosberg var. AUSTROTEXANA B.L. Turner, var. nov. TYPE: U.S.A. Texas: Karnes Co., roadside 2 mi E of H Tejano Cafe, “dry sandy, clay soil”, 22 Jun 1952, Joe C. Johnson 833 (LL).
H. nigricans (Lam.) Fosberg var. austrotexana B.L. Turner, var. nov.; similis H. nigricans var. nigricans sed calyces matun valde papillosi ubique.
Resembling var. nigricans but the mature calyces markedly papillose throughout.
Other than having markedly papillose calyces, this taxon is essentially the same as var. nigricans; indeed, it apparently replaces the latter in the region shown in Figure 1. The two varieties do, however, grow in close proximity and occasional plants appear to show intergradation of the calyx character concerned in regions of near contact (e.g. Goliad Co.: Smith 4271; San Patricio Co.: Turner 80-91M).
HEDYOTIS NIGRICANS (Lam.) Fosberg var. GYPSOPHILA B.L. Turner, var. nov. TYPE: MEXICO. Nuevo Leon: Santa Rita, 2370 m, “Sparse pine woods - gypsum hillsides,” 11 Jun 1981, Hinton et al. 18278 (HOLOTYPE: TEX).
H. nigricans (Lam.) Fosberg var. gypsophila B.L. Turner, var. nov.; similis H. nigricans var. nigricans sed plantae parviores et ramosissimi e basi; folia ovato-elliptica et saepius 2.5-3.5 plo longiores quam latiores (vice folia linearia-lanceolata usque linearia-oblanceolata et saepius 4-20 plo longiora quam latiora); calyces matun plerumque hispidi enatis capillaribus et latis basi (vice calyces glabros enatis infirme evolutis).
Resembling var. nigricans but the plants low and much-branched from the base, the leaves elliptic-ovate and mostly 2.5-3.5(4.0) times as long as wide (vs. linear- lanceolate to linear-oblanceolate, mostly 4-20 times as long as wide) and the mature calyces usually markedly hispid with broad-based hairs (vs. glabrous or merely minutely setose).
This taxon is represented by 45 or more collections at LL, TEX, mostly obtained from gypseous soils in the state of Nuevo Leén. While quite variable as concerns calyx pubescence, the branching habit and leaf shape is very diagnostic, and in combination the characters are as distinctive for recognition purposes as_ those characters in combination used by, for example, Terrell in his recognition of Hedyotis nigricans var. pulvinata (Small) Fosb., the latter superficially resembling var. gypsophila as conceived here.
HEDYOTIS NIGRICANS (Lam.) Fosberg var. PAPILLACEA B.L. Turner, var. nov. TYPE: U.S.A. New Mexico: Otero Co., northern McKittrick Canyon at first crossing of Texas-New Mexico boundary on the New Mexico side, “gravels and boulders of stream bottom. In Riparian type habitat and below protected cliffs; Big tooth maple, Ponderosa Pine, Madrone”, etc. 8 Oct 1973, Thomas F. Patterson 508 (HOLOTYPE: LL).
volume 79(1):12-21
July 1995
PHYTOLOGIA
16
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Shy, wat Sete ieet meets
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. ott Sat Pass
Po ede gwcanhia tie cist Meee Bee ga cr een
TTT 9g pee 4 COTS TT HL nee Sc‘ Pimge mY Ba aca Nene Sanesan nN SIT 600-64 re ME Alas aren Peeeee SBF x. “ ott sane 9 Ter eS a OOo «i or ‘estes, 2-Lan ; CK bre & tine oom LCS Siro eh eet Mas sy LS Se Se eo “a A= a ye \ J 1 C58
var. nigricans (closed circles); gles); var. papillacea (open
gonals); var. floridana (closed trian
n of Hedyotis nigricans in U.S.A.: circles); var. pulvinata (open triangles).
Figure 1. Distributio var. austrotexana (dia
Turner: Overview of Hedyotis 17
Figure 2. Distribution of Hedyotis nigricans complex in Texas: var. circles); var. papillacea (closed circles); + intermediates to Var. algricans and \ ai papillacea (half circles); var, austrolexana (triangles).
NIZFICANS (Open
volume 79(1):12-21
PHYTOLOGIA July 1995
18
yw V4 FR S pad y - a ae | | Ne Ne hee N ps AB (
y similar
(open circles) and the superficiall ); H. butterwickiae (triangle).
gulata ircles
Figure 3. Distribution of Hedyotis an H. nigricans var. papillacea (closed ci
Tumer: Overview of Hedyotis 19
Figure 4. Distribution of Hedyotis nigricans in México: var. nigricans (open circles); var. gypsophila (closed circles); intermediates (half circles).
20 PHYTOLOGIA July 1995 volume 79(1):12-21
H. nigricans (Lam.) Fosberg var. papillacea B.L. Turner, var. nov.; similis H. nigricans var. nigricans sed saepius 5-15 cm alta, enascens caudicibus ramosis et ligneis; caules, folia, calycesque aliquantum papillosi enatis capillaribus.
Resembling var. nigricans but the plants mostly 5-15 cm high and the stems, leaves and calyces to some extent papillose with hair-like enations.
As shown in Figure 1, the var. papillacea is largely confined to the northernmost panhandle region of Texas where it passes, rather abruptly, into var. nigricans. It also is found in the trans-Pecos regions of Texas and closely adjacent New Mexico where it reportedly occurs in and along limestone ledges and bluffs (Del Norte, Glass and Guadalupe Mts). Plants of the latter region superficially resemble Hedyotis angulata, and some of these were annotated as such by Terrell (e.g., Warnock 7978, from the Del Norte Mts [TEX]). Inclusion of the panhandle collections with the trans-Pecos material might appear moot in that the panhandle collections have somewhat longer, more linear-lanceolate leaves and occur as populational disjuncts. However, similar populational disjunctions occur in several species of Asteraceae (e.g., Chrysothamnus) and need not be cause for much concern, at least I find it difficult to distinguish between the two populational elements.
It should be noted that the type collection of var. papillacea was found growing with or near material that might be deemed to be var. nigricans (Patterson 508, 516) in that these two relatively late-flowering collections seemingly lack the papillose enations which characterize the taxon, but in all other characters these two plants resemble var. papillacea as conceived here. The same is true for occasional specimens from the
panhandle region, where the variety is apparently much more common, to judge from herbarium collections.
Finally, it should be emphasized that from among the 1000 or more specimens of var. nigricans examined by me in the present study, only a few sheets were discerned to have papillose enations of the type found in var. papillacea, at least two of these from the state of Florida (Franklin Co., sand dunes and coastal areas along the Gulf of Mexico: Henderson 63-1309, Kral 39899). Obviously such enations are under relatively simple genetic control, but in the var. papillacea these appear to be populationally “fixed” in the regions shown in Figure 1, occurring on plants with a habital display quite different from the habital display of var. nigricans over most of its eastern distribution.
ACKNOWLEDGMENTS
I am grateful to my wife, Gayle, for the Latin diagnoses, and to her and Piero Delprete for reviewing the manuscript.
Turner: Overview of Hedyotis 21
LITERATURE CITED
Lewis, W.H. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Gard. 55:31-33.
Shinners, L.H. 1949. Transfer of Texas species of Houstonia to Hedyotis (Rubiaceae). Field. & Laboratory 17:166-169.
Terrell, E.E. 1986. Taxonomic and nomenclatural notes on Houstonia_ nigricans (Rubiaceae). Sida 11:471-481. ___. 1991. Overview and annotated list of North American species of Hedyotis, ~ Houstonia, Oldenlandia, and related genera. Phytologia 71:212-243.
McVaugh, R. 1956. Edward Palmer, Plant Explorer of the American West. Univ. of Oklahoma Press, Norman, Oklahoma.
Phytologia (July 1995) 79(1):22-24.
THE STATUS OF QUERCUS ARKANSANA SARG. (FAGACEAE) IN TEXAS
David M. Hunt
New York Natural Heritage Program, 700 Troy-Schenectady Road, Latham, New York 12110-2400 U.S.A.
& Michael H. MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A.
ABSTRACT
The status of Quercus arkansana Sarg. in Texas is discussed. It is known to occur in Cass County, and there is an historical record for Jasper County.
KEY WORDS: Quercus arkansana, Texas, Fagaceae, phytogeography
Quercus arkansana Sarg., an uncommon Coastal Plain oak with scattered populations ranging from southwestern Georgia and northwestern Florida to southwestern Arkansas and northwestern Louisiana, has not been known for Texas (Bill Carr, Texas Natural Heritage Program, pers. comm.; Correll & Johnston 1970; Johnston 1990; Stanley D. Jones, Botanical Research Center (BRCH), pers. comm.; Hunt 1986; Little 1977; Nixon 1985; Sargent 1965; Simpson 1988; Vines 1977). Hunt (1990) lists the species as occurring in Texas on the basis of an historical specimen collected in 1903 in Jasper County (W.W. Ashe 1 [NCU]). More recently, two small populations of Q. arkansana have been found in Cass County.
In 1990, Hunt (Hunt TX218 [to be distributed to TEX and ND)}) first confirmed Quercus arkansana from Texas, identifying it as “Q. arkansana tending toward Q. nigra.” This collection, representing one sapling, was from the south side of TX 77, 1 km west of the Louisiana state line in sandy loam pine-oak woods dominated by Pinus echinata P. Mill., Quercus falcata Michx., Q. nigra L., and Liguidambar styraciflua L. This find was the result of searches stimulated by the presence of a large population in Louisiana about 3 km away, documented initially by G.H. Ware (Ware 1492 [DAV]) in 1965 (Hunt 1990).
In 1995, the MacRobertses found several small trees (up to 7 meters high) in a young, densely stocked, even-aged pine plantation on deep sandy soils along a 1 km
De
Hunt et al.: Status of Quercus arkansana in Texas 23
stretch of Cass County Road 4561 about 5 km northeast of McLeod, Texas, and 5 km west of Rodessa, Louisiana, which they identified as Quercus arkansana (MacRoberts & MacRoberts 2875 [NCSC], 2874, 2877 [BRCH], 2879 [VDB], 2881 [LSUS], 2873 [ND], 2878 [DAV]). Some of these specimens were sent elsewhere for confirmation. Hunt determined 2876 as “probably Quercus arkansana x Q. nigra;” 2875 and 2873 were identified as Q. arkansana by J.W. Hardin and Richard Jensen, respectively. Julia Larke determined an unnumbered specimen to be Q. arkansana. This population is within 2 km of a large population of Q. arkansana in Louisiana centering on a point where Black Bayou and State Line Creek cross LA 168 about 3 km west of Rodessa (Teague & Wendt 1994). Associated species for this site include Quercus marilandica Muenchh., Q. stellata Wang., Q. incana Bartr., Pinus taeda L., P. echinata, Vitis spp., Sassafras albidum (Nutt.) Nees, and Liquidambar styraciflua. The area was heavily shaded, had a dense pine litter and almost no herbaceous understory, and was badly damaged by commercial forestry.
The environment of the Cass County sites is typical for the species: midslope of eroding sandhills adjacent to headwater tributaries. That Quercus arkansana occurs in Cass County is not surprising since it occurs in adjacent parishes and counties in Louisiana and Arkansas (Hunt 1990; Louisiana Natural Heritage 1995; Smith 1988). Hunt (1986) has documented increased introgression between Q. arkansana and Q. nigra at the range edge of the former. Thus the mixed traits found in the Cass County collections are expected: short petioles (Q. nigra), scurfy leaf and twig pubescence (Q. arkansana), and leaf shape varying between the two provide strong evidence for putative hybridization between these species.
In addition to these recent finds, Hunt located a specimen from Jasper County (W.W. Ashe I [NCU]) collected in 1903 that is probably Quercus arkansana but may be a hybrid between Q. nigra and Q. velutina Lam. (Hunt 1990). This specimen was originally identified as Q. velutina x Q. laurifolia Michx. Unfortunately, a specific locality is not given and, although much potential mesic sandhill habitat exists in the northern third of the county, relocating this population, if it still exists, may be difficult. The east Texas landscape has been drastically altered over the past century, and Q. arkansana is notorious for its patchy distribution and its inconspicuousness, which has always made it difficult to locate (Hunt 1986).
These populations represent westward range extensions for Quercus arkansana. Additional populations of this oak are expected and should be sought in Bowie, Marion, and Cass counties. The rarity of this species in Texas and throughout its range makes it a good candidate for any protected plant list for the state, and an effort to protect its habitat should be made to ensure its survival in Texas.
ACKNOWLEDGMENTS
__ Stanley Jones (BRCH) and Bill Carr (Texas Natural Heritage Program) supplied information on the distribution of Quercus arkansana. J.W. Hardin (NCSC), Richard Jensen (ND), and Julia Larke (Louisiana Natural Heritage Program), confirmed the MacRobertses’ initial identification. Hunt’s research was supported by NSF Grant BSR-8414419 to David E. Giannasi and David M. Hunt. Thanks are due to Lon
24 PHYTOLOGIA July 1995 volume 79( 1):22-24
Hunt and Michael Moore for assistance in the field, and Nancy Coile for handling herbarium loans. Richard Jensen and Stanley Jones reviewed the manuscript.
LITERATURE CITED
Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner, Texas.
Hunt, D.M. 1986. Distribution of Quercus arkansana in Georgia. Castanea 51:183- 187.
Hunt, D.M. 1990. A Systematic Review of Quercus Series Laurifoliae, Marilandicae and Nigrae. Ph.D. dissertation, University of Georgia, Athens, Georgia.
Johnston, M.C. 1990. The Vascular Plants of Texas. A list, up-dating the Manual of the Vascular Plants of Texas. Published by the author, Austin, Texas.
Little, E.L. 1977. Atlas of United States Trees. Vol. 4. Minor Eastern Hardwoods. Misc. Publ. 1342. USDA Forest Service, Washington D.C.
Louisiana Natural Heritage Program. 1995. Rare plant species of Louisiana. Unpublished report. Louisiana Department of Wildlife and Fisheries, Baton Rouge, Louisiana.
Nixon, E.S. 1985. Trees, Shrubs, & Woody Vines of East Texas. Bruce Lyndon Cunningham Productions, Nacogdoches, Texas.
Sargent, C.S. 1965. Manual of the Trees of North America. Dover Press, New York, New York.
Simpson, B.J. 1988. A Field Guide to Texas Trees. Texas Monthly Press, Austin, Texas.
Smith, E.B. 1988. An Atlas and Annotated List of the Vascular Plants of Arkansas. Published by the author.
Teague, J. & T. Wendt. 1994. Caddo and Bossier Parishes, LA: Natural areas survey. Unpublished Report. The Nature Conservancy. Baton Rouge, Louisiana
Vines, R.A. 1977. Trees of East Texas. University of Texas Press, Austin, Texas.
Phytologia (July 1995) 79(1):25-27.
CHROMOSOME NUMBERS REPORT
Y olanda Herrera-Arrieta
CIIDIR Unidad Durango, Instituto Politécnico Nacional, Apartado Postal 57, Durango, Dgo., C.P. 34000 MEXICO Becaria de la COFAA
ABSTRACT
A first record of Muhlenbergia quadridentata chromosome number is given, a diploid chromosome number for M. virescens is reported for the first time, and the tetraploid condition of M. montana is confirmed. _—
RESUMEN
Se registra por primera vez el ntimero cromosémico de Muhlenbergia quadridentata, un nimero cromosémico diploide para M. virescens se reporta por primera vez, y la condici6n tetraploide de M. montana se confirma.
KEY WORDS: Muhlenbergia, Poaceae, cytology
INTRODUCTION
The mostly American genus Muhlenbergia Schreber is comprised by two rather distinct groups which had been cytologically studied by several authors. Pohl (1964) studied the broad-leaved, mesic, rhizomatous species from the deciduous forests of eastern North America. While Peterson (1988), did chromosome counts for the group of annuals comprising the xeromorphic caespitose species, distributed in the western
plains of America. They both reached the conclusion that the basic number for the genus is x= 10.
Chromosome counts were completed during a systematic study of the Muhlenbergia montana (Nutt.) Hitch. complex (Herrera-A. & Bain 1991; Herrera- Armieta & Grant 1993, 1994), a group of thirteen perennial, xeromorphic caespitose species. Chromosome counts for three species of the complex were successful and are here reported. Chromosome number for M. quadridentata (H.B.K.) Kunth is a first report, the M. virescens (H.B.K.) Kunth chromosome count seems to be the first
aD
26 PHYTOLOGIA July 1995 volume 79(1):25-27
diploid record of the aneuploid number reported by Reeder (1967), and the M. montana tetraploid condition reported by Reeder (1968) is confirmed.
MATERIAL AND METHODS
Chromosome determinations are based on observations of up to twelve cells from a minimum of five individuals per population, using a phase contrast microscope. Floral buds were field collected in 95 percent ethanol-glacial acetic acid (3:1) prior to fixation and storage under refrigeration in 70% ethanol. To stain: Hydrolyze in 1N
HCI at 60°C for 7 to 10 minutes, and stain in Feulgen reagent for 2 hours, rinsed in running tap water for 3 minutes. Slides were prepared in a drop of 45% acetic acid, and squashing the floral buds under a cover slip. The cover slip was temporarily sealed with a paraffin-gum arabic mixture. Attempts to grow the plants from this species complex under greenhouse conditions failed, and therefore no mitotic counts were possible.
RESULTS POACEAE:
Muhlenbergia quadridentata (H.B.K.) Kunth, n= 10. MEXICO. México: 2 km W of Rfo Fro, North exposition of Volcan Iztaccihuatl, 3100 m, Pinus-Quercus forest, Herrera & Cortés 919 (CIIDIR,MTMG).
Muhlenbergia virescens (H.B.K.) Kunth, n = 10. MEXICO. Chihuahua: 25.6 miles S of Creel on road to Batopilas, 2100 m, table rock with Arctostaphylos, Pinus and Quercus spp., Herrera with Peterson & Annable 969 (CHDIR,MTMG).
Muhlenbergia montana (Nutt.) Hitche.,n = 20. MEXICO. México: Entrance to the National Park “Lagunas de Zempoala”, 2960 m, forest of Pinus hartwegii and Abies religiosa, Herrera & Cortés 926 (CIIDIR,MTMG).
All of them showing stable microsporocytes with normal bivalents during meiosis.
Recorded chromosome numbers from Reeder (1967, 1968) are: Muhlenbergia virescens 2n = 24 and M. montana 2n = 40.
DISCUSSION
The basic chromosome number recognized for Muhlenbergia is settled as x = 10 (Pohl 1964; Reeder 1967, 1968; Peterson 1988). Diploidy (= 10) and tetraploidy (7 = 20) are the most common in this genus, however one case of octaploidy was reported by Pohl (1964) for M. californica Abrams, a rare endemic species.
Chromosome counts remain necessary to support the interpretation of evolution in this genus. One of the important findings here is that the more widely distributed species of the complex (Muhlenbergia montana) 1s a tetraploid, while the other two
Herrera-Armieta: Chromosome numbers in Muhlenbergia 27
species from more restricted geographic areas (M. quadridentata and M. virescens), are diploids. All this seems to support the theory of evolution of grasses (Stebbins 1956).
ACKNOWLEDGMENTS
The author acknowledges the Instituto Politécnico Nacional and COFAA-IPN for financial support, and thanks the reviewers.
LITERATURE CITED
Herrera A. Y. & J.F. Bain. 1991. Flavonoid profiles in Muhlenbergia montana complex. Biochem. Syst. Ecol. 19:665-672. .
Herrera-Arrieta, Y. & F.W. Grant. 1993. Correlation between generated morphological character data and flavonoid content of species in the Muhlenbergia montana complex. Can. J. Bot. 71:816-826.
Herrera-Arrieta, Y. & F.W. Grant. 1994. Anatomy of the Muhlenbergia montana (Poaceae) complex. Amer. J. Bot. 81:1038-1044.
Pohl, R.W. 1964. Cytogeography of the rhizomatous American species of Muhlenbergia. Brittonia 17:107-112.
Peterson, P.M. 1988. Chromosome numbers in the annual Muhlenbergia (Poaceae). Madrofio 35:320-324.
Reeder, J.R. 1967. Notes on Mexican grasses VI. Miscellaneous chromosome numbers. Bull. Torr. Bot. Club. 94:1-17.
Reeder, J.R. 1968. Notes on Mexican grasses VIII. Miscellaneous chromosome numbers. Bull. Torr. Bot. Club 95:69-75.
Stebbins, G.L. 1956. Cytogenetics and evolution of the grass family. Amer. J. Bot. 43:890-905S.
Phytologia (July 1995) 79(1):28-30.
A NEW COMBINATION IN MUHLENBERGIA (POACEAE)
Clifford W. Morden
Department of Botany and Center for Conservation Research and Training, 3190 Maile Way, University of Hawaii at Manoa, Honolulu, Hawaii 96822 U.S.A. cmorden@uhunix.uhcc.hawalii.edu
ABSTRACT
Systematic analyses of morphological and anatomical variation among populations of Muhlenbergia villiflora and M. villosa indicate that there is insufficient differentiation to warrant recognition of these taxa as separate species. The new combination of M. villiflora var. villosa is proposed as a more appropriate means of recognizing the habitat preference and slight differences in spikelet size that distinguish these taxa. Nomenclatural data and a key to the varieties are provided.
KEY WORDS: Muhlenbergia, Poaceae, systematics
Species of the Muhlenbergia repens Hitchc. complex are distributed throughout North America (excluding the southeastern United States), and in the Andean highlands of South America. This complex consists of eight species characterized by a rhizomatous perennial habit with short culms seldom exceeding 20 cm, short involute leaf blades, and a short contracted panicle with awnless or mucronate spikelets. Two species, M. villiflora Hitchc. and M. villosa Swallen, differ from the others by having a densely villous lemma and palea. These species appear distinct from each other in that M. villosa is slightly larger in all morphological attributes including plant height, leaf size, inflorescence length, and spikelet length. Their distnbution and _ habitat requirements are also distinct; M. villiflora is an ecological dominant in gypsum soils of northern México, and M. villosa is locally sporadic to common in alkaline or calcareous soils of west Texas and New Mexico.
Morphological and anatomical analyses of these two species (Morden 1985; Morden & Hatch 1987) have shown that specific recognition of both taxa is not warranted. Anatomically, these taxa are indistinguishable (Morden & Hatch 1987), and plants of Muhlenbergia villiflora found growing along the margins of their natural habitats (7.e., soils with a more abundant water supply or lower concentration of gypsum) are larger, and approach M. villosa in most characteristics. Therefore, these species are herein treated as a single species, M. villiflora, and the two forms are recognized as varieties, var. villiflora and var. villosa (Swallen) Morden based on_ their
28
Morden: New combination in Muhlenbergia 29
habitat preferences and morphological differences. A key to the varieties and complete descriptions are provided below.
Spikelets usually less than 2.0 mm long; plants of gypsiferous soils of northern
INICXICO scrapes Sos on iasuchatabuicenas tan beds aabdswanapelenaane 1. M. villiflora var. villiflora Spikelets usually greater than 2.0 mm long; plants of alkaline or calcareous soils, west Mexas-and NEw MEXICON 2.5. os 0 Sanus otesscaroctepearesaans 2. M. villiflora var. villosa
1. Muhlenbergia villiflora Hitchc. var. villiflora, North Amer. Fl. 17:470. 1935.-- Vilfa pubescens Fourn., Mex. Pl. 2:102. 1886. TYPE: MEXICO. Canon de las Mifias et Victoria, inter Michibuana et Tanquecillos, Karwinsky 1012 (HOLOTYPE: P; Type fragment: US!). Not Muhlenbergia pubescens (H.B.K.) Hitche. (North Amer. Fl. 17:460. 1935.).
Perennial with scaly rhizomes; the scales 3-18 mm long, acute, often deteriorating with age. Culms much-branched above, wiry, erect, seldom spreading, 7-17 cm tall (rarely higher), 0.3-0.7 mm diam., glabrous; internodes 5-33 mm long, nodulose- roughened at least below the inflorescence. Sheaths 3-16 mm long, usually about 1/2 the length of the internode, margins hyaline and clasping at the base, open and diverging from culm near the leaf collar. Ligules membranous, 0.3-1.5 mm long, erose, toothed, or acute, decurrent. Blades 3-19 mm long, 0.2-1.0 mm wide, abaxial surface glabrous, adaxial surface pubescent, strongly involute and arcuate spreading, margins scabrous, prominent midvein absent. Inflorescence a contracted panicle, 1-4 cm long (occasionally longer), 1-15 mm wide or wider if branches open or reflexed, usually exserted above the upper leaf sheath; inflorescence branches solitary at each node, with 4-11 nodes per inflorescence; branches ascending. Pedicels 0.1-1.1 mm long, minutely setose. Spikelets 1.4-2.3 mm long, not crowded on the branches, 1-15 spikelets on the lowermost panicle branch. Glumes equal, 0.6-1.7 mm long, acute, 1/2-2/3 the length of the floret, 1 (occasionally 2-cr 3-) -nerved, green or purple. Lemmas acute, 1.3-2.3 mm long, 3-nerved, densely villous near the base and along midnerve and margins to near the apex, green or becoming purple at maturity; mucro absent to 0.6 mm long. Paleas 1.0-2.1 mm long, densely villous between the nerves, with color similar to lemma. Anthers 0.7-1.6 mm long, yellow, dark green, or purple. Caryopses narrowly elliptic to linear, 0.7-1.2 mm long, 0.2-0.3 mm wide, dark brown. Chromosome number 2” =20, 22 (Reeder 1967).
Distribution. México: Chihuahua, Coahuila, Hidalgo, Nuevo Leén, San Luis Potosi, and Zacatecas. Open ground in gypsiferous to calcareous soils, often forming extensive stands across gypsum flats.
2. MUHLENBERGIA VILLIFLORA Hitche. var. VILLOSA_ (Swallen) Morden, stat. nov.-- BASIONYM: Mudhlenbergia villosa Swallen, J. Wash. Acad. Sci. 31:350. f. 2. 1941. TYPE: UNITED STATES. Texas: 15 miles south of Stanton, 11 July 1928, Tharp 5048 (HOLOTYPE: US!; Isotypes: GH!,MO!,TEX!).
_ Perennial with scaly rhizomes; the scales 5-16 mm long, acute, often deteriorating with age. Culms much-branched above, wiry, erect, seldom spreading, 4-30 cm tall, 0.3-0.7 mm diam., glabrous; internodes 5-37 mm long, nodulose-roughened at Icast below the inflorescence. Sheaths 5-15 mm long, usually about 1/2 the length of the internode, margins hyaline and clasping at the base, open and diverging from culm
30 PHY TOLOGIA July 1995 volume 79(1):28-30
near the leaf collar. Ligules membranous, 0.4-1.5 mm long, erose, toothed, or acute, decurrent. Blades 7-30 mm long, 0.2-1.2 mm broad, abaxial surface glabrous, adaxial surface pubescent, strongly involute and arcuate spreading, margins scabrous, prominent midvein absent. Inflorescence a contracted panicle, 1-5 cm long, 1-5 mm wide, usually exserted above the upper leaf sheaths; inflorescence branches solitary at each node, with 5-11 nodes per inflorescence; branches ascending. Pedicels 0.1-1.2 mm long, minutely setose. Spikelets 1.8-2.5 mm long, not crowded on the branches, with 2-9 spikelets on the lowermost panicle branch. Glumes equal, 0.6-1.8 mm long, acute, 1/2-2/3 the length of the floret, 1 (rarely 2) -nerved, green or purple. Lemmas acute, 1.8-2.4 mm long, 3-nerved, densely villous near the base and along the midnerve and margins to near the apex, green or becoming purple with maturity; mucro absent to 0.4mm long. Paleas 1.7-2.3 mm long, densely villous between the nerves, color similar to the lemma. Anthers 0.9-1.4 mm long, yellow, dark green or purple. Caryopses narrowly elliptic to linear, 1.0-1.4 mm long, 0.2-0.4 mm wide, dark brown. Chromosome number 2n =20, 40 (Morden 1985; Reeder 1967).
Distribution. United States: southern New Mexico and Texas in the Trans-Pecos, western Edwards Plateau and southern High Plains. Open ground in alkaline to calcareous soils, usually in isolated clumps and seldom forming dense stands.
LITERATURE CITED
Morden, C.W. 1985. A biosystematic study of the Muhlenbergia repens complex (Poaceae: Eragrostideae). Ph.D. Dissertation, Texas A&M University, College Station, Texas.
Morden, C.W. & S.L. Hatch. 1987. Anatomical study of the Muhlenbergia repens complex (Poaceae: Chloridoideae: Eragrostideae). SIDA 12:347-359.
Reeder, J.R. 1967. Notes on Mexican grasses VI. Miscellaneous chromosome numbers. Bull. Torrey Bot. Club. 94: 1-17.
Phytologia (July 1995) 79(1):31-34.
SEDUM BOOLEANUM (CRASSULACEAE), A NEW RED-FLOWERED SPECIES FROM NUEVO LEON, MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A new species of Sedum, S. booleanum B.L. Turner, is described from Nuevo Leén, México where it occurs in gypsum outcrops. The taxon is red- flowered and has the habit of Villadia but the petals are separate to the base, or nearly so, suggesting a position in Sedum where it apparently has no close relatives. A photograph of living material is also presented, along with a photograph of its namesake.
KEY WORDS: Crassulaceae, Sedum, systematics, México, Nuevo Leon
Routine identification of Mexican plants has revealed the following novelty, which was called to my attention by the collectors concerned.
SEDUM BOOLEANUM B.L. Turner, spec. nov. TYPE: MEXICO. Nuevo Leon: Mpio. Rayones, Cerro Blanco, 1340 m, gypsum hillsides, forming colonies, 27 Feb 1990, Hinton et al. 20468 (HOLOTYPE: TEX!).
Succulenta erecta, perennis, radicibus fibris, 5-8 cm alta. Caules 3-4 crh diametro prope basim (ubi siccati) et papillosi. Folia (siccata) ovata, papillosa, 7-10 mm longa, 2-4 mm lata, gradtim deminuta ab imo caulis (ubi mox decidua) ad apicem per 1/2-2/3 suas longitudines superposita ut caulem celent. Flores 5-10, terminaliter dispositae in ramis_ brevibus circinatisque, infloresceniam congestam, 1-2 cm latam, circa 1 cm altam facientes. Sepala 5, ovata, glabra, circa 3 mm longa, 1.5 mm lata, latissima prope medium, libra vel paene libra ad basim, costis dorsalibus prominentibus. Stamena 5, alternata, petalis circa 3 mm longis, antheris luteis in plantis maturis. Carpella 5, 2-5 mm alta per anthesin, stylis erectis, circa 1 mm longis. Fructus matun non Visi.
31
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PHYTOLOGIA July 1995
Snes
Figure 1. Sedum booleanum, photograph of type material.
volume 79(1):31-34
Turner: New species of Sedum from México 33
Figure 2. George Boole Hinton, in the field on Rancho Aguililla, Nuevo Leén, México, at the type locality of Paronychia hintoniorum (cf. p. 38 this issue).
34 PHY TOLOGIA July 1995 volume 79(1):31-34
Erect fibrous-rooted perennial (?) succulent 5-8 cm high. Stems near base 3-4 mm across (when dried), papillose. Leaves (dried) ovate, papillose, 7-10 mm long, 3-4 mm wide, gradually reduced from the bottom of the stem (where soon deciduous) upwards, overlapping for 1/2-2/3 their lengths so as to obscure the stem. Flowers 5- 10, arranged terminal on short circinnate branches, forming a congested inflorescence 1-2 cm wide, ca. 1 cm high. Sepals 5, ovate, glabrous, ca. 3 mm long, 1.5 mm wide, widest at or near the middle, free to the base, or nearly so, with a pronounced dorsal midrib. Stamens 5, alternate with the petals, ca. 3 mm long, the anthers yellow at maturity. Carpels 5, in flower ca. 2.5 mm long, the styles erect, ca. 1 mm long. Mature fruit not available.
According to the collectors, the type was collected at the date given above, subsequently flowering at their residence on 27 June 1990 from which herbarium material was made, this constituting the holotype. The photograph (Figure 1) was also made from type matenial.
Vegetatively, Sedum booleanum much resembles species of the genus Villadia, but the flowers appear to be like those of Sedum, the petals free to the base, or nearly so.
It is a pleasure to name this very attractive red flowered Sedum for George Boole Hinton, age 5 (Figure 2), the great grandson of the well-known Mexican collector, George B. Hinton (1880-1943). In spite of his relative youth, George Boole has become a fourth generation plant collector in México. He reportedly often
accompanies both his father, George, and his grandfather, James, on various |
collection expeditions to the Sierra Madre Oriental of northeastern México. Let’s hope he continues this familial tradition.
ACKNOWLEDGMENTS
I am grateful to Gayle Turner for the Latin diagnosis, and to her and Mark |
Mayfield for reviewing the manuscript.
Phytologia (July 1995) 79(1):35-37.
A NEW SPECIES OF STEVIA FROM MEXICO
Tetsukazu Yahara & Akiko Soejima
Department of Biology, Kyushu University, Hakozaki, Fukuoka 812-81, JAPAN College of Arts and Sciences, University of Osaka Prefecture, Gakuen-machi, Sakai, Osaka 593, JAPAN
ABSTRACT A new species of Stevia, S. stolonifera is described from México.
KEY WORDS: Stevia, Asteraceae, México, systematics
The genus Stevia consisting of ca. 250 species has two centers of diversification; one in México and another in the Andes Mountains (King & Robinson 1987). Mexican representatives have been comprehensively studied by Grashoff (1972, 1974) who recognized 79 species in his unpublished monograph (dissertation in 1972) and described three additional species in 1974. Since then, five additional species have been described from México (McVaugh 1982; Turner 1992, 1993a, 1993b).
From the view point of reproductive biology, herbaceous species of Mexican Stevia are particularly interesting because agamospermy is prevalent among them and specimens with irregular pollen grains (putative apomicts) are known from 32 of the 54 species (Grashoff 1972). To elucidate the evolutionary processes of agamospermy, We are carrying out studies on sexual populations of the herbaceous species. In the course of this study, we have recognized the following novelty.
STEVIA STOLONIFERA Yahara & Soejima, spec. nov. TYPE: MEXICO. Jalisco: Sierra del Halo, near a lumber road leaving the Colima highway 7 miles SW of Tecalitlan and extending southeastly toward San Isidro: Steep slopes in mesophytic forests near summits of barrancas in pine zone 13-16 miles from
highway; 2000-2200 m; 28-30 Nov. 1959, R. McVaugh & W.N. Koelz 1169 (HOLOTY PE: TEX).
Steviae origanoideae H.B.K. similis sed rhizomis stoloniferis et foliis crassis subintegris reticulatis differt.
Stoloniferous perenial herbs to 1 m tall. Stems i-several, simple below, erect, often purplish, puberulous. Internodes as long as leaf blades. Leaves opposite, thick,
35
36 PHY TOLOGIA July 1995 volume 79 1):35-37
semisessile, oblong, 3-5 cm long, 1-2 cm wide, entire or inconspicuously crenate; apex obtuse; base cuneate; upper surface glabrous or sparsely puberulous, glandular- punctate; lower surface paler, reticulate, sparsely puberulous along veins, glandular- punctate. Inflorescence a compound corymb, the total inflorescence up to 4 cm across; branches opposite, puberulous; bracts up to 2.5 cm long, foliaceous, conspicuous. Heads 7-8 mm high, nearly sessile, in small groups ca. 1.0-1.5 cm across. Involucres cylindrical, 4.5-5.5 mm high, sparsely puberulous, sessile- glandular. Phyllaries oblong, acute at apex. Florets white, glabrous, sparsely glandular; lobes ca. 1 mm long, throat plus tube ca. 4 mm long. Achenes heteromorphic, anstate, ca. 2 mm long, dark brown, glabrous except along ribs. Pappus of the 4 adelphocarps of 3 awns, ca. 4 mm long alternating with scales less than 0.2 mm long, fimbriate.
ADDITIONAL SPECIMENS EXAMINED: MEXICO. Michoacan: Volcan Paracutin, Mpio. Uruapan, 16 Nov 1983, F.R. Barrie 553 (TEX); Coalcoman, 20.9 km al Oeste de Coalcomadn hacia Coahuayana, terraceria, 550 m, 17 Dec 1984, C.P. Cowan 4908 (TEX).
This species may be related to Stevia origanoides H.B.K. but distinctively differs in stoloniferous rhizomes and thick, nearly entire leaves reticulate beneath. The specimens of S. stolonifera were collected from western Michoacan and southeastern Jalisco where typical S. origanioides occurs. Among the three specimens cited above, Cowan 4908 has normal pollen and is regarded as_ sexual while pollen is irregular in
McVaugh & W. N. Koelz 1169 (holotype) and Barrie 553 that are therefore considered to be asexual.
ACKNOWLEDGMENTS
We are grateful to Billie Turner for reviewing the manuscript.
LITERATURE CITED
Grashoff, J.L. 1972. A systematic study of the North and Central American species of Stevia. Doctoral Dissertation. The University of Texas, Austin, Texas.
Grashoff, J.L. 1974. Novelties in Stevia. (Compositae: Eupatoneae). Brittonia 26:347-384.
King, R.M. & H. Robinson. 1987. The genera of the Eupatorieae (Asteraceac). Monographs of the Missouri Botanical Garden 22: 1-581.
McVaugh, R. 1982. Stevia zacatecana McVaugh, sp. nov. in Compositarum Mexicanarum pugillo supplementum. Contnbutions from the University of Michigan Herbarium 15:196.
Turner, B.L. 1992. Two new species of Stevia (Asteraceae, Eupatoricac) from México. Phytologia 72: 127-129.
Yahara & Soejima: New species of Stevia from México 37
Turner, B.L. 1993a. A new species of Stevia (Asteraceae, Eupatorieae) from Chihuahua, México. Phytologia 74:286-288.
Turner, B.L. 1993b. A new species of Stevia (Asteraceae) from the “Antler” region of northern Jalisco. Phytologia 74:369-370.
Phytologia (July 1995) 79(1):38-42.
PARONYCHIA HINTONIORUM (CARYOPHYLLACEAE), A NEW SPECIES FROM NUEVO LEON AND VERACRUZ, MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Paronychia hintoniorum B.L. Turmer spec. nov., is described and illustrated. It occurs in the states of Nuevo Leén and Veracruz, México, and is closely related to the widespread P. mexicana, and is well differentiated by both vegetative and floral characters. Distribution maps of the two species are provided.
KEY WORDS: Caryophyllaceae, Paronychia, systematics, México, Nuevo Leon, Veracruz
Routine identification of plants from northeastern México has revealed the |
following novelty.
PARONYCHIA HINTONIORUM B.L. Turner spec. nov. Figure 1. TYPE: |
MEXICO. Nuevo Leén: Mpio. Galeana, Rancho Aguililla, ca. 1900 m, 22 Jul 1995, Hinton et al. 25368 (HOLOTYPE: TEX!).
Paronychia hintoniorum B.L. Turner, spec. nov.; similis P. mexicana Hemsl. sed foliis oblanceolatis glabriusque, apicibus non spinulosis, et calycibus majoribus glabriusque.
Perennial sprawling or recumbent herbs from lignescent tap roots to 25 cm high, |
the stems simple, numerous and procumbent from the base of the plant, very sparsely
puberulent to glabrous; stipules white-scarious, 3-4 mm long and as wide, acute to — rounded apically. Leaves opposite throughout, gradually reduced upwards, those at |
midstem oblanceolate, 10-25 mm long, 3-7 mm wide, glabrous throughout, the apices obtuse to acute, not clearly apiculate or setose. Flowers axillary, few to numerous in
fasciculate or subfasciculate offshoots or clusters. Calyces glabrous, 3.5-4.0 mm __
long; sepals united below for 1.5-2.0 mm, the lobes 1.5-2.0 mm long, white- marginate, 1-3 nervate, acute apically. Stamens 5, small, ca 1.5 mm long, united
below into a scarious sheath. Ovary ca. 1.5 mm high, sparsely short-glandular
apically, style ca. 0.3 mm long. Fruits and seeds not available.
38
Tumer: New Paronychia from México
Figure 1. Paronychia hintoniorum, a single stem and flower from the holotype.
Se)
40 PHYTOLOGIA July 1995 volume 79 1):38-42
ig
Figure 2. Distribution of Paronychia mexicana (open circles), and P. hintoniorum (closed circles).
Turner: New Paronychia from México 41
ADDITIONAL COLLECTION EXAMINED: MEXICO. Veracruz: Mpio. de Perote, Totalco, “Orilla de camino”, 2300 m, 7 Jul 1970, F. Ventura A. 1537 (LL,US).
Paronychia hintoniorum is clearly related to P. mexicana Hemsl. but is markedly different in leaf shape and vestiture (oblanceolate, glabrous, and acute to obtuse apically, vs. linear-lanceolate, markedly hirsute and apices spinulose, respectively), and larger glabrous calyces (3.5-4.0 mm long vs. 1.4-2.0 mm) having mbbed lobes (vs. nearly ribless and pubescent).
Both of the above cited collections occur along the periphery of the known range of Paronychia mexicana and because of their marked differences are unlikely to be but forms of the latter. I wrote to the Hinton family, upon whose rancho the type collection was made (cf p. 31, this issue), asking them to examine populations at the type locality, especially to ascertain if it might not be weedy at this site. George Hinton, the grandson of the legendary México collector, George B. Hinton, responded:
I went back to the locality of the Paronychia and observed the following: it grows at the base of a limestone hill in colonies of Agave lechuguilla. In these colonies it grows with Acacia sp., Acalypha monostachya, Bahia absinthifolia, Berberis trifoliolata, Dyschoriste schiedeana, Ephedra aspera, Flourensia cernua, Gymnosperma glutinosum, Loeselia caerulea, Mortonia palmeri, Opuntia phaeacantha, Yucca filifera. Less frequently it grows as above with A. Striata instead of A. lechuguilla. Its habit is procumbent although it frequently climbs up on the plants around it. The stems are about 0.25 m; the ones I sent you are much smaller because of the difficulty of getting your hand down to the base of the plant thru the Agave. I collected about 12 sheets, with several complete plants which I will send you when dry. It only grows in the agave patches, and these are strung for about 120 m. along the base of the hill. It doesn't appear to be a weed.
He also sent additional sterile material which matched that of previous collections.
I am aware of the wide geographical gap between the only two sites known for this species (Figure 2). The Veracruz specimens are, except for their somewhat smaller leaves and more floriferous condition, almost exactly like that of type material. Label data on Ventura’s specimen report the plant as “crece en lugares despejados; abundantes”. Veracruz populations of Paronychia hintoniorum are located near populational sites of P. mexicana, the latter readily recognized by the characters alluded to in the above account. It is perhaps tempting to believe that P. hintoniorum might be but a populational growth form of P. mexicana; if so, then these must rank as among the most remarkable populational segregates within a single species to my knowledge. In any case, comparable material was not detected elsewhere among the broad range of P. mexicana examined in this study. Indeed, Chaudhri (1968) recognized two subspecies under P. mexicana, one of these with two varieties. | examined type material of these taxa and all are essentially alike (except for variation in stamen number, a variable organ set as noted by Core [1943]). Apparently, Chaudhri did not examine material of what is here called P. hintoniorum or else he would have
42 PHY TOLOGIA July 1995 volume 79(1):38-42
surely dubbed this with a name, to judge from his annotations on a broad range of specimens at F, GH, LL, TEX, US.
It is a pleasure to name this taxon for the Hinton family, upon whose property the type locality occurs.
ACKNOWLEDGMENTS
I am grateful to Gayle Turmer for the Latin diagnosis, and to Piero Delprete and Mark Mayfield for reviewing the paper. The drawing was executed by Marcia Thompson. Distributional maps (Figure 2) are based upon specimens at F, GH, LL, TEX, US.
LITERATURE CITED
Chaudhri, M.N. 1968. A revision of the Paronychiinae. Revis. Paronychiinae. 440
PP. Core, E.L. 1943. The North American species of Paronychia. Amer. Midl. Naturalist 26:369-397.
Phytologia (July 1995) 79(1):43-46.
A NEW SPECIES OF PITTOCAULON (ASTERACEAE, SENECIONEAE) FROM OAXACA, MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Pittocaulon calzadanum B.L. Turner, spec. nov. is described and illustrated. It is a tree 3-S m high with fore-shortened stems, rayless white heads and pinnate leaves and is known from only a restricted area of Oaxaca, México (Mpio. Santos Reyes Tepejillo). The species does not appear to be especially closely related to yet other taxa of Pittocaulon, and future workers might treat the species as belonging to a monotypic genus.
KEY WORDS: Asteraceae, Senecioneae, Pittocaulon, systematics, México, Oaxaca
Routine identification of Mexican Asteraceae has revealed the following novelty.
PITTOCAULON CALZADANUM B. L. Turner, spec. nov. Figure 1. TYPE: MEXICO. Oaxaca: Distr. Santiago, Juxtlahuaca, Mpio. Santos Reyes Tepejillo, 3
km N of Santos Reyes Tepejillo “a Corral de Piedra” (17 27’ N x 97 57’ W), ca.
1770 m, 21 Apr 1995, J.I. Calzada 19872 (HOLOTYPE: TEX; Isotypes: to be distributed).
Arbor 3--5 m alta. Caules terminales non angustati, cicatricibus foliorum delapsorum notati; cortex semisucculenta, textura intenor lignea, cellulis resiniferis inspersa. Folia alterna; petioli 3--5 cm longi; laminae latae, ovato- ellipticae, pinnatinerviae, 10--20 cm longae, 3--7 cm latae, tomentosae; margine irregulariter lobatae. Capitulescentia paniculato-corymbosa 50--100- cephala, aut triangulata aut ovata, pedunculis ultimis tomentosis, saepius 3--8 mm longis. /nvolucra cylindro-campanulata; bracteae interiores 8, lanceolatae, 6--7 mm longae, 1--2 mm latae, tomentosae aetate glabratae, marginibus chartaceis candidis. Receptaculum circa 2.5 mm longum, alveolatum, palcis nullis. Flosculi radiantes nulli. Flosculi disci cujusque capituli 8; corollae candidae, glabrae, 7--9 mm longae, lobis irregularibus 2--3 mm longis, ut
—
videtur lactiferis. Achenia (immatura) columnania, circa 2.5 mm longa, glabra;
43
Ad PHY TOLOGIA July 1995 volume 79 1):43-46
carpopodia bene evoluta, annulata; pappi setae capillares, numerosae candidae, 6--7 mm longae, marginibus (praecipue inferne) scabridiusculae.
Tree 3-5 m high. Stems (terminal), abruptly fore-shortened, the bark semisucculent, and the interior with hard woody tissue, the latter suffused with resinous cells. Mature leaves alternate, densely velvety-tomentose, deciduous at anthesis, leaving a pronounced scar; petioles 3-5 cm long; blades broadly ovate to deltoid, pinnately nervate, 10-20 cm long, 3-7 cm wide, moderately tomentose on both surfaces, the margins irregularly lobate. Capitulescence a terminal, ovoid or trianguloid, corymbose panicle of numerous (50-100) heads, the ultimate peduncles tomentose, mostly 3-8 mm long. Involucres cylindrocampanulate, the inner bracts 8, lanceolate, 6-7 mm long, 1-2 mm wide, tomentose, glabrate with age, the margins white-chartaceous. Receptacle ca. 2 mm across, epaleate, alveolate. Ray florets absent. Disk florets 8-10 per head; corollas reportedly white, glabrous, 7-9 mm long; tubes 4-5 mm long; the throat 2-4 mm long, irregularly lobed, the lobes 2-3 mm long, apparently lactiferous. Achenes (immature) columnar, ca. 2.5 mm long, glabrous; carpopodia well-developed, annulate; pappus of numerous white capillary bristles 6-7 mm long, the margins minutely scabridulate, especially below.
Label data describe the tree as 3 m high having white corollas and yellow stamens. It also states that the plant occurs in tropical deciduous forests and is “mass bien escasa”. Calzada, who collected the type, revisited the site and tree concerned in July of 1995 (Calzada s.n. [TEX]) so as to collect mature leaves (not shown in Figure 1); leaf measurements in the present description were obtained from this collection. José Panero, who also visited the site concerned, states (pers. comm.):
The new Pittocaulon is a very remarkable plant. It is a small tree of the tropical deciduous forest. It can grow to 5 m tall. The leaves are kind of gray- green, somewhat silvery. The plant is an inhabitant of rocky, limestone outcrops. | first saw the plant in March of this year and asked Ismael [Calzada] to collect it. At first, I thought it was going to be a weird Parthenium, later | was surprised to see it was a Senecio.
It grows with frexinus purpusii, Conzaitia multiflora, Xylosma flexuosum, Schoepfia angulata, Erythrina petrea, Croton sp., Jatropha sp., Bunchosia trifoliata, Quercus glaucoides, among others.
As noted by Panero, this is a remarkable Pittocaulon, the latter a generic segregate from Senecio first proposed by Robinson & Brettell (1973), who recognized five species in the genus, all confined to south-central México. Jeffrey (1992) also recognized the genus as distinct, emphasizing its subumbellate inflorescences, cortical resin ducts and palmately veined leaves. Barkley (1985), however, retained Pittocaulon in Senecio (s.\.) although he now accepts its generic status (pers. comm.). Pittocaulon calzadanum has a corymbose-paniculate capitulescence, pinnately veined leaves, eradiate heads, and relatively deeply lobed, white corollas. In short, a very different looking Pitfocaulon than those described to date. When I first examined the plant [ took it to be, because of its narrow white discoid heads, a species of Digiticalia, but the woody habit, abruptly foreshortened stems, semisucculent bark and attainment of anthesis before the leaves appear, strongly suggest that it belongs to the Pittocaulon
Turner: New Pittocaulon from México
Xe
i\ 7
ee iy.
~/ ) YY, yp
Figure 1. Pittocaulon calzadanum, from holotype.
45
46 PHYTOLOGIA July 1995 volume 79(1):43-46
alliance, although some workers, because of its differing capitulescence, pinnately veined leaves, narrow involucres, and rayless white corollas might treat it as a monotypic genus.
It is a pleasure to name this remarkable new species for J. Ismael Calzada, premier collector working out of UNAM, who first collected the taxon concerned.
ACKNOWLEDGMENTS
I am grateful to my wife, Gayle Turner, assisted by Rupert Barneby, for the Latin diagnosis, and to José Panero for calling the plant to my attention. Rupert Barneby and José Panero reviewed the manuscript. The illustration was provided by Marcia Thompson.
LITERATURE CITED
Barkley, T. 1985. Infrageneric groups in Senecio s.l., and Cacalia s.1. (Asteraceae: Senecioneae) in Mexico and Central America. Bnittonia 37:211-218.
Jeffrey, C. 1992. The tribe Senecioneae (Compositae) . . . Notes on Compositae: VI. Kew Bull. 47:49-109.
Robinson, H. & R.D. Brettell. 1973. Studies in the Senecioneae (Asteraceae). I. A new genus Pittocaulon. Phytologia 26:451-453.
Phytologia (July 1995) 79(1):47-50.
TWO NEW VARIETIES OF HEDEOMA PALMER! (LAMIACEAE) FROM NORTHEASTERN MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Two new infraspecific taxa of Hedeoma palmeri are described: _ var. santiagoanum B.L. Turner, var. nov., and var. zaragozanum B.L. Turner, var. nov. The former is largely confined to central Nuevo Le6én (Mpio. Villa Santiago) and closely adjacent Coahuila; the latter is largely confined to southern Nuevo Leén (Mpio. Zaragozana) and closely adjacent Tamaulipas (Mpio. Hidalgo). Their relationships to the two other varieties of H. palmeri (var. palmeri and var. galeanum) are discussed, and the distribution of each in the area concerned is depicted.
KEY WORDS: Lamiaceae, Hedeoma, systematics, México
Hedeoma palmeri Hemsl., a member of the subgenus Poliomenthoides of Hedeoma, is typified by materials from San Luis Potosi. It was treated by Irving (1980) as a single variable species, although he called attention to populations of diversely tomentose individuals from Nuevo Leén which Turner (1991) subsequently described as var. galeanum Turner, sufficiently distinct so as to be placed in its own subspecies. Additional collections from the more montane regions of Coahuila, Nuevo Ledn, and Tamaulipas have revealed two additional infraspecific morphogeographical taxa that appear to warrant names, and these are described herein. Both appear to be closely related to the var. palmeri and are positioned within the
subsp. palmeri. A key to these four varieties is provided below, along with a map showing the distribution of each (Figure 1).
KEY TO THE SUBSPECIES AND VARIETIES OF H. PALMERI
1. Leaves bicolored, the lower surfaces densely white-pilose; mostly gypsum outcrops in the vicinity of Galeana, Nuevo Leon (subsp. galeanum). ...........66...
Beds ohia echme a Mtoe a eae Sastinte daittecs eta cre sars deite Yaw ae pee dala ee anes Bet A ane ded var. galeanum 1. Leaves not bicolored, about equally green on both surfaces, the lower surfaces moderately to sparsely hirsute (subsp. palmeTi)..........0...000.000cc cece cece nece ees (2)
47
48 PHYTOLOGIA July 1995 volume 79(1):47-S0
2. Calyx lobes with spreading hairs 0.4-0.6 mm long; midstem leaves mostly 1-2
cm long; San Luis Potosf, Guanajuato, Querétaro, and Hidalgo. ... var. palmeri
2. Calyx lobes glabrate or with hairs appressed, if somewhat spreading then the
hairs 0.2 mm long or less; midstem leaves mostly 2-4 cm long; Nuevo Leén and closely adjacent Coahuila and Tamaulipas..................cceeeceeeeeee eens (3)
3. Calyces mostly 4.5-5.5 mm long, the lobes greenish with short spreading hairs 0.1-0.2 mm long; central Nuevo Leon and closely adjacent Coahuila.
cdadiventanednd Mla beaial Sacvensaueu ee sas cucone sees se iedansite es var. Santiagoanum
3. Calyces mostly 5.5-6.5 mm long, the lobes reddish to purplish, glabrate or nearly so (any hairs minute and appressed); southern Nuevo Leén (Mpio. Zaragozana) and closely adjacent Tamaulipas. ............. var. zaragozanum
HEDEOMA PALMERI Hemsl. var. SANTIAGOANUM B.L. Turner, var. nov. TYPE: MEXICO. Nuevo Leén: Mpio. Villa de Santiago, between Las Ajuntas
and Potrero Redondo, abundant in pine forest, 15 Aug 1939, C.H. Muller 2702 (HOLOTYPE: TEX!).
H. palmeri Hemsl. var. santiagoanum B.L. Turner, var. nov., similis H.
p. var. palmeri sed habens folia majora et calyces parviores, hirsutos, 0.1-0.2 mm longos.
ADDITIONAL SPECIMENS EXAMINED: MEXICO. Coahuila: Mpio. Arteaga, road from Los Linos to El Cercado, 2095 m, 29 Jul 1995, Hinton et al. 25446 (TEX). Nuevo Leén: Mpio. Villa de Santiago, Pasaje de los Osos al Pte. del Yebanis, Santiago, 19 May 1966, Marroquin 1311 (TEX); ca. 18 km al S de
Monterrey, 16 Sep 1966, Marroquin 1383 (TEX); Cafion la Boca (100 19’ W x 25
24’ N), 1600 m, 10 Sep 1983, Villarreal 2341 (TEX); 5 km SE of La Tninidad, in Canyon Cebolla, 2000 m, 8 Aug 1988, Patterson 6321 (TEX); Mpio. Montemorelos, trail up Sierra Cebolla from La Trinidad, 1600 m, 6 Sep 1992, Patterson 7163 (TEX).
This variety is distinguished from var. palmeri by its relatively small calyces, the lobes of which have a short spreading, pubescence, and its relatively large leaves. Occasional specimens appear to weakly approach var. galeanum (e.g., Patterson 71631), but overall the vestiture of such plants is more like that of var. palmeri.
HEDEOMA PALMERI Hemsl. var. ZARAGOZANUM B.L. Turner, var. nov. TYPE: MEXICO. Nuevo Leon: ca. 30 mi NE of Dr. Arroyo along Hwy 29 along the first pass; “open pastureland and heavily forested N-facing slopes . . . infrequent perennial, in clearings”, 24 02' N, 99 58’ W, ca. 6000 ft, 9 Sep 1971, James Henrickson 6628 (HOLOTYPE: LL!; Isotype: MEXU).
H. palmeri Hemsl. var. zaragozanum B.L. Turner, var. nov., similis H. p. var. palmeri sed habens folia majora et lobos calycum pacne glabros, rubellos.
Tumer: New Hedeoma from México 49
tOl~ te
et or SAN A iolei2' 22°06'
Figure 1. Distribution of varietics of Hedeoma palmeri in northeastern México: vat. galeanum (closed circles), var. palmeri (closed triangles), var. santiagoanum (open
re var. zaragozanum (open triangles). Localities vouchered by material at | |
50 PHY TOLOGIA July 1995 volume 7% 1):47-50
ADDITIONAL SPECIMENS EXAMINED: MEXICO. Nuevo Leén: Mpio. Zaragoza, Cerro El Viejo, 2400 m, 7 Jul 1992, Hinton et al. 22103 (TEX); Cerro H Viejo, 2200 m, 29 Jul 1992, Hinton et al. 22245 (TEX); Cerro El Viejo, 2405 m, 12 Oct 1992, Hinton et al. 22486 (TEX); Los Potreritos, 1390 m, 2 Aug 1994, Hinton et al. 2454] (TEX). Tamaulipas: Mpio. Hidalgo, Los Caballos, 1750 m, 21 Sep 1994, Hinton et al. 24824 (TEX).
The var. zaragozanum 1s a distinctive populational element of the Hedeoma palmeri complex and, so far as known, is largely confined to the environs of Cerro El Viejo, mostly between 1400 to 2400 meters where it occurs in pine-oak woodlands. It is readily distinguished from var. palmeri by its nearly glabrate, reddish-hued calyx lobes, which characters also serve to distinguish it from var. santiagoanum. Future workers might wish to treat the taxon as a monotypic element of 1ts own subspecies.
ACKNOWLEDGMENTS
I am grateful to Gayle Turner for the Latin diagnoses, and to Mark Mayfield and Piero Delprete for reviewing the paper.
LITERATURE CITED
Irving, R. 1980. The systematics of Hedeoma (Labiateae). Sida 3:278-294. Turner, B.L. 1991. Novelties and new combinations in Mexican Hedeoma (Lamiaceae). Phytologia 71:32-37.
Phytologia (July 1995) 79(1):51-57.
A NEW SPECIES OF CERATOZAMIA (ZAMIACEAE) FROM OAXACA, MEXICO WITH COMMENTS ON DISTRIBUTION, HABITAT, AND RELATIONSHIPS
Jeffrey Chemnick & Timothy J. Gregory
114 Conejo Road, Santa Barbara, California 93103 U.S.A.
ABSTRACT
Ceratozamia whitelockiana spec. nov., from Oaxaca, Meéxico is described and illustrated. The species differs from others in the genus in the upright habit of its few, large, glaucous, pea-green leaves with comparatively long petioles and relatively small megastrobili and microstrobili. It is most closely related to Ceratozamia miqueliana Wendland (Vovides et al. 1983; Stevenson et al. 1986); having similar cones, caudex, and leaf color but differs in the habit, size, and shape of the leaves. Ceratozamia whitelockiana is known only from the drainage of the Rio Valle Nacional, at elevations from 335 to 975 m.
KEY WORDS: Ceratozamia, México, Oaxaca, Zamiaceae, systematics
CERATOZAMIA WHITELOCKIANA Chemnick & Gregory, spec nov. TYPE: MEXICO. Oaxaca: Vicinity of Metates, south of Valle Nacional, May 1995, Chemnick & Gregory 5 (HOLOTYPE: HNT; Isotypes: to be distributed to FTG & XALU. Cultivated specimens at GannaWalska Lotusland, Santa Barbara, California; Mildred Mathias Botanic Garden, UCLA, California; and UCSB Greenhouse, Santa Barbara, California.
Truncus semihypogaeus, ad 30 cm altus; cataphylla lanata, triangularia, 5 cm longa basi 5 cm lata: folia pauca, usque 5, glauca; petiolus teresve, 2.0-2.5 m longus, parte infima dilatatus, pauca spinis armatus; rachis subteres, supra bisulcata, in dimido inferiore, paucis spinis armata, supra fere inermis vel inermis, in cuspidem 10-25 mm longam excurrens; foliala opposita vel subopposita, 30-40 juga, lanceolata vel falcata, 30-50 cm longa, 30-38 mm lata, papyracea, pisacea, tenula, basi attenuata, apicem attenuata, margine integerrima, revoluta; 22-27 nervis moderata; strobilus microsporangiatis lineari-cylindricus, 26-28 cm longus, 15-28 mm_ latus; pedunculus tomentosus, 20-30 mm longus, 11-15 mm latus; strobilus megasporangiatis cylindricus, apice mucronatus, 14-18 cm longus, 7.5-10.0 cm_ latus; pedunculus 1-2 cm longus.
5]
ay PHY TOLOGIA July 1995 volume 79(1):51-57
MORPHOLOGY
Stem solitary, semihypogeous, moderately short (20-30 cm), cylindric (12-18 cm in diameter), covered by rough, irregular persistent leaf and cataphyll bases, brownish-red; cataphylls wrinkled, stipulate, triangular, densely white hairy at crown, irregularly arranged on lower portions of stem, 5 cm wide and 5 cm long; leaves 2.0- 2.5 m long, usually in whorls of 2-4, recently-emerged and juvenile leaves glaucous on both surfaces, light pea-green, older leaves glabrous, uniformly medium-green on both surfaces, adult plants with up to 2 previous whorls of leaves; petiole 1.00-1.25 m long, terete with an expanded base, 15 mm in diameter at base and tapering gradually to 8 mm at the first leaflet, sparsely armed with simple spines (1-3 mm), spines more densely distributed proximally and becoming sparse distally; rachis nearly straight, subterete, very sparsely armed on proximal 25%, ending in conical-linear apex 10-25 mm long and unarmed; adaxial surface is flattened and shallowly bisulcate with leaflets inserted in the paired grooves up to 5 mm apart, the paired grooves arising distally to the first pair of leaflets; leaflets linear lanceolate to falcate, papyraceous, the median leaflets 30-50 cm in length, gradually attenuate, 30-38 mm in width with 22-27 veins slightly raised on abaxial surface, 30-40 “pairs” inserted on 25-50 mm centers, opposite to sub-opposite, 9-12 mm wide at point of attachment on rachis, margins are slightly revolute and turned upward, basal 25-30% of leaf keeled becoming flattened distally, leaflets gradually reduced in length towards apex; microsporangiate strobilus elongate-conical, solitary, 26-28 cm in length, 28 mm in diameter at base, 15 mm in diameter distally, mucronate, peduncle 20-30 mm in length and 11-15 mm in diameter, tomentose to wooly; microsporophylls 8 mm wide and 3 mm long, sporangia in a single patch, olive green; megasporangiate strobilus cylindrical to ovoid with a large apiculum, solitary, overall length 14-18 cm and diameter 7.5-10.0 cm at maturity, apiculate cap 1.5-3.0 cm in length and 3-5 cm in width, megastrobilus borne on a short peduncle 30-38 mm long and 18-20 mm wide; megasporophyll length 2.5-3.0 cm, sporophyll face 3.5-5.0 cm wide and 17-23 mm long, inner face somewhat glabrous except for the moderately rolled margins which are gray tomentose; sporophyll horns divergent to either side of the sporophyll up to 10 mm long, only slightly raised from the sporophyll face, outer edges grey and tomentose, horns joined by a wrinkled raised edge; megastrobilus with short purple hairs sparsely scattered on sporophyll face and sarcotesta where exposed between megasporopyhlls which are widely separated at maturity by the fully-developed seeds; sarcotesta white, soon turning brown as it ripens; 31-33 mm long, 25-27 mm wide; sclerotesta irregular, ovoid, tan, 24-26 mm long, 18-20 mm wide, smooth with 8-9 indistinct longitudinal ridges.
Etymology: The species is named to honor Mr. Loran Whitelock of Los Angeles, CALIFORNIA for his remarkable dedication and contribution to cycad biology and awareness throughout the world.
DISTRIBUTION AND HABITAT Ceratozamia whitelockiana is known only from the drainage of the Rfo Valle
Nacional in montane tropical forest within the range of 335-973 m, but occurs more commonly at lower elevations (335-600 m). Habitat consists of very steep slopes
Chemnick & Gregory: New species of Ceratozamia 53
with small pockets of remnant primary forest now covered mostly by coffee and banana groves and secondary growth. The patchy canopy consists of emergent trees to 40 m covered with epiphytes. Ceratozamia whitelockiana occurs on heavily shaded east- and west-facing slopes in primary forest with Chamaedorea sp., Geonoma sp., Melastoma spp., Acanthus sp., Ficus sp., Begonia sp., Selaginella sp. Soil is light- colored crumbly, rocky clay with outcroppings of sedimentary rock. Ceratozamia whitelockiana growing in exposed, deforested areas have extremely bleached, yellow leaves. The entire locality is rapidly being cleared and burned and thus this cycad must be considered endangered. In our most recent survey of the locality in May, 1995 we found approximately 250 plants during 3 days of field work. The same areas were visited several times in 1979, 1980, and 1981 and the population of Ceratozamia whitelockiana was considerably larger then, perhaps by twice as many individuals. Since this cycad is seldom seen in collections, it appears that habitat destruction is the greatest threat to its existence. The more inaccessible reaches of the Rio Valle Nacional drainage are likely to contain many pocket populations of Ceratozamia whitelockiana but the rapid rate of deforestation will soon reach areas that are currently inaccessible. In May 1995, the smoke from clearing fires was intense and recently cleared fields, as evidenced by still fresh, charred remains, were spread throughout the drainage like a patchwork quilt. This cycad does not seem to persist in open situations or in second growth forest for very long. The only plants we found in cleared areas were artificially maintained by local farmers and appeared bleached and chlorotic.
RELATIONSHIP TO OTHER SPECIES OF CERATOZAMIA AND DISCUSSION
The current state of taxonomy within the genus Ceratozamia is confused, ambiguous, and incomplete. Three of the most widespread taxa, both in the wild and in cultivation, C. mexicana Brongniart (Vovides et al. 1983; Stevenson et al. 1986), C. robusta Miquel (Vovides et al. 1983; Stevenson ef al. 1986), and C. latifolia Miquel (Vovides et al. 1983; Stevenson ef al. 1986) are based on vague and obscure descriptions and neotypifications. Locality information is either non-existent or too generalized. Important morphological data such as male and female cone descriptions are incomplete or omitted. When considered from historical perspective, the neotypifications assign the above specific epithets to localities of Ceratozamia which do not necessarily correspond to the most likely localities where the original authors and collectors might have been in the mid 1800’s when access into México was much more restricted than today. The many isolated populations, forms, ecotypes, and varieties of the large-leaved Ceratozamia have been treated within the above three taxa with apparently little regard for valid character differences that in some cases might Suggest separation at the species level.
Ceratozamia_ whilelockiana is distinguished from the other large-leaved Ceratozamia as follows: C. mexicana has smooth, dark brown, globose stems to 1 m tall and 20 cm in diameter; numerous, glabrous, dark-green, arching leaves which are heavily armed with numerous spines; megastrobili which are on average 35 cm long and 12 cm in diameter borne on a peduncle 10 cm long; microstrobili which are on average 38-43 cm long and 7-8 cm in diameter borne on a peduncle 8-10 cm long and 2.5 cm in diameter. Ceratozamia whitelockiana has rough, cylindrical reddish stems that are much smaller than C. mexicana and its few, sparsely-armed, upright, glaucous, pea-green leaves with long petioles are strikingly different than the leaves of C. mexicana as are the much smaller male and female cones of C. whitelockiana.
54 PHYTOLOGIA July 1995 volume 79(1):51-57
>
~=*
,
he ade
GROWTH HABIT
Seamer Do9s Fig.1-Ceratozamia whitelockiana. a, megasporangiate strobilus at pollination b, microsporanaiate strobilus after shedding, pollen. c, arowth habit.
| | |
Chemnick & Gregory: New species of Ceratozamia 55
Ceratozamia latifolia stems are globose, light brown, and frequently sucker, especially in cultivation; leaves are 90-150 cm; leaflets are coriaceous, unequally attenuate, slightly overlapping, 20-30 cm long and 33-43 mm wide. Ceratozamia whitelockiana stems are solitary, even in cultivation; leaves are 2.0-2.5 m long; leaflets are papyraceous, 30-50 cm long, 30-38 mm wide, and not overlapping.
Ceratozamia robusta has very large stems to 1.5 m, numerous, heavily-armed, glabrous dark-green leaves to 2.25 m, megastrobili on average 38 cm long and 15.25 cm in diameter borne on a peduncle 7.5 cm long and 28 mm in diameter, microstrobili 45 cm long and 8 cm in diameter. Ceratozamia whitelockiana is a much different plant than C. robusta based on many characteristics, but especially in the detail of the male and female cones which, as reproductive structures, are characters of the highest weight.
We stress the differences between these two taxa because in Stevenson ef al. (1986), figure 7, indicates three populations of Ceratozamia robusta in north central Oaxaca. One of these populations appears to occur in the drainage of the Rio Valle Nacional. Similarly, in their paper on the distribution of Ceratozamia, Moretti et al. (1980), figure 1, identifies several populations in northern Oaxaca belonging to the C. mexicana complex. The localities are not described in the detailed text that precedes the illustration but the placement of one of those populations would appear to be in the Rio Valle Nacional drainage. We have searched extensively for other Ceratozamia in the drainage of the Rio Valle Nacional, from the municipality of Valle Nacional up to 2,200 meters but have only found C. whitelockiana.
It is noteworthy that Ceratozamia whitelockiana, C. robusta, and C. mexicana retain their respective phenotypes even when cultivated for many years under varying conditions. We have grown all three taxa for over fourteen years and found that cultivated individuals are easily distinguished. We make this observation with respect to remarks in Stevenson et al. (1986a) regarding the validity of C. microstrobila Vovides & Rees. Stevenson et al. (1986a) assert that C. latifolia and C. microstrobila, are the same because “when cultivated in conditions of high moisture and deep shade, plants assignable to C. microstrobila ‘turn into’ plants of C. latifolia. Conversely, when plants assignable to C. latifolia are exposed to conditions that are dry with high light intensity, they ‘turn into’ plants of C. microstrobila. In our opinion, the plants that have been referred to C. microstrobila are nothing more than forms of C. latifolia that are phenotypical expressions of environmental conditions. Therefore, we recognize only C. latifolia and consider C. microstrobila to be a synonym.” However, a careful character examination of these two taxa reveals a host of differences that justify separation at the species level. We have similarly cultivated both taxa for seventeen years and have observed cultivated specimens of numerous individuals of both taxa in other gardens and collections, and have never seen the alleged change of phenotypic expression whereby one taxon “turned into” the other, regardless of whether the individuals were grown in full sun, heavy shade, or even in the greenhouse. Therefore we reject the assertion that C. latifolia and C. microstrobila are synonymous but rather that each is a distinct species. Similarly, we reject any ad hoc hypothesis that C. whitelockiana is merely an ecotype of C. robusta or C. mexicana.
Ceratozamia miqueliana has 7-10 leaves that are distinctively different from those of C. whitelockiana. The leaflets are fewer (about 15 pairs), wider (60-65 mm),
56 PHY TOLOGIA July 1995 volume 79(1):51-57
unequally and abruptly attenuate. The petiole is heavily armed with long, curved spines which is in stark contrast to that of C. whitelockiana, which is much longer overall and sparsely armed with much shorter spines. However, there are many similarities between C. whitelockiana and C. miqueliana that suggest an affinity between the two taxa. Both species have subterranean to shortly arborescent stems of similar size, shape, and color; each with rough, wrinkled, irregular leaf bases and brownish-red cataphylls. Both species have juvenile and adult emergent foliage which is a very distinctive glaucous, pea-green color which matures into papyraceous, slightly revolute leaflets. The mature foliage retains the glaucous coating for some time, eventually giving way to a more glabrous, medium green color in old age. Male and female cones of both taxa are of similar size. The megastrobilus in C. miqueliana averages 11 cm long and 6.5 cm wide and is borne on a short peduncle 30 mm long. In C. whitelockiana, it averages 15 cm long and 8 cm wide and is borne on a short peduncle 30-38 mm long. The microstrobilus is 20 cm long and 4.5 cm wide in C. miqueliana and 26 cm long and 25 mm wide in C. whitelockiana. The closest population of C. miqueliana to C. whitelockiana is approximately 150 km.
Since cytological and genetic evidence currently does not yield any measurable character differences upon which to base species differentiation within the genus (Walters et al. 1991), classic taxonomic consideration of characters and weighting of those characters is our basis for conferring specific status to Ceratozamia whitelockiana and assigning it to the “miqueliana group” which also includes the various forms of C. miqueliana and C. euryphyllidia Vazquez Torres, Sabato, & Stevenson. It 1s our hope that workers will continue to investigate Ceratozamia in detail to determine the disposition of the many populations and types currently being lumped into vaguely conceptualized and incompletely described taxa that generate confusion and uncertainty rather than create the order, predictability, and sense that responsible taxonomy 1s supposed to serve.
ACKNOWLEDGMENTS
We are grateful to Sherwin Carlquist and Dieter Wilken for reviewing the manuscript and providing valuable assistance. We are indebted to Jim Melli for providing the illustrations and to Loran Whitelock for providing details of cone dimensions.
LITERATURE CITED
Moretti, A., M. Vazquez-Torres, & S. Sabato. 1980. The distribution of Ceratozamiu Brongn. (Zamiaceae). Delpinoa 21:13-21.
Stevenson, D.W., S. Sabato, & M. Vazquez-Torres. 1986a. A new species of Ceratozamia (Zamiaceae) from Veracruz, Mexico with comments on_ species
relationships, habitats, and vegetative morphology in Ceratozamia. Brittonia 38: 17-26.
| Chemnick & Gregory: New species of Ceratozamia a7
Stevenson, D.W. & S. Sabato. 1986. Typification of names in Ceratozamia | Brongn., Dion Lindl., and Microcycas A. DC. (Zamiaceae). Taxon 35:578-584. —Vovides, A.P., J.D. Rees, & M. Vazquez-Torres. 1983. Zamiaceae in Flora de | Veracruz., Fasiculo 26, INIREB, Xalapa, Veracruz, México.
Walters, T.W. & D.S. Decker-Walters. 1991. Patterns of allozyme diversity in the | West Indies cycad Zamia pumila (Zamiaceae). Amer. J. Bot. 78:436-445.
Phytologia (July 1995) 79(1):58-64.
Rexford F. Daubenmire (1910-1995)
“Dauby” was the usual appellation applied by graduate students to Dr. Daubenmire, Professor at Washington State University, Pullman, Washington, during the years 1950-1953 while I was working under the aegis of the late Prof. Maron Ownbey (1910-1974) in the area of plant systematics.
I first read about Dauby's death in the obit section of the New York Times (8 September 1995). This was a short but well-wnitten account of his professional life and contributions to ecology. Unfortunately it conveyed very little about the man himself. Indeed, most scientists are largely remembered by bnef obits prepared by their professional colleagues in which their lives are summed up as lines culled from their latest CV. Subsequent biographers have to invent their other attributes, especially for scientists who are reluctant to write personal letters or expose their psyches.
Perhaps, for many workers, that is as it should be. But I feel otherwise. Indeed, the only previous obits to have been penned by me (Turner 1972, 1975) were both highly personal, although both were solicited. In these I wished to portray the inner essence of the person, his weaknesses and strengths, beauties, foibles, whatever. Whether or not I succeeded in these endeavors is not so important as the attempt, for these will surely provide future biographers with at least some material by which to humanize their subjects. To me, at least, an individual's work cannot be understood solely by publications and their contents.
The present obit is obviously unsolicited. It is written simply because I thought Dauby was a fine researcher, a commendable undergraduate teacher, and a remarkable professional. Certainly, any deep appreciation I have of the field of ecology comes from my enrollment in all of the courses he taught in botany at W.S.U. during the ume of my attendance at that institution. These included autecology, synecology, field ecology, and plant geography; I also served as his T.A. in undergraduate courses in general botany, sitting in on all of his freshman lectures on that subject.
Dauby was, for the most part, a calm, even-tempered, rather handsome man. He wore a full mustache above a seemingly perpetual Gioconda-like smile (unusual for most competitive males of my acquaintance, at the tme or since). Even when exceedingly irritated he retained that sphinxious grin: along with his expressive eyes, and thin lips, he exuded a detached serenity that belied his inner turmoils.
At the tme I knew him, during the pnme of his professional career, aged 40-43, Dauby was lean and well-proportioned, about 5 feet ten or so and perhaps 150 pounds. He wore an academic costume to all of his formal lectures: well-creased pants, a professorial tweed coat with leather covered elbows, bowtie, and freshly polished shoes. I remember this well, for the late Art Cronquist (1919-1992), his colleague at the time, for whom I was also a T.A., dressed in just the opposite
58
Tumer: Obituary for R.F. Daubenmire 59
fashion, usually a slip-over, much-abused sweater, baggy pants and coat, that looked slept in, occasionally an off-angled mussed tie, and large military-type shoes in various stages of repair. In short, Dauby believed in appearances; Art did not. Like their attires, they were antagonists, but most of the antagonism drifted downward from Dauby. I can still recall a brief statement or two made to himself by Dauby upon hearing the approach of Art along the lower floor of the botany building as Dauby ascended the stairs leading to the second floor, myseif along his side. Cronquist, with his six foot eight inch Swedish frame, would usually enter the building with a large booming voice singing whatever song entered his mind, operetta or ballad. On this particular day it was “Oh, she jumped in bed and covered up her head and said | couldn't find her. . ..” and carried on through the whole verse (which I myself sang upon occasion, having learned it as a teenager in Texas). Dauby paused for a second, looking at me with grimaced eyes and no smile, saying “That man! God, that man!” Then he trudged on up to the second floor with a perplexed expression.
In Dauby's formal undergraduate lectures he spoke at a slow clip, very precisely, everything biological presented as black or white, with little, if any, gray areas. He drew precise figures on the chalk board and labeled their parts with easily read names. Excellent teacher, answering questions from the floor briefly but adequately.
In upper undergraduate and graduate level courses he was less effective. For example, in autecology, having written the text himself, Dauby did not feel it necessary to lecture on the subject, rather he would meet his classes so as to answer questions about any ambiguities in the text chapters, which we were all expected to have pored over prior to attendance. Most of these classes lasted 10-15 minutes, though sometimes they were prolonged by an overly querulous student. This permitted him to shorten his teaching load and retire to his office (door nearly always closed) so that he might get on with his research or textbook wnitings.
Dauby took a different tack for his course in synecology (lectures from which he was hoping to develop a text on the subject, and did). He often became rather enlivened by his own spontaneous insights into the field of community ecology, holding forth on succession, its history, comparing community classification to systematic classification, but always with the admonition to accept such comparisons as “analogous to,” not “the same as,” efc. At such times he could be brilliant, but, sadly, he often took himself too seriously. Indeed, I think he did so much of the time, for he seemed to lack a sense of humor, at least where his utterances about ecology were concerned.
To give an example: holding forth on the contribution of F.E. Clements to the field of ecology, especially as regards climax concepts, Dauby suddenly became reiterative, stating that the trouble with American ecology was that everything important in the field of synecology was discovered by Clements, so much so that one might characterize its history as “Before Clements, B.C., B.C., B.C. . ..” he finally added, “before Christ” with a full grin, Cheshire-like, something unusual for him; clearly, he much appreciated his effective presentation and original commentary. The class (about 60, mostly graduate students from several disciplines, for Dauby's classes were very popular) laughed appreciatively, including myself, but I raised my hand almost immediately after his riveting delivery and interjected rather loudly, and with much glee, and some laughter, “I now take it we're entering A.D., after Daubenmire!”
60 PHY TOLOGIA July 1995 volume 79(1):58-64
Instead of appreciating my joshing spontaneity, he became suddenly furious. Red faced and with grin-turned scowl, he ordered me out of the classroom “Out,” he said, “Get out.” The class was bewildered, for they had all chortled loudly at my retort, so was I, for I never meant to be disrespectful, merely entertaining, attempting to add to the pedagogic verbalization he’d seized upon.
I did leave the class as instructed, very embarrassed of course, although pleased that my peers had perceived my spontaneous remarks as somehow appropriate. Afterwards I tried to apologize to Dauby, but he would have none of it, although he did relent and permitted me back in his class the following week.
My interpersonal relationships with Dauby were largely developed because of my interest and background in plant systematics. I believe he sought out my conversation, both during field courses in connection with his formal classes in synecology, where sack lunches were the rule, and following this or that class lecture in which allusions were made to the views of systematists generally. I believe he mostly wanted feed- back on his many attempts to make plant community classification “analogous” to organismal classification. “But they are very different,” | would assert, “Community ecologists do not have evolutionary theory as a direct underpinning by which to arrange and classify.” “Ah,” he would respond, “communities evolve, they are made up of plants and animals, all of which coevolve,” etc. And he would usually wrap up the conversation pretty quickly with terse sentences that made his points; (Dauby would have made an excellent tnal lawyer speaking before an educated jury). Deep down, I think he knew these analogies were basically misleading, dishonest even, for he not only was well aware of Gleason’s (a systematist!) individualistic concepts on community structure but, at the tme also coexisted with Prof. R.H. Whitaker, his nemesis at Washington State University during my formative years there.
Like most academic professionals, Dauby had considerable concern about his standing in the field of plant ecology, especially as perceived by his peers. I remember well his deep sense of betrayal by the ecological community, if not the man, when the article by Frank Egler, “A commentary on American plant ecology, based on the textbooks of 1947-1949,” first appeared in the October, 1951 issue of Ecology (32: 673-695). Egler, a very perceptive, erudite, human, to judge from his well-turned article, compared the ecological texts of F.E. Clements, Dynamics of Vegetation, 1949; H.J. Oosting, The Study of Plant Communities, 1948; and Daubenmire, Plants and Environment (A Text Book of Plant Ecology). Not only did Egler compare these texts (as indicators of the state of American plant ecology and its development over half a century), he also commented rather freely on the psyches of the authors concerned, especially as related to their academic beginnings. In preparing the present “obit”, I re-read Egler’s article (after a 44 year hiatus!) and it stills reads as I remember it from my first reading in 1951: a very personal evaluation by a highly skilled communicator with a broad grasp of his field. And he was clearly aware of the controversial nature of his commentaries, noting in his “Postlude,” near the end of his article:
I have been accused in this manuscript, both of being holier-than-thou, and of being satanic. With either accusation, I plead that to be both forceful and modest at the same time is a difficult task. If 1 appear to claim that I can see farther and from greater heights than some others, it is only - to use Newton's oft-quoted analogy - that those few cubits of stature have been attained by
Turner: Obituary for R.F. Daubenmire 61
climbing on the backs of giants. The giants are there for others to climb, even though the shoulders may bear us ungraciously.
In the fall of 1951 I was enrolled in Daubenmire’s course in autecology, for which his text was mandatory, as noted above. I had not given much thought as to how the text might have been written, but after reading Egler’s comments, I developed a greater interest in Dauby’s style.
Dauby was undoubtedly flattered that Egler possibly ranked him as among the “giants” of American ecology, but Egler was surely correct that the “shoulders [of such workers] may bear us ungraciously.” At least that seemed true of Dauby, who brought up Egler’s article time and again during the late fall of 1951, complaining that the editors of Ecology should ever have published such a commentary. But what most galled him was Egler’s paragraph on Dauby’s “style of writing,” which, in contrast with Clement’s style, was said to have
. succeeded to a high degree in developing a terseness, a paucity of words, a fact-crammed grammatical structure that is the goal of many a scientific writer. It is as functional, as devoid of decorative flourishes and artistic omamentation as the layercake skyscrapers built lately in New York. AS was said by the romanticist against the classicist, his writing had become correct and soulless, learned and uninspiring, scientific and godless, virtuous and cold. One can almost imagine that this author, beginning with terse abbreviated lecture notes, kept building through the years in card-catalogue style, inserting abstracts and summanies in their appropriate places as the new literature appeared. For these reasons, the book will long serve as a well- organized reference work for the American literature on the effects of environmental factors on plants.
And that was the way he lectured too, in both undergraduate and graduate courses, except in his autecology course, in which he never lectured, as noted in the above (the text seemingly written from abbreviated sentences on stacks of cards) with practically no sidebar diversions, even when controversy arose from among the students. And, too, that was the way he must have composed his text on Plant Geography (Academic Press, 1978). I attended his first class towards this new textbook venture in the spring of 1953, just before my doctoral defense scheduled for that same semester. My final personal insights into the man’s ouvre and psyche involves that class.
I truly looked forward to Dauby’s course. Having had a firm background in both plant geography and geology as a result of my master's work at Southern Methodist University in 1949-50, to say nothing of my courses in geomorphology and genetics at W.S.U., I felt primed and excited. Dauby even questioned my “need” to take his course, especially since I had made top grades in nearly all of my courses, and he was well aware of my conversational ability in systematics generally. “Concentrate on your doctoral thesis” he advised, knowing that I was scheduled to finish that same semester. But I told him my thesis was essentially written and that 1 would truly enjoy the class, efc. As a member of my doctoral committec, he relented.
_ Everything went fine in the course on Plant Geography. Dauby each day perfectly poised and academic, covering the topic from 5 x 8 cards with information not especially new or novel, throwing in this or that study called to the fore since Cain's
62 PHY TOLOGIA July 1995 volume 79(1):58-64
fine text on the subject, Foundations of Plant Geography, which first appeared in 1944, Nothing new really, until suddenly one day he digressed. Lecturing upon the origin of American deserts and their likely age, he bedazzled me (but perhaps not the class) with his observation that the deserts had developed very recently in North America, and that their floras were probably derived out of mostly recently extinct if not extant elements of the more temperate Artemisia shrublands and grasslands of the western Rocky Mountains, if not from conifer forests. The kingpin in this hypothesis, he reckoned, was the fossil Opuntia described by Chaney from the Green River shales of Utah, “the earliest and perhaps only fossil cactus from the New World” he noted. “We have to be objective and acknowledge the evidence,” he continued, drawing the words out tersely, and afterwards donning that smug Gioconda smile he was so adept at when playing his verbal trump cards.
I disagreed, of course, noting in class, lawyer-like perhaps, that all of the floristic evidence argued against his views: the Cactaceae is not well developed in temperate North America, anyway, if an Opuntia had happened to become fossilized in Eocene time, then it merely proved the cacti had been around for eons, and that the center of diversity of cacti in North America lay to the south in Anzona, New Mexico, Texas, mostly subtropical regions, much as suggested by Chaney in his paper, and what about Fouquieria, Idria (both belonging to the Fouquieriaceae, a family of only two genera confined to the hot deserts of North America without clear familial relationship elsewhere) and many other genera too numerous to mention, to say nothing of the genus Larrea which dominates the deserts of two continents, etc. On like that I held forth, and Dauby fumed, even entered this fray with a dead look of castigation. “I stand on the fossil data” he said, but noting at the same time that the state of Florida has as many cacti nearly as Anizona or New Mexico, and “certainly Florida is not a desert.” “But the Florida cacti mostly belong to the genus Opuntia,” | said, “many of these, if not most, of recent introduction or else the results of Small’s taxonomic splitting of this or that variable entity. Anyway,” I retorted, “The cacti of Florida, so far as evidence bearing on the age and ongin of the family Cactaceae, is meaningless.” And I forget, now, how our 15 minute debate went, but it ended with a stony silence on Dauby’s part, and “I wish you weren’t here” - look and an early closure of the lecture for that day.
After that venture into Dauby’s card session, upon the advice of my graduate student peers, I kept strictly quiet, dutifully recording his lectures in my own shorthand in preparation for our final exam, which was soon upon us.
The exam was well-structured, very fair, and straightforward, as were all of the exams in the four courses I took from him. But for me, on this particular exam, there was a problem. Dauby asked the question (assigning it 10 points): Give the age and origin of the family Cactaceae (not worded so as to be answered, according to Daubenmire!). Nevertheless, I placed in the appropriate space provided the answer according to Daubenmire, recounting his views very nicely I thought. But at the bottom of my answer I wrote “This is the answer which you might wish, Dr. Daubenmire, but for the correct answer, see the backside of this sheet.” There | defended my point of view (and those of many others) regarding this issue.
When the final exam was graded and the semester grades posted, | was surprised to see that I had received a 90 on my final exam (the entire cactus question graded as incorrect) and a B in the course. I inquired of him why he did not accept my answer to
Tumer: Obituary for R.F. Daubenmire 63
the cactus question concerned. His response was “Well, Turner, you got the answer, but you didn’t believe it, or else why did you give an additional answer on the back side of the sheet; in short, you only get to give one answer, not two, that’s why you missed the whole question!”
“OK,” I said, “But what about the B in the course. I had A’s in my earlier exams, and a low A (90) on the final, why a B? Other students with much lower averages received A’s [I’d made comparisons among my peers].” “Well,” he responded, “let’s put it this way, you got a B for Bad Behavior,” his eyes full on me dead as a desert duck, no water anywhere.
“Fine,” I responded, laughing, “now that I know the standards I won’t complain, considering the criteria I’m sure I got it fairly.” That was one of the few B’s I received in my university education and one that I am proudest of.
But the cactus question did not end there. Daubenmire attended my final defense (of a systematic thesis, a cytotaxonomic study of the genus Hymenopappus). After most of my committee members had finished asking this or that question, Dauby, who had said nothing to this point, suddenly said, “Turner, when and where do you think the Cactaceae arose?” I was taken aback, but rising to the occasion (I hoped then), I said strongly and affirmatively, without a glimmer of a smile, “Well, Dr. Daubenmire, do you want my answer, or yours?”
Dauby looked very distressed at my response, folded his papers, got up from the large table which was surrounded by about ten professors, and left the room. He did not approve my performance, but (so I was told) the upper administration, appraised his evaluation negatively and I passed my defense without undue rancor.
As a postscript to the cactus story recounted above, I can’t help but add that the fossil Opuntia described by Chaney from the fossil beds was, some 18 years later, found to be to a fossilized rhizome and associated root system of a monocot, possibly a sedge (Becker 1962). Upon reading this “inspiring” revelation I sent copy of the article to Dauby, with a little memo, merely stating, “Remember this?” He never responded. Nor did he include an account of his views on the origin of the Cactaceae in his text on Plant Geography. Indeed, published some 25 years after that first class on the subject, Dauby’s outlook re American deserts changed considerably, even introducing in his text some of the very same views which I propounded in his first course on the subject.
I hope the above account is not viewed by the reader as a “get-even” article. It is not intended as such (to my knowledge). Rather, I hope in this telling to capture an aspect of the man not generally known. Like most of us he had a mixture of traits some admirable, some not. But, surely some of these affected his research and teaching. In fact, | consider him with his often adamant views and determination to be the foremost ecologist in America (during his heyday) the essential ingredients of most successful scientists. Even at the time I admired his competitive personality, although disagreeing, upon occasion, with his behavior. Certainly he was one of the most
organized, clearly focused graduate level teachers to position information in my neural lodgings.
64 PHYTOLOGIA July 1995 volume 79( 1):58-64
Dauby was the academic father of numerous doctoral students in ecology, many of these friends of mine. For the most part he kept them at a distance; some he favored with warm, but detached, smiles and relatively brief office conferences; others he simply ignored, doubting their competence, begrudgingly entering into their research projects and practically never into their personal problems. Most of his students appeared to stand in awe of the man, even forming cabals among themselves and their leader, constituting a solid phalanx whenever Dauby's views were attacked by W.H. Whitaker or yet others. But that is another telling.
LITERATURE CITED
Becker, H. 1962. Reassignment of Opuntia to Cyperacites. Bull. Torrey Bot. Club 89:3 19-330.
Turner, B.L. 1972. Lowell David Flyr, 1937-1971. Sida 5:54-58.
___.. 1975. Marion Ownbey 1910-1971, an appreciation. Pl. Sci. Bull. 5:56-58.
umer--Department of Botany, University of Texas, Austin, Texas 78713
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Phytologia (August 1995) 79(2):65-67.
NEW NAMES AND COMBINATIONS, PRINCIPALLY IN THE ROCKY MOUNTAIN FLORA--IX
William A. Weber
University of Colorado Museum, Campus Box 350, Boulder, Colorado 80309 U.S.A.
The eighth paper in this series was published in Phytologia 70:23 1-233. 1991.
ABSTRACT
New combinations are proposed in Azaleastrum, Boechera, Oreobatus, and Picradenia. Validations are provided for previously published new combinations in Coriflora.
KEY WORDS: Azaleastrum, Boechera, Coriflora, Oreobatus, Picradenia, Rocky Mountains
Azaleastrum albiflorum Rydb. subsp. warrenii (A. Nelson) W.A. Weber, comb. nov. Based on Azaleastrum warrenii A. Nelson, Bot. Gaz. (Crawfordsville) 56:67. 1913.
Boechera pallidifolia (Rollins) W.A. Weber, comb. nov. Based on Arabis pallidifolia Rollins, Cruciferae of Continental North America, p. 181. 1993.
Oreobatus deliciosus (James ex Torrey) Rydb. subsp. neomexicanus (A. Gray) W.A. Weber, comb. nov. Based on Rubus neomexicanus A. Gray, Pl. Wrightianae 2:55. 1853. Synonyms: Oreobatus neomexicanus (A. Gray) Rydb., Rubus deliciosus James ex Torrey var. neomexicanus Kearney. This subspecies replaces the Coloradan race, Oreobatus deliciosus subsp. deliciosus in the southern edge of eastern Colorado and ranges south through New Mexico into southeastern Anzona. Subsp. neomexicanus has more distinctly lobed, larger, leaves which are soft-pubescent on both surfaces. Gray was under the misapprehension that R. deliciosus had purple flowers, and his major distinction appears to be that in neomexicanus the petals are white.
Picradenia richardsonii subsp. floribunda (A. Gray) W.A. Weber, comb.
nov. Based on Actinella richardsonii (Hook.) Nutt. var. floribunda A. Gray, Mem. Amer. Acad. Arts Sci., Ser. 2, 4:101. 1849 (Plantae Fendlerianae).
65
66 PHY TOLOGIA August 1995 volume 79(2):65-67
Validation of the generic name Coriflora (Ranunculaceae)
Coriflora W.A. Weber, Phytologia 51:372-374. 1982, was invalidly published as a result of the omission of certain information, rectified here.
Coriflora W.A. Weber, nom. nov. sViorna Spach, Hist. Nat. Végetaux: Phanérogames 7:268. 1839, type Clematis viorna L., Sp. Pl. 543. 1753 (Viorna urnigera Spach), nom. illeg., non Viorna (Pers.) Reichenbach, Handb. 277. 1837, nom. illeg., superfl. renaming of Muralta Adams. 1763, nom. rej.
Clematis cirrhosa L. is the type of (Pers.) Reichbach’s name; this type was explicitly excluded by Spach by citation (l.c. p. 261) as a synonym of Cheiropsis elegans Spach. According to Article 48, ICBN, Spach’s name is a validly published later homonym for which I am providing a replacement name. While irrelevant to this transaction, according to Pfeiffer, Nomenclator Botanicus 1588. 1874, Clematis, section Viorna antedates Gray, Syn. Fl. N. Amer. 1:5. 1895. Formal transfer of the species is effected below.
Coriflora addisonii (Bntt. ex Vail) W.A. Weber, comb. nov. BASIONYM: Clematis addisonii Britt. ex Vail, Mem. Torrey Bot. Club 2:28, footnote and pl. 3. 1890.
Coriflora albicoma (Wherry) W.A. Weber, comb. nov. BASIONYM: Clematis albicoma Wherry, J. Wash. Acad. Sci. 21:198, fig. 1. 1931.
Coriflora baldwinii (Torrey & A. Gray) W.A. Weber, comb. nov. BASIONY M: Clematis baldwinii Torrey & A. Gray, Fl. N. Am. 1:8. 1838.
Coriflora beadlei (Small) W.A. Weber, comb. nov. BASIONYM: Viorna beadlei Small, Man. Southeast. Fl. 527, 1504. 1933.
Coriflora bigelovii (Torrey) W.A. Weber, comb. nov. BASIONYM: Clematis bigelovii Torrey, Pacific Railroad Rep. 4:61. 1857.
Coriflora crispa (L.) W.A. Weber, comb. nov. BASIONYM: Clematis crispa L., Sp. Pl. 543. 1753.
Coriflora fremontii (James) W.A. Weber, comb. nov. BASIONYM: Clematis ochroleuca Ait. var. fremontii James, J. Cincinnati Soc. Nat. Hist. 6:120. 1883.
Coriflora gattingeri (Small) W.A. Weber, comb. nov. BASIONYM: Clematis gattingeri Small, Bull. Torrey Bot. Club 24:209. 1897.
Coriflora glaucophylla (Small) W.A. Weber, comb. nov. BASIONYM: Clematis glaucophylla Small, Bull. Torrey Bot. Club 24:337. 1897.
Coriflora hirsutissima (Pursh) W.A. Weber, comb. nov. BASIONYM: Clematis hirsutissima Pursh, Fl. Amer. Sept. 2:385. 1814.
Coriflora integrifolia (L.) W.A. Weber, comb. nov. BASIONYM: Clematis integrifolia L., Sp. Pl. 544. 1753.
Coriflora morefieldii (Kral) W.A. Weber, comb. nov. BASIONYM: Clematis morefieldii Kral, Ann. Missouri Bot. Gard. 74:665. 1987.
Coriflora ochroleuca (Ait.) W.A. Weber, comb. nov. BASIONYM: Clematis ochroleuca Ait., Hort. Kew. 2:260. 1789.
Coriflora palmeri (Rose) W.A. Weber, comb. nov. BASIONYM: Clematis palmeri.Rose, Contr. U.S. Natl. Herb. 1:118. 1891.
Weber: New combinations in Rocky Mountain flora IX 67
Coriflora pitcheri (Torrey & A Gray) W.A. Weber, comb. nov. BASIONYM: Clematis pitcheri Torrey & A. Gray, Fl. N. Am. 1:10. 1838.
Coriflora reticulata (Walt.) W.A. Weber, comb. nov. BASIONYM: Clematis reticulata Walt., Fl. Carol. 156. 1788.
Coriflora scottii (Porter) W.A. Weber, comb. nov. BASIONYM: Clematis scoittii Porter, Synops. Fl. Colorado, p. 1. 1874.
Coriflora texensis (Buckl.) W.A. Weber, comb. nov. BASIONYM: Clematis texensis Buckl., Proc. Acad. Nat. Sci. Philadelphia. 13:448. 1862.
Coriflora versicolor (Small ex Britt.) W.A. Weber, comb. nov. BASIONYM: Clematis versicolor Small ex Bnitt., Man..Fl. Northern States and Canada. 421. 1901.
Coriflora viorna (L.) W.A. Weber, comb. nov. BASIONYM: Clematis viorna L., Sp. Pl. 543. 1753.
Coriflora viticaulis (Steele) W.A. Weber, comb. nov. BASIONYM: Clematis viticaulis Steele, Contr. U. S. Natl. Herb. 13:364. 1911.
ACKNOWLEDGMENTS
I am indebted to Dan Nicolson for his advice on the Coriflora problem.
Phytologia (August 1995) 79(2):68-76.
NEWLY REQUIRED SUPRAGENERIC NAMES IN VASCULAR PLANTS
James L. Reveal
Department of Plant Biology, University of Maryland, College Park, Maryland 20742-5815 U.S.A.
ABSTRACT
Several supra-ordinal names in current use in textbooks and the more technical literature are not validly published. The following are now established: Cycadidae, Cycadophytina, Ephedridae, Ephedropsida, Equisetidae, Equisetophytina, Ginkgoidae, Ginkgoophyta, Ginkgoophytina, Gnetidae, Gnetophyta, Gnetophytina, Isoetidae, Lycopodiophytina, Magnoliophyta, Magnoliophytina, Ophioglossidae, Pinophyta, Pinophytina, Polypodiophytina, Psilotidae, Psilotophyta, Psilotophytina, Salviniidae, Taxidae, and Welwitschiidae. My own failures in 1992 require formal validation of the superorders Cornanae, Cyclanthanae, Loasanae, Nepenthanae, Primulanae, Rafflesianae, Sarracenianae, and Trochodendranae. Several ordinal names attributed to G.T. Bumett are invalid as they were proposed at the misplaced rank of section. The following names now in current use are validated: Acorales, Araliales, Aspleniales, Buxales, Calycerales, Connarales, Hippuridales, Nelumbonales, and Vitales. Cyphocarpaceae, a provisional family name proposed by Miers in 1848 is in current use; it is now validated. The revelation that Scrophulanaceae is polyphyletic requires the acceptance of Rhinanthaceae Juss. and recognition of Schlegeliaceae.
KEY WORDS: nomenclature, Magnoliophyta
When the three great workers on higher plant phylogeny and nomenclature, Cronquist, Takhtajan, and Zimmermann (1966), joined forces to promote a new system of classification for plants and the use of generic stems throughout all ranks above that of genus, they established a new era of botanical nomenclature for these oft used but rarely fully evaluated names. It was therefore a surprise to discover that several of their, and others (e.g., Tippo 1942; Bold 1957; Ehrendorfer 1971) now commonly used names were not validly published. In all instances noted here, the authors failed to provide a full and direct reference to a Latin description or diagnosis (Art. 36.1; Greuter et al. 1994). Many of the names proposed as new by the three were validated earlier by others, most notably Bessey (1907, 1910) and Boivin
68
Reveal: New names in Magnoliophyta 69
(1956); of course, several of the names proposed in 1966 are valid. Nonetheless, the following require validation:
Cycadidae Reveal, subclass nov., validated by a reference to the Latin diagnosis of a J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956) isonym of class Cycadopsida A.T. Brongniart (Enum. Pl. Mus. Paris xxxii, 136. 12 Aug 1843, as Cycadoideae, validated by a diagnosis in French).
Cycadophytina Cronquist, Takht., & Zimmerm. ex Reveal, subdiv. nov., validated by a reference to the Latin diagnosis of a later J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956) isonym of Class Cycadopsida A.T. Brongniart (Enum. Pl. Mus. Paris xxxii, 136. 12 Aug 1843, as Cycadoideae, validated by a diagnosis in French).
Ephedridae Cronquist, Takht., & Zimmerm. ex Reveal, subclass nov., validated by a reference to a H.G.L. Reichenbach (F/. Germ. Excurs. 1(2):156. Jan-Apr 1831, as Tribe Ephedreae) name with a diagnosis in Latin.
Ephedropsida Reveal, class nov., validated by a reference toa H.G.L. Reichenbach (Fl. Germ. Excurs. 1(2):156. Jan-Apr 1831, as Tribe Ephedreae) name with a diagnosis in Latin.
Equisetidae Reveal, subclass nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956, as Division Equisetophyta [“Equisophyta’’}) name with a diagnosis in Latin. :
Equisetophytina Reveal, subdiv. nov., validated by a reference toa J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956, as Division Equisetophyta {“Equisophyta”]) name with a diagnosis in Latin.
Ginkgoophyta Bold ex Reveal, div. nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956, as Class Ginkgoopsida) name with a diagnosis in Latin.
Ginkgoophytina Cronquist, Takht., & Zimmerm. ex Reveal, subdiv. nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956, as Class Ginkgoopsida) name with a diagnosis in Latin.
Gnetidae Cronquist, Takht., & Zimmerm. ex Reveal, subclass. nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:494. Dec 1956, as Class Gnetopsida) name with a diagnosis in Latin.
Gnetophyta Bold ex Reveal, div. nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:494. Dec 1956, as Class Gnetopsida) name with a diagnosis in Latin.
Gnetophytina Cronquist, Takht., & Zimmerm. ex Reveal, subdiv. nov., validated by a reference to the Latin diagnosis of a later J.R.B. Boivin (Bull. Soc. Bot. France 103:494. Dec 1956) isonym of Class Gnetopsida H.G.A. Engler (Nat. Pflanzenfam., I], 1:2. 26 Mar 1887, as Gnetales, validated by a diagnosis in German).
70 PHYTOLOGIA August 1995 volume 79(2):68-76
Isoetidae Reveal, subclass. nov., validated by a reference to a Latin diagnosis associated with a later J.R.B. Boivin (Bull. Soc. Bot. France 103:493. Dec 1956, as Isopsida) isonym of Class Isoetopsida H.G.A. Engler (in H.G.A. Engler & K.A.E. Prantl, Die Pflanzenfam. Nacht.:. 5. July 1897 with a diagnosis in German).
Lycopodiophytina O. Tippo ex Reveal, subdiv. nov., validated by a reference to a F.G. Bartling (Ord. Nat. Pl.: 14, 19. Sep 1830, as Class Lycopodiopsida [“Lycopineae”]) name with a description in Latin.
Magnoliophyta Cronquist, Takht., & Zimmerm. ex Reveal, div. nov., validated by a reference toa C.A. Agardh (Classes Pl. (2:] 13. 1825, as Class Polycarpellae) name with a description in Latin.
Magnoliophytina D. Frohne & U. Jensen ex Reveal, subdiv. nov., validated by a reference to a C.A. Agardh (Classes Pl. [2:] 13. 1825, as Class Polycarpellae) name with a description in Latin.
Ophioglossidae Takht. ex Reveal, subclass nov., validated by a reference to a rankless R. Brown (Prodr.: 136. 27 Mar 1810, as Ophioglosseae) name with a diagnosis in Latin.
Pinophyta Cronquist, Takht., & Zimmerm. ex Reveal, div. nov., validated by a reference to a F.G. Bartling (Ord. Nat. Pl.: 90,92. Sep 1830, as Class Coniferae) name with a description in Latin.
Pinophytina Cronquist, Takht., & Zimmerm. ex Reveal, subdiv. nov., validated by a reference to a F.C. Bartling (Ord. Nat. Pl.: 90, 92. Sep 1830, as Class Coniferae) name with a description in Latin.
Polypodiophytina Reveal, subdiv. nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:494. Dec 1956, as Subdiv. Pteridophytina [“Pterophytina”]) name with a diagnosis in Latin.
Psilotidae Reveal, subclass nov., validated by a reference to the Latin description of a later T. Nakai (Chosakuronbun Mokuroku (Ord. Fam. Trib. Nov.]: 206. 20 Jul 1943) isonym of Order Psilotales H.G.A. Engler (in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. Nachtr. 1:5. Jul 1897 with a diagnosis in German).
Psilotophyta B. Boivin ex Reveal, div. nov., validated by a reference to the Latin description of a later T. Nakai (Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 206. 20 Jul 1943) isonym of Order Psilotales H.G.A. Engler (in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. Nachtr. 1:5. Jul 1897 with a diagnosis in German).
Psilotophytina O. Tippo ex Reveal, subdiv. nov., validated by a reference to the Latin description of a later T. Nakai (Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 206. 20 Jul 1943) isonym of Order Psilotales H.G.A. Engler (in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. Nachtr. 1:5. Jul 1897 with a diagnosis in German).
Reveal: New names in Magnoliophyta 71
Salviniidae Pic. Serm. ex Cronquist, Takht., & Zimmerm. ex Reveal, subclass nov., validated by a reference to the Latin description associated with the type genus by M. Adanson (Gen. Pl. 2:15. Jul-Aug 1764).
Taxidae F. Ehrendorfer ex Reveal, subclass nov., validated by a reference to the Latin description of the type genus given by S.L. Endlicher (Syn. Conif:: 242. Mai-Jun 1847).
Welwitschiidae Cronquist, Takht., & Zimmerm. ex Reveal, subclass nov., validated by a reference to a J.R.B. Boivin (Bull. Soc. Bot. France 103:494. Dec 1956, as Class Welwitschiopsida [“Welwopsidia”]) name with a diagnosis in Latin.
My own failure (Reveal 1992) to provide a reference to a validating Latin description or diagnosis means that several superordinal names are not available.
Cornanae Thome ex Reveal, superord. nov., validated by a reference to a S.L. Endlicher (Gen. Pl. Suppl. 5:17. 1850, as Subfam. Cornoideae [“Corneae”’]) name with a diagnosis in Latin.
Cyclanthanae Thorne ex Reveal, superord. nov., validated by a reference to a F.G. Bartling (Ord. Nat. Pl.: 67. Sep 1830, as Tribe Cyclantheae [“Cyclanthea”]) name with a diagnosis in Latin.
Loasanae R. Dahlgren ex Reveal, superord. nov., validated by a reference to a P.F. Horaninow (Char. Ess. Fam.: 147. 1847, as Tribe Loaseae) name with a descniption in Latin.
Nepenthanae Takht. ex Reveal, superord. nov., validated by a reference to a J.H.F. Link (Handbuch 1:369. Jan-Aug 1829, as Subfam. Nepenthoideae (“Nepenthinae”]) name with a diagnosis in Latin.
Primulanae R. Dahlgren ex Reveal, superord. nov., validated by a reference to a A.J.G.C. Batsch (Tab. Regni Veg.: 206. 2 Mai 1802, as Order Cyathinae) name with a description in Latin.
Rafflesianae Thome ex Reveal, superord. nov., validated by a reference to a description in Latin for the Tribe Rafflesieae H.W. Schott & S.L. Endlicher ex E. Spach (Hist. Nat. Vég. 10:551. 20 Mar 1841, as “Rafflesiaceae”) given by R. Brown (Trans. Linn. Soc. London 19:242. 6 Nov 1844).
Sarracenianae Thome ex Reveal, superord. nov., validated by a reference to the Latin description of Sarraceniaceae given by G. Bentham & J.D. Hooker (Gen. PI. 1:48. 7 Aug 1862).
Trochodendranae Takht. ex Reveal, superord. nov., validated by a reference to an A.L. Takhtajan ex A.J. Cronquist (Integr. Syst. Class. Fl. Pl: 157. 10 Aug 1981, as Order Trochodendrales) name with a description in Latin.
72 PHYTOLOGIA August 1995 volume 79 2):68-76
In reviewing other suprageneric names of vascular plants I discovered that several names proposed by G.T. Bumett in 1835 and previously considered to have been validly published at the rank of order (Cronquist 1981; Reveal 1993), are invalid as they were proposed at the misplaced rank of section (Art. 33.5; Greuter et al. 1994). The following names, now in current use, are validated.
Acorales Reveal, ord. nov., validated by a reference to a J.H.F. Link (Handb. 1:144. Jan-Aug 1829, as Subfam. Acoroideae [“Acorinae”]) name with a description in Latin.
Araliales Hutch. ex Reveal, ord. nov., validated by a reference to an A.L. de Jussieu (Gen. Pl: 217. 4 Aug 1789, as Fam. Araliaceae [“Araliae”]) name with a description in Latin.
Aspleniales Pic. Serm. ex Reveal, ord. nov., validated by a reference to a C.B. Pres! (Abh. Konigl. Bohm. Ges. Wiss., ser. 4, 5:91. 2 Dec 1836, as Tribe Asplenieae [“Aspleniaceae”]) name with a description in Latin.
Buxales Takht. ex Reveal, ord. nov., validated by a reference to the Latin diagnosis given by F.G. Bartling (Ord. Nat. Pl.: 370. Sep 1830, as “Buxea”) for the Tnbe Buxeae Dumort. (Comment. Bot. xx. 1822).
Calycerales Takht. ex Reveal, ord. nov., validated by a reference to a R. Brown ex L.C.M. Richard (Mém. Mus. Hist. Nat. 6:74. Nov 1820, as Fam. Calyceraceae (“Calycereae”) name with a description in Latin.
Connarales Takht. ex Reveal, ord. nov., validated by a reference to an A.P. de Candolle (Prodr. 2:84. mid Nov 1825, as Tnbe Connareae) name with a description in Latin.
Hippuridales Pulle ex Reveal, ord. nov., validated by a reference to a J.H.F. Link (Enum. Hort. Berol. Alt. 1:5. 16 Mar-30 Jun 1821, as Fam. ana {“Hippurideae”]) name with a description in Latin.
Nelumbonales Nakai ex Reveal, ord. nov., validated by a reference to an A.P. de Candolle (Syst. Nat. 2:43. late Mai 1821, as Tribe Nelumboneae) name with a description in Latin.
Vitales Takht. ex Reveal, ord. nov., validated by a reference to an A.L. de Jussieu (Gen. Pl.: 267. 4 Aug 1789, as Fam. Vitaceae [“Vites”]) name with a descnption in Latin.
In preparing the list of family names for consideration under the rubric “NCU” (Hoogland & Reveal 1993), we failed to note that Cyphocarpaceae was a provisional name and thus not validly published (Art. 34.1[b]; Greuter et al. 1994). In order that this name may continue in use, as was our intent in 1993, it is validated here.
Cyphocarpaceae (Miers) Reveal & Hoogland, stat nov., based on Subfam. Cyphocarpoideae Miers, London J. Bot. 7:61. 1848, as Cyphocarpaceae.
Reveal: New names in Magnoliophyta 73
One goal of systematics is to recognize monophyletic taxa. The recent discovery (Olmstead & Reeves 1995) that Scrophulaniaceae, as defined by most modern workers, is polyphyletic requires a redefinition of that family. Two approaches can be taken, the reduction of numerous commonly accepted families to synonymy under a single, broadly defined Scrophulanaceae, or a fragmentation of the family into smaller groups reminiscent of the family treatments proposed by Jussieu (1789) and subsequent early nineteenth century authors. In reviewing the options, I have decided to take the latter course and propose the following linear sequence within a broadly defined Scrophulariales:
Scrophulaniales Lindl. (1833) Acanthales Lindl. (1833) Bignoniales Lindl. (1833) Gesneriales Dumort. (1829) Globulariales Dumort. (1829) Lentibulaniales Lindl. (1833) Pinguiculariales Dumort. (1829) Plantaginales Lindl. (1833) Rhinanthales Dumort. (1829) Veratrales Dumort. (1829) . Buddlejaceae K. Wilh. (1910) . Retziaceae Bartl. (1830) . Stilbaceae Kunth, nom. cons. (1831) . Bignoniaceae Juss., nom. cons. (1789) Crescentiaceae Dumort. (1829) Paulowniaceae Nakai (1949) . Schlegeliaceae Reveal (1996) . Verbasaceae Raf. (1821) . Scrophulariaceae Juss., nom. cons. (1789) Antirrhinaceae Pers. (1807) Capraniaceae Martinov (1820) Chelonaceae Martinov (1820) Gratiolaceae Martinov (1820) Limosellaceae J. Agardh (1858) Linanaceae Martinov (1820) Oxycladaceae (Miers) Schnizl. (1843-1870) 9. Rhinanthaceae Vent., nom. cons. prop. (1799) Aragoaceae D. Don (1835) Buchneraceae (Benth.) Lilja (1870) Digitalidaceae Martinov (1820) Erinaceae Duvau ex Pfeiff. (1873) Euphrasiaceae Martinov (1820) Melampyraceae Rich. ex Hook. & Lindl. (1821) Pedicularidaceae Juss. (1789) Sibthorpiaceae D. Don (1835) Veronicaceae Durande (1782) 10. Oftiaceae Takht. & Reveal (1993) Spielmanniaceae J. Agardh, nom. illeg. (1858) 11. Ellisiophyllaceae Honda (1930) 12. Orobanchaceae Vent., nom. cons. (1799)
ONAKH AWNe
74 PHY TOLOGIA August 1995 volume 79(2):68-76
Aeginetiaceae Livera (1927) Phelypaeaceae Horan. (1834)
13. Selaginaceae Choisy, nom. cons. (1823) Hebenstretiaceae Horan. (1834)
14. Globulariaceae DC., nom. cons. (1805)
15. Gesneriaceae Dumort., nom. cons. (1822) Belloniaceae Martinov (1820) Besleriaceae Raf. (1838) Cyrtandraceae Jack (1823) Didymocarpaceae D. Don (1822) Ramondaceae Godr. (1850)
16. Plantaginaceae Juss., nom. cons. (1789) Littorellaceae Gray (1821)
Psylliaceae Horan. (1834)
17. Pedaliaceae R. Br., nom. cons. (1810) Sesamaceae R. Br. ex Bercht. & J. Presl (1820)
18. Martyniaceae Stapf, nom. cons. (1895)
19. Trapellaceae Honda & Sakisaka (1930)
20. Myoporaceae R. Br., nom. cons. (1810) Bontiaceae Horan. (1834)
21. Acanthaceae Juss., nom. cons. (1789) Justiciaceae Raf. (1838) Mendonciaceae Bremek. (1954) Meyeniaceae Sreem. (1977) Nelsoniaceae (Nees) Sreem. (1977) Thomandersiaceae Sreem. (1977) Thunbergiaceae (Dumort.) Lilja (1870)
22. Lentibulariaceae Rich., nom. cons. (1808) Pinguiculaceae Dumort. (1829) Utriculariaceae Hoffmanns. & Link, nom. cons. (1809)
All names necessary for the proposed revision of Scrophulanales are available except for the following:
Schlegeliaceae (Gentry) Reveal, fam. & stat. nov., based on Tribe Schlegelieae Gentry, Fl. Neotrop. Monogr. 25:48. 19 Sep 1980.
The problematic relationship of Schlegelia, Gibsoniothamnus, and Synapsis with Bignoniaceae and Scrophulanaceae is well known (Monachino 1949; Williams 1970; Gentry 1980; Armstrong 1985). With the discovery of a fourth genus, Exarata (Gentry 1992), the distinctiveness of the taxon has become clear, and its nearness to Bignoniaceae confirmed. The fragmentation of Scrophulariaceae, the mandatory recognition of Paulowniaceae, and the unique position of the genera related to Schlegelia in the data presented by Olmstead & Reeves (1995) requires the recognition of Schlegeliaceae as a distinct family. Failure to fragment the traditional Scrophulaniaceae into smaller families would mean that recognition of Oftiaceae, Ellisiophyllaceae, Orobanchaceae, Selaginaceae, and Globulaniaceae is impossible, Bignoniaceae becomes doubtful, and the continued recognition of Pedaliaceae, Martyniaceae, Trapellaceae, Myoporaceae, and even Plantaginaceae dubious. Even the
Reveal: New names tn Magnoliophyta 75
continued acceptance of Gesneriaceae renders a broadly defined Scrophulariaceae paraphyletic.
The family name Rhinanthaceae (1799) will be proposed for conservation against the earlier Veronicaceae (1782) as the former was widely accepted in the early literature and its generic stem is the basis for Subfam. Rhinanthoideae Link.
ACKNOWLEDGMENTS
Work on the Indices Nominum Supragenericorum Plantarum Vascularum Project is supported in part by the Intemational Association for Plant Taxonomy and the University of Maryland at College Park in cooperation with the National Agricultural Library, U.S. Department of Agniculture, Beltsville, Maryland. Dr. Bryan E. Dutton and Dr. Kerry A. Barringer reviewed the manuscript. This is Scientific Article 9162, Contribution No. A-7834, of the Maryland Agricultural Experiment Station.
LITERATURE CITED
Armstrong, J.E. 1985. The delimitation of Bignoniaceae and Scrophulanaceae based on floral anatomy, and the placement of problem genera. Amer. J. Bot. 72:755- 766.
Bessey, C.E. 1907. A synopsis of plant phyla. Univ. Nebraska Stud. 7:275-373.
Bessey, C.E. 1910. The phyla, classes, and orders of plants. Trans. Amer. Microscop. Soc. 29:85-96.
Boivin, J.R.B. 1956. Les familles de Trachéophytes. Bull. Soc. Bot. France 103:490-50S.
Bold, H.C. 1957. Morphology of Plants. Harper and Row Publishers, New York, New York.
Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Columbia University Press, New York, New York.
Cronquist, A., A.L. Takhtajan, & W. Zimmermann. 1966. On the higher taxa of Embryobionta. Taxon 15:129-134.
Ehrendorfer, E. 1971. “Spermatophyta”, pp. 584-745. In: D. von Denffer, W. Schumacher, K. Magdefrau, & F. Ehrendorfer, Lehrbuch der Botanik, ed. 30, Stuttgart, Germany.
Gentry, A. 1980. Bignoniaceae. I. Crescentieae and Tourtettieae. FI. Neotrop. Monogr. 25.
Gentry, A. 1992. Exarata (Bignoniaceae), a new genus from the Choc6é Region of Ecuador and Colombia. Syst. Bot. 17:503-507.
Greuter, W., F.R. Barrie, H.M. Burdet, W.G. Chaloner, V. Demoulin, D.L. Hawksworth, P.M. Jgrgensen, D.H. Nicolson, P.C. Silva, P. Trehane, & J. McNeill, (eds.). 1994. International Code of Botanical Nomenclature (Tokyo Code) adopted by the Fifteenth International Botanical Congress, Yokohama, August-September 1993. Regnum Veg. 131.
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Hoogland, R.D. & J.L. Reveal. 1993. Vascular plant family names in current use. Regnum Veg. 126:15-60
Jussieu, A.L. de. 1789. Genera Plantarum. Paris, France.
Monachino, J.V. 1949. A note on Schlegelia and Dermatocalyx. Phytologia 3:102- 10S.
Olmstead, R.G. & P.A. Reeves. 1995. Evidence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and ndh sequences. Ann. Missouri Bot. Gard. 82:176-193.
Reveal, J.L. 1992. Validation of subclass and superordinal names in Magnoliophyta. Novon 2:235-237.
Reveal, J.L. 1993. A preliminary list of validly published automatically typified ordinal names of vascular plants. Taxon 42:825-844.
Takhtajan, A.L. 1967. Systema et Phylogenia Magnoliophytorum. Moscow, U.S.S.R.
Williams, L.O. 1970. An overlooked genus of the Scrophulariaceae. Fieldiana, Bot. 32:211-214.
Phytologia (August 1995) 79(2):77-79.
TWO NEW MEXICAN SPECIES OF SENECIO (ASTERACEAE)
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. |
ABSTRACT
Two new species of Senecio are described from México: S. ozolotepecanus B.L. Turner, from western Oaxaca, and S. viejoanus B.L. Turmer from southern Nuevo Leén and closely adjacent Tamaulipas. The former is closely related to S. -picridis Schauer of the Triangularis species- group and the latter is closely related to S. loratifolius Greenm. of the Lugentes species-group (sensu Barkley 1985).
KEY WORDS: Asteraceae, Senecio, México, systematics
Routine identification of Mexican comps has brought to fore the following novelties in Senecio.
SENECIO OZOLOTEPECANUS B.L. Turner, spec. nov. TYPE: MEXICO. Oaxaca: “Dirt road between La Cienegilla and San Gregorio Ozolotepec. Pine forest or cloud forest dominated by Clethra, Pinus and Quercus’ 2500-3000 m, 12 Dec 1989, Andrew McDonald 2970 (HOLOTYPE: TEX!; Isotype: MEXU).
Senecioni picridi Schauer similis sed differt foliis mumerosioribus majonbus angustioribus tenuiorbusque, bracteis involucn ut videtur multiseriatis, cal yculo longitudine bracteas intenores paene aequanti.
Suffruticose shrubs ca. 1 m high. Stems tomentose at first but soon glabrate. Leaves numerous and much overlapping, gradually reduced upwards and extending into the capitulescence. Midstem leaves sessile, briefly clasping, linear to linear- lanceolate, mostly 10-15 cm long, 0.5-1.0 cm wide, markedly white-tomentose beneath, less so or glabrate above, minutely denticulate to entire, the apices narrowly acute. Heads 30 or more arranged in open very leafy corymbose panicles, the ultimate peduncles tomentose, mostly 1-4 cm long. Involucres narrowly campanulate, 11-14 mm high, ca. 10 mm wide (pressed), the outermost bracts (calyculus) nearly as long as the inner, shaggy-white tomentose, the innermost pubescent at the apices with coarse hairs. Ray florets 8; ligules yellow, 8-10 mm long, ca. 3 mm wide. Disk
ae
78 PHYTOLOGIA August 1995 volume 79(2):77-79
florets ca. 40 (est.), 8-9 mm long, glabrous throughout; tube ca. 3.5 mm long; lobes triangular, ca. 1 mm long. Achenes (immature) columnar, ca. 2 mm long, pubescent throughout with appressed hairs; pappus of numerous white fragile slender bristles ca. 8 mm long.
This species belong to the ser. Fruticosa of Senecio (sensu Barkley 1985) and is seemingly most closely related to S. picridis Schauer, having most of the features of that species, except that the leaves are longer, thinner, more numerous, and markedly overlapping. Additionally, the involucral bracts (including the outermost) are nearly all of the same length and very loosely tomentose throughout, giving the involucre a
multiseriate appearance.
Senecio ozolotepecanus might also be mistaken for S. stoechadiformis, the latter readily distinguished by its thicker, fewer, entire leaves, naked capitulescence, and well-developed calyculus, the outermost bracts half as long as the inner or less.
SENECIO VIEJOANUS B.L. Turner, spec. nov. TYPE: MEXICO. Nuevo Leén: Mpio. Aramberni, Cerro Viejo, 3400 m, pine woods, 20 Nov 1993, Hinton et al. 23969 (HOLOTYPE: TEX!; Isotypes: GH,NY).
Senecioni loratifolio Greenm. similis sed capitulis majonbus (involucris plerumque 9-11 mm altis vs. 6-8 mm altis) dispositis plerumque in capitulescentia racemoidea et foliis anguste lineanbus (3-6 mm latis) non amplexicaulibus differt.
Simple-stemmed perennials 30-40 cm high from thick woody rhizomes, leaves linear, mostly basal, exauriculate, 0.3-0.6 cm wide, 10-20 cm long, tomentose above and below, with age the upper surface often glabrate. Heads 5-20, arranged in terminal raceme-like corymbs, the ultimate peduncles mostly tomentose, 1-3 cm long. Involucres broadly campanulate, 9-11 mm high, 10-12 mm wide (pressed); bracts ca. 23, linear-lanceolate, apically tufted; calyculus a series of loose bracts which grade into the inner series. Ray florets 13-32, the ligules yellow, 10-20 mm long, 2-4 mm wide. Disk florets numerous (80+), the corollas yellow, glabrous, 6-8 mm long, the tube 2.5-3.5 mm long with lobes ca. 0.8 mm long, somewhat warty on the outer surfaces. Achenes columnar, ca. 3 mm long, pubescent in lines; pappus of numerous white fragile capillary bristles 8-10 mm long.
ADDITIONAL COLLECTIONS EXAMINED: MEXICO. Nuevo Leon: Pefia Nevada, west side of Picacho Onofre, 3230 m, 4 Jul 1959, Beaman 2687 (TEX); Mpio. Zaragoza, Cerro Viejo, 3310 m, 5 Oct 1992, Hinton et al. 22394 (TEX); summit of Pefia Nevada, 2700-2900 m, “abundant in fir zone”, 5 Aug 1983, Nesom 4805 (TEX). Tamaulipas: Mpio. Miquihuana, 5 km N of Aserradero, ca. 2500 m, 25 Oct 1986, Herndndez S. 2078 (TEX); E side of Pefia Nevada, 3500-3600 m, 5 Jul 1985, McDonald 1614 (TEX); Cerro Pefia Nevada, 1 Jun 1975, Patterson 1523 (TEX); Pefia Nevada, 19 Jul 1949, Stanford et al. 2591 (TEX).
This species is obviously a sister-taxon of Senecio loratifolius, differing from the latter in having larger heads which are mostly arranged in raceme-like corymbs, and by the very linear-leaved foliage throughout, those along the stem not at all clasping. Senecio loratifolius, so far as known, is confined to the higher peaks of central Nuevo
Turner: New species of Senecio from México 79
Leon (Cerro Potosf and closely adjacent peaks in Coahuila) while S. viejoanus is restricted to the higher peaks of southern Nuevo Ledn (Cerro Pefia Nevada and Cerro Viejo).
ACKNOWLEDGMENTS
I am grateful to Guy Nesom for the Latin diagnoses, and to him and Mark Mayfield for reviewing the manuscript.
LITERATURE CITED
Barkley, T. 1985. Infrageneric groups in Senecio s.1., and Cacalia s.1. (Asteraceae: Senecioneae) in Mexico and Central America. Bnrittonia 37:21 1-218.
Phytologia (August 1995) 79(2):80-82.
A NEW SPECIES OF SALVIA (LAMIACEAE) FROM NUEVO LEON, MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Salvia jorgehintoniana Ramamoorthy, spec. nov. is described and illustrated from southern Nuevo Leén. It belongs to the sect. Curtiflorae of Salvia, where it relates to S. longistyla, a wide spread, variable species of western and south central México. It differs from the latter in having much larger corollas and smaller, abruptly acuminate calyx lobes.
FEY WORDS: Lamiaceae, Salvia, México, Nuevo Leén, systematics
Routine identification of Mexican plants has revealed the following novelty. To judge from notes and annotations accompanying type maternal, Dr. T.P. Ramamoorthy, in an earlier independent study, came to the same conclusion. Because of this I have credited him with the name and authorship, although the description and views as to its sectional relationship are those of my own.
SALVIA JORGEHINTONIANA Ramamoorthy, spec. nov. Figure 1. TYPE: MEXICO. Nuevo Leén: Mpio. Galeana, along road from Agua Blanca to San Miguel, 2020 m, “mixed forest of pine and oak”, 28 Aug 1991, Hinton et al. 23148 (HOLOTYPE: TEX!)
S. longistyla Benth. similis sed corollis 40-50 mm longis (vice corollae 25- 40 mm longae), lobis calycum 5-6 mm longis (vice lobi 6-12 mm longi), apicibus abrupte acutatis (vice apicum gradatim acuminatorum).
80
Tumer. New Salvia from México
Figure 1. Salvia jorgehintoniana (Hinton 22456).
82 PHYTOLOGIA August 1995 volume 79(2):80-82
Perennial herbs 0.8-1.0 m high. Midstems sparsely puberulous with mostly down-curved eglandular hairs. Leaves 10-25 cm long, 5-13 cm wide; petioles 4.5-9.0 cm long; blades broadly ovate to subdeltoid, pinnately nervate, sparsely to moderately pubescent above and below, especially along the veins, the margins serrate. Flowers in terminal racemes 20-30 cm long, arranged 4-6 to a node, the pedicels mostly 10-15 mm long, densely pubescent with spreading hairs 0.3-0.5 mm long, mostly eglandular but at least some with weakly developed terminal viscid glands. Calyces 2.1-2.5 cm long, sparsely to moderately pubescent with spreading, mostly glandular hairs to 1 mm long; lobes 5-6 mm long, deltoid, abruptly acute, the upper lobes 3-ribbed. Corollas red, 40-55 mm long; upper lobes 8-10 mm long; lower lobes 5-6 mm long. Stamens exserted for 5-10 mm beyond the apex of the upper lobes; anthers purple, ca. 2mm long. Style glabrous, extending somewhat beyond the stamens. Seeds ovoid, ca. 3 mm long, 1.5 mm wide, pale yellow, glabrous.
ADDITIONAL SPECIMEN EXAMINED: MEXICO. Nuevo Leoén: Mpio. Zaragoza, Cerro El Viejo, 1935 m, 6 Oct 1992, Hinton et al. 22456 (TEX).
According to label data, the type was collected from a “large colony”. The species is quite spectacular, with very large crimson corollas (up to 55 mm long, not counting the extended stamens and style branches). It belongs to the subgenus Calosphace, sect. Curtiflurae, where it relates to Salvia longistyla Benth., having the general habit, large leaves, and inflorescence of that species, but it differs markedly in having much larger corollas (40-55 mm long vs. 25-40 mm long) and shorter calyx lobes (5-6 mm long vs. 6-12 mm long) with abruptly acuminate apices (vs. gradually narrowing apices). In addition, the vestiture is less glandular-viscid and the styles are glabrous throughout, or nearly so.
Salvia jorgehintoniana is apparently endemic to southern Nuevo Leén, while S. longistyla is fairly widespread, occurring from Durango to Guerrero and across the trans-volcanic belt to Veracruz.
The apellation honors George Hinton, son of James Hinton, and grandson of the
late G.B. Hinton, who, in conjunction with his father, has collected many extraordinary plants from the state of Nuevo Leén.
ACKNOWLEDGMENTS
I am grateful to Gayle Turmer for the Latin diagnosis, and to her and Piero Delprete for reviewing the paper. Marcia Thompson provided the illustration.
Phytologia (August 1995) 79(2):83-88.
TAXONOMY OF THE HEDYOTIS ACEROSA (RUBIACEAE) COMPLEX
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Hedyotis acerosa, a species of the southcentral U.S.A. and northern México, is treated as having four morphogeographical vanities: var. acerosa, a widespread very common stiffly erect, fasciculate plant occurring mostly in Texas and Coahuila, México; var. polypremoides, an erect nonfasciculate plant of New Mexico, western trans-Pecos Texas and Chihuahua and westernmost Coahuila, México; var. potosina B.L. Turner, var. nov., a low, mat-forming taxon with elongate corollas, occurring from southemmost Coahuila to San Luis Potosi, México; and var. tamaulipana B.L. Turner, var. nov., an open, much-branched, wirey-stemmed plant with relatively small flowers occurring in westcentral Tamaulipas, México. A key to these taxa is provided along with maps showing their distr bution.
KEY WORDS: Rubiaceae, Hedyotis, Houstonia, Texas, México, systematics
Attempts to classify Mexican collections of Hedyotis acerosa assembled at LL, TEX has prompted the present study. Terrell (1991) provided a bnef overview of this complex, which he included in his concept of the genus Houstonia. While not pretending to understand fully the taxonomic limits of these two closely related genera, my taxonomic intuition, after comparing representative species of the groups concerned, is that they are best treated as but a single genus, Hedyotis having priority.
Distributional maps are based upon specimens on file at LL, TEX, all of these annotated accordingly.
KEY TO THE HEDYOTIS ACEROSA COMPLEX IN U.S.A.
1. Stems with leaves decidedly fasciculate; flowers mostly on pedicels 0.3-1.5 mm long; vestiture of stems various, but not uniformly minutely pubescent with down curved hairs; Culberson Co., Texas and eastwards. .................0000- var. acerosa
1. Stems with leaves not fasciculate, or very weakly so; flowers, at least some of them, on pedicels 2.0-20.0 mm long; vestiture of stems uniformly minutely pubescent with down-curved hairs ........5......0.0cccceeeeee sees var. polypremoides
84 PHYTOLOGIA August 1995 volume 79(2):83-88
KEY TO THE HEDYOTIS ACEROSA COMPLEX IN MEXICO
1. Stems with internodes much-shortened, forming low pulvinate mat-like plants mostly 2-5 cm high; corolla tubes mostly 8-10 mm long; southernmost Coahuila
and southwards to San Luis Potosf. ............... se cseeeeeeeeeeeeeeeeeenes var. potosina 1. Stems not as described in the above, mostly 5-15 cm high, forming well-defined rather naked stems; corolla tubes mostly 3-7 mm long....................eeee eee ee es (2)
2. Leaves markedly fasciculate; pedicels 0.3-1.5 mm long (rarely not so on lanky new growth of secondary shoots); vestiture various, but not minutely pubescent with down-curved hairs; common in Coahuila and closely adjacent central NUCVO: LeOns ictise Snes e eegs co eaee eset: egos eee eee var. fasciculata
2. Leaves weakly fasciculate, if at all; pedicels mostly 2-20 mm long; vestiture uniformly minutely pubescent with mostly down-curved hairs (rarely subglabrous in var. tamaulipana); Chihuahua, Coahuila and Tamaulipgs. ....(3)
3. Corolla tubes mostly 3-4 mm long; calyx lobes 1.5-2.0 mm long; Tamaulipas. .....
Sra bscnndiee at oki uak we reaiiean a rigednest igesedeea cers Rue miaer anew var. lamaulipana
3. Corolla tubes mostly (4-)5-6(-7) mm long; calyx lobes 3-4 mm long; Chihuahua, COMMU aisitistiuacincatane masat ergs eerste cons eae rena var. polypremoides
HEDYOTIS ACEROSA A. Gray, Pl. Wright. 1:81. 1850. Houstonia acerosa (A. Gray) Benth. & Hook. f. (for additional synonymy cf Terrell 1991).
HEDYOTIS ACEROSA A. Gray var. ACEROSA
As described by Gray, this is a ngidly erect, fasciculate, sparingly branched plant ca. 15 cm high. Type matenal was collected by C. Wnght in late June of 1849, presumably in present day Kinney or Val Verde County, Texas where Wright would have first encountered the taxon. Wright, in his protologue, also cited a specimen from near Buena Vista, Coahuila, collected by Gregg, among others alluded to; clearly lectotypification is needed, but from the description there is little doubt as to the application of the name.
This variety, in habit, is relatively uniform throughout its range, but its vestiture varies considerably as shown in Figure 2. In the latter illustration, specimens with a mixture of both long and very short, mostly straight hairs are depicted as open circles; those with + uniformly small straight hairs, and/or + glabrous are shown as closed circles; specimens intermediate to these extremes shown as half circles. At least a few of the specimens here accepted as var. acerosa were annotated by Terrell as subsp. polypremoides (e.g., Ector Co., Tex; Rowell 5605 [LL}).
HEDYOTIS ACEROSA A. Gray var. POLYPREMOIDES (A. Gray) W.H. Lewis, Ann. Missouri Bot. Gard. 55:397. 1969. Hedyotis acerosa A. Gray var. bigelovii (Greenm.) W.H. Lewis Hedyotis polypremoides (A. Gray) Shinners Houstonia acerosa A. Gray subsp. polypremoides (A. Gray) Terrell Houstonia polypremoides A. Gray Houstonia polypremoides A. Gray var. bigelovii Greenm.
85
Taxonomy of Hedyotis acerosa complex
Tumer:
>
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86 PHYTOLOGIA August 1995 volume 79(2):83-88
ihe
| |
Figure 2. Distribution of Hedyotis acerosa in México: var. acerosa (open circles); var. polypremoides (closed triangles); plants + intermediate to var. acerosa but tending to var. polypremoides (half solid triangle); var. potosina (closed circle); var. tamaulipana (Open square).
Tumer: Taxonomy of Hedyolis acerosa complex 87
Lewis (1968) lectotypified this taxon by Pringle 356 (GH), collected in the Santa
Eulalia Mts., Chihuahua (ca. 28° 35’ N, 105° 53’ W). Shinners (1949) accepted this taxon as a good species, but Lewis (1968, 1969) recognized it as but varietally distinct, although Terrell (1991) recognized it as a subspecies. Lewis (1969) took up the name Hedyotis acerosa var. bigelovii for this taxon, but the current code mandates the varietal name polypremoides as correct, much as inadvertently supplied by Lewis.
I accept its varietal status because the taxon clearly grades into the var. acerosa in the trans-Pecos region of Texas (e.g., Whitehouse s.n. [TEX]; Young s.n. [TEX]- both from the Guadalupe Mts.) and in México (e.g., western Coahuila, Johnston 389 {LL}; etc.), mostly along the western periphery of var. acerosa, as noted by Terrell (1979).
Occasional plants of var. acerosa, either aberrant late-flowering, or cut-back shoots showing new growth, are apt to be mistaken for var. polypremoides but individuals of the former can be readily sorted out by their pubescence, as noted in my key.
HEDYOTIS ACEROSA A. Gray var. POTOSINA B.L. Turner, var. nov. TYPE: MEXICO. San Luis Potosi: Charcas, Jul-Aug 1934, C.L. Lundell 5048 (HOLOTYPE: LL!).
H. acerosa A. Gray var. acerosa similis sed plantae 2-4 cm altae (vice 5-15 cm), breves, ramosissimae e basi sunt, tubis corollarum plerumque 8-10 mm longis (vice 4-6 mm). .
ADDITIONAL SPECIMENS EXAMINED: MEXICO. Coahuila: mouth of San Lorenzo Canyon, SE of Saltillo, 6200 ft, 2 Aug 1975, Engard 690 (LL); 4 mi S of Saltillo, 6000 ft, 18 Nov 1958, Rollins 58125 (LL). Nuevo Leén: 18 mi E of Matehuala, road to Dr. Arroyo, 5 Aug 1970, Flyr 1536 (TEX). San Luis Potosi: 16 mi N of Matehuala, 11 Feb 1960, Johnston 5088A (TEX); 70 mi S of Matehuala, 2 Sep 1975, Simpson 7036 (TEX). Tamaulipas: Mpio. Bustamante, 38.8 km N of Tula, 2 Jun 1983, Barnett 83071 (TEX).
All of the above cited plants, including the type, were annotated by Terrell as subsp. acerosa. But, as indicated by label data on Simpson 7036, var. potosina is a mat-forming plant, having a very different growth habit than found in var. acerosa; additionally, the corolla tubes are nearly twice the length of those of the latter, and it occupies a decidedly different geographical region. While treated at the varietal level, it might ultimately prove to be specifically distinct, at least no clear intermediates were found linking var. potosina to var. acerosa, although the two taxa come in close proximity in the region about Saltillo, Coahuila.
HEDYOTIS ACEROSA A. Gray var. TAMAULIPANA B.L. Turner, var. nov. TYPE: MEXICO. Tamaulipas: Mpio. Villagran, 1 mi E of Ejido de San Lazaro (ca. 24° 35'N x 99° 13’ W), ca. 1500 ft, 11 Oct 1959, M.C. Johnsion (with J. Graham) 4281k (HOLOTYPE: TEX!).
88 PHYTOLOGIA August 1995 volume 79(2):83-88
H. acerosa A. Gray var. polypremoides (A. Gray) W.H. Lewis similis, sed plantae laxe divaricateque ramosae sunt, lobis calycum brevioribus (plerumque 1-2 mm longis vice 3-4 mm), et tubis corollarum brevioribus (plerumque 2-4 mm longis vice 4-6 mm).
ADDITIONAL PLANTS EXAMINED: MEXICO. Tamaulipas: Mpio. San Carios, 6 mi S of San Carlos on the road to Padilla, 1600 ft, calcareous terraces of Arroyo de San Carlos, 13 Dec 1959, Johnston 5007A (TEX); Mpio. Casas, “S mi E of Casas on the new Victono-Soto la Marina highway”, 28 Sep 1960, Johnston 5784B (TEX).
This taxon resembles Hedyotis acerosa var. polypremoides but the plants are loosely divaricately branched, the calyx lobes shorter (mostly 1-2 mm long vs. 3-4 mm long), and have shorter corolla tubes (mostly 2-4 mm long vs. 4-6 mm long).
Terrell annotated all of the maternal cited above, including the type, as Houstonia
acerosa subsp. polypremoides. Considering the differences between the latter and var.
tamaulipana, as outlined in the above, and the aaa nature of the populations concemed, varietal status for the latter seems justified
ACKNOWLEDGMENTS
I am grateful to Gayle Turner for the Latin diagnosis, and to her and Piero Delprete for reviewing the paper.
LITERATURE CITED
Lewis, W.H. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Gard. 55:3 1-33.
Lewis, W.H. 1969. Hedyotis acerosa var. bigelovii, comb. nov. (Rubiaceae). Ann. Missouri Bot. Gard. 55:397.
Shinners, L.H. 1949. Transfer of Texas species of Houstonia to Hedyolis (Rubiaceae). Field & Laboratory 17:166-169.
Terrell, E.E. 1991. Overview and annotated list of North American species of Hedyotis, Houstonia, Oldenlandia, and related genera. Phytologia 71:212-243.
Phytologia (August 1995) 79(2):89-92.
TAXONOMIC STUDY OF HEDYOTIS PALMERI (RUBIACEAE)
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Hedyotis palmeri (A. Gray) W.H. Lewis (=Houstonia palmeri A. Gray) is a species of northcentral México occurring in mostly shallow soils from 1000- 2200 m. Two morphogeographical infraspecific categones are recognized: var. palmeri (including H. longipes S. Wats.), occurring in southeastern Coahuila and most of Nuevo Leén from 1600-2200 m; and var. muzquizana B.L. Turner, var. nov., occurring in northcentral Coahuila mostly at 1000- 1600 m. The latter differs from the former in being a taller plant with larger corollas and longer pedicels. A map showing the distributions of the two taxa is included. Lectotypification for H. longipes and H. palmeri is provided.
KEY WORDS: Rubiaceae, Hedyotis, Houstonia, México, systematics
Terrell (1991) provided an overview of the genera Hedyotis, Houstonia, and Oldenlandia for North America. In this, Hedyotis palmeri (A. Gray) W.H. Lewis was positioned in the genus Houstonia, subgenus Fricotis Terrell. This subgenus includes Hedyotis acerosa A. Gray which I have recently studied (Turner, 1996), concluding that it is seemingly adequately treated as an element of Hedyotis. In working over that complex I became interested in the regional variation displayed by H. palmeri, hence the present paper.
Terrell did not recognize infraspecific categories under Hedyotis palmeri, but a sorting of the specimens available to me, most of these annotated by him, showed that there was a series of populations in northcentral Coahuila made up of larger wirier plants, with larger corollas on more elongate pedicels than occurs in_ typical populational elements of H. palmeri. The two series of populations apparently do not coexist, and while clear intermediates between these are not known, their close relationship is so obvious that I have described the new taxon as but varietally distinct.
_A key to these two varieties follow, along with a map showing their distribution (Figure 1), based upon material on file at GH, LL, SRSC, TEX.
89
90 PHYTOLOGIA August 1995 volume 79(2):89-92
is
we o
Figure 1. Distribution of Hedyotis palmeri: var. muzquizana (open circles); var. palmeri (closed circles).
Tumer: Study of Hedyotis palmeri 91
Key to varieties of Hedyolis palmeri
Mature corolla tubes mostly (6-)8-10 mm long; pedicels mostly 20-30 mm long; n
SCAU Ao 2a as or sey pacaia sea Sen Sati ocus ina tasesaas dp eseiweeeasaes var. muzquizana Mature corolla tubes mostly 4-5(-8) mm long; pedicels mostly 5-20 mm long; s OAHU ae NEV OA SCOI nds sis nn 0 ote ase onde ies asus onesie daw nas eeu vetoes var. palmeri
HEDYOTIS PALMERI (A. Gray) W.H. Lewis var. PALMERI, Rhodora 63:222. 1961. BASIONYM: JHoustonia palmeri A. Gray, Proc. Amer. Acad. Arts 17:202. 1882. TYPE: MEXICO. Coahuila: “Lerios, 45 mi E of Saltillo”, Jul 1880, Edward Palmer 397 (LECTOTYPE [here selected]: GH!). The lectotype is mounted on the same sheet with two other collections, all of these cited in the protologue: Palmer 398, Jul 1880, from “40 mi S of Saltillo”, and Palmer 2117, Jul 1880, from “6 mi E of Saltillo”. The several specimens are very similar but the collection selected as the lectotype is better developed as to flowering and fruiting material.
Houstonia longipes S. Wats., Proc. Amer. Acad. Arts 18:97. 1883. Hedyotis longipes (S. Wats.) W.H. Lewis, Rhodora 63:222. 1961. TYPE: MEXICO. Nuevo Le6n: Monterrey, Feb 1880, Edward Palmer 395 (LECTOTYPE [here selected]: GH!). The lectotype 1s mounted on the same sheet with two other collections: E. Palmer 394, Sep 1880, from Monclova, Coahuila, and Gregg s.n., 29 May 1847, from “Cerralbo”, Coahuila (?). All of these are very similar and all were cited in the protologue. The Gregg specimens are the only ones having both flowers and fruits, the corolla tubes being ca. 4 mm long. Corollas are absent on the remaining collections.
This is the commonly collected widespread vanety of the species and is known to me only by collections obtained from the south and east of var. muzquizana (Figure
1).
HEDYOTIS PALMERI (A. Gray) W.H. Lewis var. MUZQUIZANA B.L. Tumer, var. nov. TYPE: MEXICO. Coahuila: Mpio. M. Muzquiz, ca. 130 road km NW of Muzquiz on Hwy 2A, “midslope of Sa. La Encantada along road up to tunnel entrance to La Encantada basin and mining area. Steep NW - facing slope, .
. . common but scattered, mostly in moister microsites”, 28° 30’ 40” N x 102°
19’ 30” W, 3 Jun 1992, Guy Nesom 7380 (with M. Mayfield) (HOLOTYPE: TEX!; Isotype: MEXU).
Similis H. palmeri (A. Gray) W.H. Lewis var. palmeri sed differt tubis corollarum plerumque 8-10 mm longis (vice tuborum plerumque 4-6 mm longorum) et pedicellis plerumque 20-30 mm longis (vice 5-20 mm longis).
ADDITIONAL SPECIMENS EXAMINED: . MEXICO. Coahuila: Mpio. M. Muzquiz, Cuesta del Plomo, 1000 m, 7 Jun 1972, M.C. Johnston et al. 7550p (TEX); Muzquiz, spring 1935, Marsh 330 (GH,SRSC,TEX); Santa Rosa Mts., 8 Jul 1938, Marsh 1251 (GH,TEX); 15 air km NW of La Babia, 18 May 1992, Mayfield 1426
92 PHYTOLOGIA August 1995 volume 79(2):89-92
(TEX); SW margin of Serranias del Burro, 1400-2100 m, 23 Jun 1991, Ruiz 47 (TEX); Rancho Agua Dulce, 1 Jul 1936, Wynd & Mueller 400 (GH).
Nearly all of the specimens cited above have the characters alluded to in the diagnosis, and it is clear that these represent populational units distinct from var. palmeri. Additionally, the plants concerned, in general, appear to be taller, wirier, with a less branched inflorescence than occurs in var. palmeri.
Terrell annotated several or more of the above sheets as Hedyotis palmeri without comment. In spite of numerous collections of var. palmeri on file at LL, TEX (40 sheets), I have not detected any clear intermediates between these allopatric entities except for a single collection from “Alamar”, Pablillo, SE of Galeana, Nuevo Le6én (Pennell 17191 {GH]), having corolla tubes 6-8 mm long, otherwise it is similar to var. palmeri. Late-flowering specimens of var. muzquizana, however, occasionally produce small flowers (e.g., Wynd & Muller 400).
It is possible that future field workers will elevate var. muzquizana to specific rank, typical specimens differing markedly from var. palmeri.
ACKNOWLEDGMENTS
I am grateful to GH and SRSC for the loan of materials. Gayle Tumer provided the Latin diagnosis, and she and Piero Delprete reviewed the article.
LITERATURE CITED
Terrell, E.E. 1991. Overview and annotated list of North American species of Hedyotis, Houstonia, Oldenlandia, and related genera. Phytologia 71:212-243. Tumer, B.L. 1996. Taxonomy of the Hedyotis acerosa (Rubiaceae) complex.
Phytologia 79:83-88.
Tumer: New Hedyolis from Texas 95
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. less we ° ; xR ons a= ° Ise ° € <0 "oO Px eng : W ac lo 2s a ees : zy jo Bos Es <3 i. : 3 3) ff ae ad a's . Pro) l ej— BD y : s > wv | ge | = : re Ww w T4 Ss ° zl ce ze : ola pe: = gm ox "Ee ‘ Ly “a eon | - ~ = + Sos. 1 u 3 ! Fw ro) oa Sen re eae 26 we if, Oe ° <S iz z “8 x: c x3Zz z ; Paaeionn 2 pe Sees Sms 5S 26 ood 2 sO haa) > 2 © ammo 5 fiw Me = v, S = e why (eo) Zz me a ae a%es- m. Gat ad 2 < b z z4 Vv Po @ 5 fEY EY 5 . 3 . > Ww F 62085 g a « > z L2é& j8& Acs | z e e e [a) °o bf if Q lz z og _ S : , | eee as N55 y Panna w * oO «<° w tet oe} 49. Ce HR a Se = ® © z2 4 ; watts j= Ze 8 ° of Pent ee Ww sl > bs Ps S 2 * “aS ‘inye aS N \ ie yore 28S lo e 3 ME a OA OU E i= a 56 TREE PS es BO ow we ee < e.- W at ae FS i. “ aia? © —zeom re o< ze D i Ete te WS re z < e 3 ice, Sap gem 4: ‘es a7 5 om? S3A3 ow F « ORE ie ws A Pkt A ie oN OE ia Frwwern., o = © om ae" 4 Ss a Me a No = e eo ue ans tenes < oe ows, sa: TA” 2 pe ® 1 ‘ =: Ee E ie GEG. SIGS Big AES OO = fo) eh tin leg 2 Fr e\ 2S Y f? w OS. ee EP jae “Zammss oeaeN, ehg eh? 2 MSs Whee = pace AC Oh 2"*sQ,, NM s mm 3 ag fr es i tr ae Ff > yw ny See QRS ON Pe 7/6 oe? oss ww ea et w iy - =e Zarate ms ay iS oe tt) pasate swat Fes CT Se PPFD anh EIR lea WOE ase ears ot Ou : te PY aw ee r\ S * Jo fe Of we w Ke Er Fe Ae Le SIERC pio SF yt 1g en 9 Bid gti teeing eee me a} “0 yo" svn a Rate) oe saint Je v aN aay lay p> Ly wt? ARN sop AY a , ee sR SIERRAE SS O% Qos os Ae a ER DIABLO, Se a WIE 5 s %, oe 4 = a Rt) * a ° Rr SE See me = | +> asc bi ae <2S OS c U(r r) gee Se (os os loo"3e ° < re eo 5 es i ‘ & Pins 22 2 a or pe € eee ae Raa x 3 <= 3< < 2“: yr. 8 | ad “we = Seas eA - bs ad an Fae aS ey @o a et SS o € ] Ww = z i z= rY) a 7) | 55 : ia ao - > =) te Es : : ae : : ° poe e 7" , 19 : ne 5 ] qs° & a’ z | z< 3 ast ——— KB xo pWze Wes ol wy,
Figure 2. Map showing locations of the localized endemics, Hedyotis butterwickiae (circle) and H. pooleana (triangle).
96 PHY TOLOGIA August 1995 volume 79(2):93-96
Mat-forming perennial herbs 2-3 cm high. Stems glabrous and much-branched from the base, the internodes mostly 1-3 mm long. Stipules lanceolate, 1-2 mm long. Midstem leaves thick, lanceolate, 5-7 mm long, 1.0-1.4 mm wide, strongly 1-nerved, glabrous except for minutely hispidulous margins, the apices decidedly apiculate. Flowering branches not much extending beyond the leaves, if at all. Pedicels 0.1-0.4 mm long. Calyces ca. 2 mm long, the 4 lobes lanceolate, ca. 1 mm long. Corollas 2.5-3.0 mm long, white, the 4 lobes lanceolate, ca. 2 mm long, hispidulous externally at the apices, moderately pilose within. Anthers included within the tube. Styles excurrent for ca. 2 mm. Capsules orbicular ca. 1 mm high, dehiscing loculicidally across the disk. Seeds not available.
Hedyotis pooleana much resembles H. mullerae Fosberg of northcentral México but the leaves are markedly different, as noted in the above diagnosis. When first collected Ms. Jackie Poole (conservation biologist and one-time curator at LL, TEX) thought the plant might be H. mullerae, which it superficially resembles. Comparisons of her material with ten or more collections of the latter at LL, TEX has shown the distinctiveness of the taxon proposed here. Terrell (1991: Phytologia 71:212-243.) provided an overview of the North American species of Hedyotis and related genera, but did not have material of the present species.
Jackie Poole collected the species again at the type locality on 25 May 1985 (Poole 2527 ([SRSC,TEX]). She also informed me that she observed the species on 27 Nov 1987 along the same ridge at about 4840 ft elevation within the boundary of the Big Bend National Park, about a mile or so from the type locality. The Dead Horse Mountains is an extension of the loftier Sierra del Carmen range across the Rio Grande in Coahuila, México. Wells (1965: Southwestern Naturalist 10:256-260.) has provided a vegetational account of this Texas extension.
Northcentral México and closely adjacent trans-Pecos Texas harbor a large array of endemic taxa. This is especially so for Hedyotis for the present novelty is apparently restncted to the Dead Horse Mountains, not too far removed from the recently described Hedyotis butterwickiae Terrell, the two occurring in close proximity (Figure 2). I can’t help but add that both of the individuals for which these two taxa were named, obtained advanced degrees under my direction years ago now, and both are still ardent field workers. Bless such students! |
ACKNOWLEDGMENTS
I am grateful to Jackie Poole for calling the novelty to my attention, to Gayle Tumer for the Latin diagnosis, and to her and Piero Delprete for reviewing the manuscript.
Phytologia (August 1995) 79(2):97-101.
A NEW SPECIES OF SALVIA (LAMIACEAE) FROM NORTHERN MEXICO
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Salvia jaimehintoniana Ramamoorthy spec. nov. is described and illustrated. It occurs in Durango, Hidalgo, Nuevo Le6n, and Tamaulipas, and belongs to the sect. Farinaceae sensu Epling. It has previously been described as S. azurea var. mexicana Epling. When elevated to specific status it must take on another epithet, the name S. mexicana L. having priority at the species level. Because of this, the new name, S. jaimehintoniana, with new typification is proposed. The distinctions between S. azurea and S. jaimehintoniana are discussed, and a map showing their distributions in Texas and México 1s provided.
KEY WORDS: Lamiaceae, Salvia, México, Salvia azurea, systematics
Routine identification of Mexican salvias has prompted the present study.
SALVIA JAIMEHINTONIANA Ramamoorthy, spec. nov. Figure 1. TYPE: MEXICO. Nuevo Leén: Mpio. Zaragosa, 4.3 road mi. S of Zaragosa on the road to Aserradero la Encantada, 20 May 1988, Burford L. Westlund 24 (HOLOTYPE: TEX!}).
_ Similis S. azurea Lam. sed differt lobis superis (conjunctis) calycum acutis (vice loborum obtusorum), bracteis floralibus late ovatis et persistentibus (vice bractearum lanceolatarum et mox deciduarum).
Perennial herbs 30-50 cm high, forming fascicles of tuberous roots. Stems stiffly erect, relatively unbranched or remotely branched, minutely hispidulous to subglabrate, the nodes pilose with hairs 0.6-1.0 mm long. Midstem leaves elliptical, lance-elliptical to lance-obovate, pinnately nervate; petioles 3-15 mm long; blades mostly 4-10 cm long, 1.2-3.5 cm wide, subglabrous or pubescent along the major veins, undersurfaces markedly glandular-punctate, the margins crenulodentate to nearly entire. Flowers 4-8 to a node, arranged in terminal interrupted spikes. Bracts ovate, persistent, 6-10 mm long, 3-5 mm wide, appressed-pubescent dorsally, the margins ciliate. Calyces 6-8 mm long, flaring upwards, 3-5 mm wide at orifice (pressed); upper lip ca. 2 mm long with 7 well-defined hispidulous ribs. Corollas
oy
| 98 PHYTOLOGIA August 1995 volume 79(2):97-101 |
blue, 15-18 mm long; tubes 2-3 mm long; throats abruptly bulging below, 4-6 mm | long (tube and throat 6-10 mm long); lower lip, 3-lobed, 6-9 mm long; upper lip densely puberulous, 3-4 mm long; tubes and throat not papillose within, or but weakly | so. Anthers included within the upper lip, attached near the orifice of the throat. | Styles pubescent above, the upper style branches ca. 3 mm long, the lower branches | ca. 1 mm long. Fruits ovoid, smooth, ca. 2 mm long, 1.5 mm wide.
REPRESENTATIVE SPECIMENS EXAMINED: MEXICO. Durango: Mpio. | de Tepehuanes, El Tarahumar, 2720 m, 27 Aug 1983, Tenorio 4200 (TEX). Hidalgo: | 6.5 air km ENE of Jacala, 1700 m, 13 Jul 1991, Mayfield et al. 820 (TEX). Nuevo | Ledén: Mpio. Galeana, above El Carnzo, 1900 m, 16 Oct 1983, Hinton et al. 18615 | (TEX); along road from Agua Blanca to San Miguel, pine-oak forests, 2030 m, 28 Aug 1991, Hinton et al. 21276 (Hinton herbarium); above Agua Blanca, oakwoods, 2305 m, 4 Jul 1992, Hinton et al. 22285 (Hinton herbarium); ca. 30 mi S of | Monterrey, 13 Aug 1934, C.H. & M.T. Mueller 1335 (TEX); area of Cerro Pefia | Nevada, ca. 12 km NE of San Antonio Pefia Nevada, N and NW slopes of mt. known | locally as Picacho Onofre, Jul 1977, Wells & Nesom 345, 374, 440 (TEX). | Tamaulipas: ca. 6 km NW of Rancho El Cielo, ca. 12 km NW of Gémez Farias, 1900 | m, 12 Aug 1991, Iltis 30724 (TEX). |
This species 1s represented at LL, TEX by twenty or more collections and is presumably the same as Salvia azurea subsp. mexicana Epling, the latter typified by collections from near Galeana, Nuevo Le6én (Photoisotypes: TEX!; Paratype: Mueller 1335 [TEX!]). I have given the plants concerned a new specific name with new typification since the name S. mexicana L. is preoccupied, precluding the elevation of Epling’s subspecific epithet.
Salvia jaimehintoniana belongs to the sect. Farinaceae as circumscribed by Epling (1939, 1940), having the perennial habit, interrupted inflorescence with persistent bracts, upper lip of the calyx with 5-7 ribs, and corolla features of species belonging to that difficult complex.
The specimen cited from Hidalgo differs from the other collections in having spreading pilose hairs, the vestiture along the stems mostly 0.5-1.0 mm high; in all other characters, however, it is like the type matenal. The rather isolated specimen from Durango has all of the features of typical forms of Salvia jaimehintoniana except for its somewhat larger corollas and smaller floral bracts.
Salvia jaimehintoniana differs from S. azurea in having calyces with the upper fused lobes acute (vs. obtuse) and floral bracts broadly ovate and persistent (vs. lanceolate and early deciduous). The former is confined to México; the latter to the U.S.A. (Figure 1).
Ramamoorthy (by annotation) first called attention to this specific novelty, and I have retained the name which he proposed. He did not, however, recognize its affinities nor provide information as to its sectional affiliation; these are my own. The epithet honors Jaime Hinton, son of the late George Hinton, renown collector of Mexican plants.
Tumer. New Salvia from México
/ ae 7
ny WZ v= :
VEY, YE
Figure 1. Salvia jaimehintoniana, from holotype.
| 100 PHY TOLOGIA August 1995 volume 79(2):97-101 |
x : wey HL. au int Bere
Figure 2. Distribution of Salvia azurea (open circles, in Texas; it is absent in México); and S. jaimehintoniana. Based upon specimens at LL, TEX.
Turner: New Salvia from México 101
ACKNOWLEDGMENTS
I am grateful to Gayle Turner for the Latin diagnosis, and to her and Piero Delprete for reviewing the manuscript.
LITERATURE CITED
Epling, C. 1939. A revision of Salvia, subgenus Calosphace. Feddes Repert. Sp. Nov. Beith. 110: 1-388.
Epling, C. 1940. Supplementary notes on American Labiatae V. Bnittonia 7:129- 142.
Phytologia (August 1995) 79(2):102-107.
A NEW SPECIES OF LUPINUS (FABACEAE) FROM OAXACA, MEXICO: A SHRUB OR TREE MOSTLY THREE TO EIGHT METERS HIGH
B. L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A new species, Lupinus jaimehintoniana B.L. Turner is described and photographed from near timberline on Cerro Quiexobra, Oaxaca, where it is a subdominant shrub or small tree up to 8 m high, the lower trunks markedly woody and up to 30 cm across. It appears to be closely related to Lupinus montanus, having most of the characters of that species, except for its woody habit, much-reduced leaves and stipules, silvery pubescence, and longer floral bracts.
KEY WORDS: Fabaceae, Lupinus, México, Oaxaca, systematics
Identification of Mexican plants has revealed the following novelty.
LUPINUS JAIMEHINTONIANA B.L. Turner, spec. nov. Figures 1, | TYPE: MEXICO. Oaxaca: Mpio. Miahuatldn, Cerro Quiexobra, 3575 m, 15 Oct 1995, Hinton et al. 26160 (HOLOTYPE: TEX!; Isotype: TEX!).
Similis Lupino montano H.B.K. sed frutex vel arbor est, 8 m alto, foliis calium superorum 5-7 foliola habentibus, stipulis 2-10 mm secus petiolos connatis, indumento pilorum curtorum et argenteorum sursum appresso.
Shrubs or trees mostly (1-)3-8 m high, the lower trunks up to 30 cm across. Flowering stems (of new growth) nodose, the vestiture of short silvery, upwardly | appressed hairs ca. 0.2 mm long. Leaves at 2-4 nodes below the inflorescence having — 5-7 leaflets; stipules 2-3 cm long, fused at the base to the petioles for 2-10 mm; petioles 4-7 cm long; leaflets narrowly elliptic, 3-6 cm long, 0.6-1.2 cm wide, , moderately silky appressed-pubescent on both surfaces, the apices acute. Inflorescence a terminal spike 15-30 cm long, ca. 5 cm across. Bracts linear- lanceolate, much exceeding the flowers, markedly pubescent with appressed hairs above and below, the apices mostly narrowly acuminate. Pedicels mostly 7-8 mm |
102
Turner: New Lupinus from Oaxaca 105
long, pubescent like the stems. Calyx with lower sepals lanceolate ca. 9 mm long (including the short tube), upper (united) lobes broadly ovate, ca. 6 mm long. Corollas reportedly purple; wing petals with claws ca. 3 mm long; blades broadly oval, glabrous, ca. 13 mm long, 9 mm wide, the upper 1/3 corrugate near its base; banner glabrous throughout, sessile or nearly so, ca. 12 mm long and as wide; keel petals glabrous with claws ca. 4 mm long, their blades ca. 7 mm long as measured along the basal axis, then sharply arcuate upwards, the apical axis nearly at nght angle to that of the basal axis, the apices acute. Lower stamens with anthers ca. 2 mm long. Pods (immature) ca. 4 cm long, 0.8 cm wide, densely tomentose with contorted subtawny hairs. Mature seeds not available. ,
ADDITIONAL SPECIMENS EXAMINED: MEXICO. Oaxaca: Mpio. Miahuatldn, Quiexobra, 3070 m, 19 Oct 1995, Hinton et al. 26228 (TEX); 35 km ESE of Miahuatlén, 5 km NE of Santo Domingo Ozolotepec, Cerro Quiexobra, “Timberline vegetation in open glades along ridges and in mountain saddles”, 3650-3800 m, 10 Dec 1989, McDonald 2923 (TEX).
When first collected by Dr. Andrew McDonald (collection cited above), perhaps the first botanist to collect on Cerro Quiexobra, I was too busy with other projects to pursue its identification. Had I known its remarkable habit (as shown in figures 1 and 2) I most certainly would have sought its identity, although McDonald did describe or label the collection as being “Common subarborescent shrubs often forming dense stands along ridges, 1-3 m tall.”
Lupinus jaimehintoniana appears to belong to the L. montanus Cerv. ex Lag. species complex, which was treated in some detail by Dunn & Harmon (1977). These authors recognized five species in the complex, one of these L. montanus having five infraspecific categories. Most of these taxa are confined to México and Guatemala and most were onginally accepted as “good” species by yet earlier well known mavens of the genus in North Amenica, mainly, C.P. Smith. As species are defined by most current workers in Lupinus, the vanous segregates from L. montanus (s.1.) rendered by Dunn & Harmon are about as distinct as yet other species in this or that complex. Nevertheless, were it not for the extraordinary habit of L. jaimehintoniana | would probably have followed Dunn & Harmon in recognizing it as but another infraspecific category of L. montanus, although having stipules quite different from the latter. In their treatment (1977) L. jaimehintoniana will key to L. montanus var. nelsonii (Rose) C.P. Smith, a taxon known only from eastern Oaxaca, mainly in the pine-fir forests in the sierras to the east of Cd. Oaxaca. In addition to its small stipules, it differs from L. montanus in having a silvery upturned vestiture on its stems (vs. tawny and down- curved or glabrous), and mostly longer floral bracts with somewhat contorted apices. Lupinus montanus is consistently described as a coarse herb or shrub 1-2 m high, the stems fistulose.
According to Dunn (1984), the largest lupine known to him at that time was a collection of Lupinus (the species not named) from Pert, said to be about 4.5 m hi gh and possessing pendant flowering branches. As shown in figures 1 and 2, Lupinus Jjaimehintoniana can develop into relatively large trees, the inflorescences clearly borne terminal and erect. .
106 PHYTOLOGIA August 1995 volume 79(2): 102-107
As communicated by Jaime Hinton (nearly 80 years of age at the time of his ascent —
of Cerro Quiexobra, accompanied by his faithful friend and colleague, Anacleto Lugo):
. we trailed McDonald’s [Dr. Andrew McDonald, currently Research Associate at Harvard University, who made the first extensive collections from Cerro Quiexobra in 1990] redoubtable footsteps over Quiexobra and up to the top of La Sirena (where, two years ago, a great fire reduced the four summits to tall grass and a few trees) .. . [I found myself] staring in “wild surmise” at the Lupine trees, as astonishing to the density of my ignorance as sudden fire to the human skin. (One badly burned and dying old Lupine graciously balanced its thirty feet of height on a real wooden trunk twelve inches across.)
Never in Quiexobra could I grow used to the miraculous blue Lupines held with such accomplished and heart-wringing majesty so high up in the air, as if, by God, who could ever have doubted that a Lupine could even more easily become a magnificent tree than a lovely herb? And I gazed with undying wonder at the beige-colored trunks of dead Lupines burning in the huge fires we built against the icen winds that drove us to bed, only a bit after sundown, as they mercilessly swept up at us from those Oaxacan sierras that by daylight appear tossed like the most gorgeous heaps of pale blue jewels across the whole wide northern world.
I reckon, Billie, Quiexobra does show the mightiest pines and firs still standing in Mexico. And I must admit my wonder at how well the Zapotec’s earthen superstition has protected those mossy sprawling _ giants (Chirathrodendron pentadactylon, famed and held in awe for its flowers shaped like little human hands, richly yellow on one side and richly scarlet on the other) from the last few hundred years of mindless ax and fire.
Alas, the enterpnsing marihuaneros of to-day are no more able to control the rampage of the fires they themselves set to burn the forest for their secret and forbidden plantations, than their fabled stone gods were able to control the marauding rampage of the fiery Ibenans.
Judging from its local abundance on Cerro Quiexobra, its resistance to fire and its adaptation to near timberline climates in México, it is likely that the species will prove hardy in the more temperate regions. At least it should prove interesting to ascertain through DNA analysis, efc., what genes might be involved that permit its development into such a bizarre woody member of this otherwise mostly herbaceous genus.
It seems fitting that this remarkable lupine should bear the name of James Hinton, for he has collected with his father numerous lupine species, many of which are types and some of which already bear their names, including Lupinus hintonii C.P. Smith (for G.B. Hinton, the elder) and L. hintoniorum B.L. Tumer (for the extended family).
Tumer: New Lupinus from Oaxaca 107
ACKNOWLEDGMENTS
I am grateful to Gayle Tumer for the Latin diagnosis and to Piero Delprete and Mark Mayfield for reviewing the manuscript.
LITERATURE CITED
Dunn, D.B. 1984. Genetic resources cytotaxonomy and distribution of New World lupin species. A paper presented at the 3rd Intemational Conference on lupines, 4- 8 Jun 1984, La Rochelle, France (copy on file at TEX!).
Dunn, D.B. & W.E. Harmon. 1977. The Lupinus montanus complex of Mexico and Central America. Ann. Missouri Bot. Gard. 64:340-365.
Phytologia (August 1995) 79(2):108-113.
NOTES ON COSTA RICAN PEPEROMIA (PIPERACEAE), INCLUDING FOUR NEW SPECIES
Michael H. Grayum
Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri 63166-0299 U.S.A.
ABSTRACT
Four new, ostensibly endemic, Costa Rican species of Peperomia are descnbed: Peperomia hammelii Grayum, P. saintpauliella Grayum, and P. trichomanoides Grayum are all terrestnal or epilithic species restnicted to the southern Pacific slope, mainly in limestone habitats. Peperomia ursina Grayum comprises epilithic or epiphytic plants from the lower Atlantic slope of the Cordillera de Talamanca. Peperomia tenuifolia C. DC., heretofore considered a synonym of P. lignescens C. DC., is reinterpreted as an older name for the species heretofore called P. killipii Trel. Revised synonymies are provided for P. lignescens and P. tenuifolia.
KEY WORDS: Costa Rica, Peperomia, Piperaceae, systematics
The Costa Rican members of the huge, pantropical genus Peperomia (Piperaceae) | were most recently treated by Burger (1971), who accounted for 66 species. Since that time, intensive collecting efforts in previously underexplored regions of the country (particularly the Cordillera de Talamanca) have resulted in the addition of at least fifteen species to this total. Most of these additions have involved species already . described from other countries, but a few appear to represent new taxonomic entities. Four new Costa Rican species of Peperomia are described hereunder, and a fifth additional species is freed from synonymy under a name accepted by Burger.
PEPEROMIA HAMMELII Grayum, spec. nov. TYPE: COSTA RICA. Puntarenas: Canton de Osa, Fila Costefia, Fila Cruces, cabeceras del Rio Piedras |
Blancas, Cerro Anguciana, faldas al oeste, bosque en roca de cal, 8° 48’ 56” N,
83° 10’ 37” W, 1,400-1,600 m, 10 Dec 1993, Hammel 19274 (HOLOTYPE: - INB!; Isotypes: BM!,COL!,CR!,F!,MO!).
108
Grayum: Notes on Costa Rican Peperomia 109
P. lignescens C. DC. affinis, a que imprimis differt caule trichomatibus multiseriatis vesicariis vestito petiolis in longitudinem late alatis laminis foliorum hirsutis pedunculis longioribus.
Plants terrestrial or epilithic. Stems erect, to ca. 23 x 0.2-0.5 cm, densely clothed with stout, multiseriate, inflated hairs to ca. 1 mm long. Leaves alternate. Petiole 1.2- 4.6 cm, broadly alate throughout its length, ca. 2-6 mm wide, hirsute on both sides. Lamina 5.5-11.0 x 2.5-7.2 cm, ovate to broadly elliptic, impeltate, broadly cuneate to rounded or subcordate at base, subacute to subacuminate apically, pinnately nerved with ca. 5-6 primary lateral veins per side, dark-gland-dotted and hirsute on both surfaces. Inflorescences solitary at stem apex. Peduncle 2.7-3.8 cm, to ca. 1 mm wide, glabrous or with few scattered hairs toward base. Spike 1.7-8.5 x 0.2-0.4 cm, white. Flowers moderately separated; rachis glabrous; bracts 0.5-0.6 mm _ wide, suborbicular, densely glandular-punctate; anthers broadly elliptic to oblong, ca. 0.25 mm. Fruits unknown.
Peperomia hammelii is known only from the type locality, on the western slope of Cerro Anguciana, the highest peak in the Fila Costefia in the southern Pacific region of Costa Rica. Here, it grows on or near limestone cliffs or outcrops at 1,400-1,600 m elevation.
Peperomia hammelii is an unusually well-marked species in uniquely combining two features which, even by themselves, are anomalous within the genus: an indument of odd, inflated hairs, and broadly and extensively alate petioles. In its terrestrial or epilithic habitat, erect, caulescent habit, alternate, pinnately veined leaves and dark, sessile laminar glands it most closely resembles P. lignescens C. DC. and allies, to which it is perhaps intimately related. Peperomia lignescens, which is parapatric and at least conceivably syntopic with P. hammelii, differs from the latter in having generally puberulent or glabrescent (rather than hirsute) foliage and shorter peduncles (in addition to the characters mentioned previously).
I take great pleasure in dedicating this new species to its discoverer, Dr. Barry E. Hammel of the Missouri Botanical Garden, a long-time student of the Neotropical flora and my colleague on the “Manual to the Plants of Costa Rica” project.
Numerous Costa Rican collections have accrued in recent years of yet another Peperomia species that agrees in a general way with the description of P. lignescens, but which differs in having consistently palmate leaf venation. These collections are all from the humid Pacific lowlands (O-1,600 m), south from the Rio Grande de Tarcoles. They key out easily to Peperomia killipii Trel. in Yuncker’s (1950) Flora of Panama treatment, and are an excellent overall match for the holotypes of P. killipii and its synonym (fide Yuncker) P. hymenodes Trel.
Peperomia lignescens was not treated by Yuncker (1950), while P. killipii was only briefly mentioned by Burger (1971: 65) in comparison with P. pseudodependens C. DC. (=P. asarifolia Schltdl. & Cham.), a somewhat similar species that also has palmate venation. Due to the venation difference, P. killipii will not key out anywhere near P. lignescens in Burger’s (1971) treatment. Nevertheless, type material of both Peperomia aguacatensis C. DC. and P. tenuifolia C. DC., two of the five heterotypic names listed in synonymy under P. lignescens by Burger (1971), agrees in all critical
110 PHYTOLOGIA August 1995 volume 79(2): 108-113
details with that of P. killipii. As both P. aguacatensis and P. tenuifolia substantially —
predate P. killipii, the last-mentioned name must fall into synonymy.
The following paragraphs provide what | presently consider to be complete synonymies for the two species I propose be called Peperomia lignescens C. DC. and
P. tenuifolia C. DC. This is necessary not only to clanfy the confusion detailed —
above, but also to establish precedent in two cases of equal priority.
PEPEROMIA LIGNESCENS C. DC., J. Bot. 4:137. 1866. Peperomia carlosiana C. DC., J. Bot. 4:140. 1866.
Peperomia carthaginensis C. DC., Linnaea 37:377. 1872. Peperomia lignescens
C. DC. var. carthaginensis (C. DC.) Trel., Contr. U.S. Natl. Herb. 26:193. 1929. Peperomia lignescens C. DC. var. subcuneilimba Trel., Contr. U.S. Natl. Herb. 26:193. 1929.
Peperomia jilotepequeana Trel. & Standl. in Standl. & Steyerm., Fieldfana, Bot. 24(3):254. 1952.
PEPEROMIA TENUIFOLIA C. DC., Linnaea 37:371. 1872. Peperomia aguacatensis C. DC., Linnaea 37:376. 1872. Peperomia killipii Trel., Bot. Gaz. 73:143. 1922. Peperomia hymenodes Trel., Contr. U.S. Natl. Herb. 26:43. 1927.
Peperomia tenuifolia differs from P. lignescens not only in its palmate leaf venation, but also in its usually epiphytic habit (it may occasionally be epilithic), absence of conspicuous dark, sessile laminar glands, and minutely papillate inflorescence rachis. Furthermore, it is a species of generally lower elevations (though there is considerable overlap). I select the names P. lignescens and P tenuifolia because they have already been more widely applied in herbaria than their alternatives, and because both P. carlosiana and P. aguacatensis are inappropriate toponyms.
PEPEROMIA SAINTPAULIELLA Grayum, spec. nov. TYPE: COSTA
RICA. Puntarenas: along short-cut road to Golfito from Villa Bricefio on Interamerican Hwy., W side of Fila Gamba, ca. 6 km from Golfito airport, 8° 41’
30” N, 83° 12’ W, < 100 m, 6 Mar 1985, Croat & Grayum 59911 (HOLOTYPE: CR!; Isotypes: BM!,MO!).
P. insueta Trel. affinis, sed differt laminis foliorum (1.7-)2.0-3.9 cm longis ovatis vel suborbicularis pedunculis 1.6-3.7 cm longis spicis 7.3-16.1 cm x 0.4-1.0 mm.
Plants terrestrial or epilithic. Stems erect to + decumbent, 0.8-1.3. x 0.2-0.3 cm. Leaves alternate in basal rosette. Petiole 1.3-7.4 cm, spreading-hirsute with uniseriate hairs. Lamina (1.7-)2.0-3.9 x 1.80-4.65 cm, broadly ovate to orbicular (or rarely obovate), impeltate, cordate or (rarely) subsagittate at base with sinus to 0.7 cm deep
and posterior lobes rounded to subtruncate or (rarely) obtuse, nearly truncate or —
Grayum: Notes on Costa Rican Peperomia 111
rounded to obtuse apically, palmately (5-)7(-9)-nerved, pellucid-gland-dotted on both surfaces, sparsely to moderately hirsute on both sides (especially along major veins abaxially). Inflorescences solitary, basal. Peduncle 1.6-3.7 cm, with hairs like petiole. Spike 7.3-16.1 cm x 0.4-1.0 mm, pinkish. Flowers + crowded at first,
becoming distant; rachis virtually glabrous; bracts 0.4-0.5 mm wide, + peltate, elliptic, covered with orange, sessile glands; anthers broadly elliptic, ca. 0.2 mm. Fruits ca.
0.5-0.6 x 0.4-0.5 mm, + globose-bodied, broadly narrowed to substipitate base, beakless; stigma apical. ,
Additional specimens examined. COSTA RICA. Puntarenas: Cant6n de Osa, forest along Quebrada Benjamin, near crossing of trail from Palmar Norte to Jalisco,
8° 58’ N, 83° 28’ W, ca. 160 m, 14 Dec 1989, Grayum & Hammel 9543 (BM,INB, MO); Cant6n de Osa/Buenos Aires, western part of main ndge of Fila Retinto, along and near trail (not on current maps) from Palmar Norte to Jalisco, 8° 59’ 30” N, 83°
28’ W, ca. 780-960 m, 9 Dec 1988, Grayum & Herrera 9150 (MO).
Peperomia saintpauliella is apparently confined to a small area to the north and east of Golfo Dulce in Puntarenas Province, from near Palmar Norte to the vicinity of Golfito. Here, it grows near forest creeks, often on vertical rock (usually specified as limestone) faces, at ca. 50-800 m elevation.
Peperomia saintpauliella comprises smallish, acaulescent plants with impeltate, suborbicular leaves and solitary, basal inflorescences. As the specific epithet implies, living specimens bear a strong vegetative resemblance to smaller forms of the cultivated African violet (Saintpaulia ionantha H. Wendl.), and have a similarly compact, ornamental appearance. Living material of P. saintpauliella is in cultivation at the Missouri Botanical Garden, and plants have been put on display in the Climatron.
Other Peperomia species most resembling P. saintpauliella are the Colombian P. macrotricha C. DC. and the Panamanian P. umbrigaudens Yunck. and, especially, P. insueta Trel. The last-mentioned species differs from P saintpauliella in having longer
(4.0-7.5 cm), narrowly elliptic to + ovate leaf blades, absolutely and relatively much
longer peduncles (about as long as the spikes), and generally shorter and thicker spikes (6-12 cm x 1.0-1.5 mm).
PEPEROMIA TRICHOMANOIDES Grayum, spec. nov. TYPE: COSTA RICA. Puntarenas: Cantén de Osa, Fila Costefia, Fila Cruces, cabeceras del Rio
Piedras Blancas, Cerro Anguciana, faldas al Oeste, bosque en roca de cal, 9° 48’
56" N, 83° 10’ 37” W, 1,400-1,600 m, 10 Dec 1993, Hammel 19273 (HOLOTYPE: INB!; Isotypes: BM!,CR!,MO!).
Differt a P. saintpauliella Grayum dimensionibus uniformiter parvioribus pedunculis relative longioribus rhachidi inflorescentiae dense pubescenti; a P. tuerckheimii C. DC. laminis foliorum impeltatis basi cordatis relative latioribus venis primariis basalibus plerumque 7.
ii PHYTOLOGIA August 1995 _ volume 79(2):108-113
Plants epilithic, the leaves and spikes flattened against rock. Stems short and thick, subcormose, ca. 0.2-0.8 x 0.15-0.25 cm. Leaves apparently alternate, in basal rosette. Petiole 0.3-1.9 cm, spreading-hirsute with uniseriate hairs. Lamina 0.5-1.8 x 0.5-1.8 cm, broadly ovate to suborbicular or subreniform, impeltate or scarcely peltate, cordulate or cordate at base with sinus to 0.25 cm deep and posterior lobes rounded to subtruncate, broadly rounded to subacute apically, palmately 3-5-nerved, pellucid-gland-dotted at least above, appressed-hirsute on both sides (more sparsely so above). Inflorescences solitary, basal. Peduncle ca. 1.3-4.1 cm, pubescent as petiole.
Spike 2.7-7.0 cm x 0.2-0.9 mm. Flowers becoming distant; rachis + densely
spreading-pubescent; bracts 0.3-0.4 mm wide, + peltate, suborbicular, densely dark- pellucid-punctate; anthers broadly elliptic-oblong to suborbicular, ca. 0.25-0.30 mm. Fruits ca. 0.5-0.6 x 0.3-0.4 mm, ellipsoidal to subglobose, narrowed to substipitate base, beakless; stigma apical.
This species is known only from the type locality, at 1,400-1,600 m eleVation on the steep limestone ramparts of Cerro Anguciana, the highest peak in the Fila Costefia of southern Pacific Costa Rica.
Peperomia trichomanoides is so named because its habitat (epilithic and growing among mosses), appressed habit, and small size recall some species of the fern genus Trichomanes L. (Hymenophyllaceae). Plants of this species resemble, in general aspect, miniature versions of P. saintpauliella (descnbed above), from which they differ not only in their uniformly smaller dimensions, but also in having proportionately longer (relative to the spike) peduncles and densely pubescent (rather than essentially glabrous) inflorescence rachises. In the latter respect, P. trichomanoides approaches some specimens of P. tuerckheimii C. DC. (including P. hispidorhachis Y unck. and P. tecticola C. DC.), another small calciphile that occurs in the same vicinity; however, P. tuerckheimii has clearly peltate, non-cordate, more elongate leaf-blades with generally 7 (rather than 5) primary basal veins.
PEPEROMIA URSINA Grayum, spec. nov. TYPE: COSTA RICA. Lim6n: Cordillera de Talamanca, along mdge descending to main fork of Quebrada Cafiabral from divide between basin of Rio Madre de Dios and that of Rio Barbilla, 10° 02’ N, 83° 25’ W, 280-400 m, 6 Sep 1988, Grayum, Herrera, & Robles 8842 (HOLOTYPE: INB!; Isotypes: BM!,COL!,F!,MO!).
Differt a P. alata Ruiz & Pav. pubescentia dense uniformiterque hirsuta; a P. tuisana C. DC. atque P. montecristana Trel. petiolis brevioribus inflorescentiis multo brevioribus.
Appressed-climbing trunk epiphytes or epilithic, stoloniferous. Stems erect to + decumbent, ca. 2-8 x 0.10-0.15 cm, spreading-hirsute with uniseriate hairs. Leaves alternate. Petiole 0.1-0.3 cm, pubescent as stems. Lower leaves + reduced; medial and distal laminae 1.0-3.6 x 0.5-1.5 cm, narrowly elliptic to rhombic, impeltate, acute at base, subacute to subacuminate at apex, + obscurely 3-nerved from base, hirsute on both surfaces. Inflorescences solitary at stem apex. Peduncle 0.1-1.0 cm, spreading-
Grayum: Notes on Costa Rican Peperomia 113
hirsute. Spike 1.3-5.0 cm x 0.7-1.5 mm, yellow-green. Flowers moderately separated; rachis glabrous; bracts 0.3-0.4 mm wide, suborbicular, densely glandular- punctate; anthers broadly elliptic, 0.15-0.25 mm. Fruits ca. 0.5-0.6 x 0.5-0.6 mm, globose-bodied, rounded at base, exserted on triangular stipe ca. 0.5-0.6 mm, papillate, with stout, conical beak to ca. 0.15 mm.
Additional specimens examined. COSTA RICA. Limon: Reserva Indigena Talamanca, camino a Soki entre la Quebrada Amubri, margen izquierda de Rio Lan, 9° 29’ 40” N, 82° 59’ 40” W, 200 m, 28 Jun 1989, A. Chacdn 20 (BM,CR,MO).
As far as is presently known, Peperomia ursina is restricte to the Atlantic slope of the Costa Rican Cordillera de Talamanca from ca. 200-400 m. According to collectors’ notes, the plants may be either epilithic or epiphytic on trunks.
Peperomia ursina is most similar and perhaps most closely related to P. alata Ruiz & Pav. and allied species characterized by alternate, distichous leaves with thin, -palmately veined blades, and solitary inflorescences. It differs sharply from most species in this group in its dense, uniform hirsute pubescence, reflected in the specific epithet. This species will key to the vicinity of P. tuisana C. DC. and P. montecristana -Trel. in Burger’s (1971) treatment of Costa Rican Piperaceae, but differs from both in its shorter petioles and much shorter inflorescences.
ACKNOWLEDGMENTS
Field work resulting in the discovery of all four new species described in this paper was supported by National Geographic Society grants 3317-86 and 4682-91, to the author, and National Science Foundation (NSF) grant BSR-8700068, to the author and B.E. Hammel. Publication was supported by NSF grant DEB-9300814, to B.E. Hammel and the author. I thank Ramblin’ Joe Evans and George E. Schatz for their critical reviews of the manuscript.
LITERATURE CITED
Burger, W. 1971. Piperaceae. Jn, W. Burger (editor), Flora costaricensis.
: Fieldiana, Bot. 35:5-218.
Yuncker, T.G. 1950. Piperaceae. In, R.E. Woodson, Jr. & R.W. Schery (editors), Flora of Panama. Ann. Missouri Bot. Gard. 37: 1-120.
Phytologia (August 1995) 79(2):114-122.
NEW ADDITIONS TO THE GENUS PINGUICULA (LENTIBULARIACEAE) OF MEXICO
Hans Luhrs Krayenhoffstr. 51, 1018 RJ Amsterdam, HOLLAND
ABSTRACT
Two new species of Pinguicula from México are described and illustrated: P. stolonifera (subgen. Pinguicula) from the state of Oaxaca, and P. laxifolia (subgen. Pinguicula) from the state of Tamaulipas. Pinguicula stolonifera belongs to the section Orcheosanthus, and subsect. Caudatopsis. A new section (Orchidioides) is proposed to include P. laxifolia. The taxonomic status of P. jorgehintonii B.L. Turner, P. hintoniorum B.L. Turner, and P. reticulata Schlauer is discussed. They are considered to be synonymous with previously described species.
KEY WORDS: Lentibulariaceae, Pinguicula, Flora of México, systematics
Research on the extensive herbarium collections of the genus Pinguicula, from the University of Texas, has revealed the following results.
Pinguicula stolonifera Luhrs, spec. nov. (Figure 1). TYPE: MEXICO. Oaxaca: ca. 3 km. se. of Ixtlan de Juarez, on steep banks in pine and oak woods, ca. 2300- 2400 m, 14 Aug. 1966, R.W. Cruden 1177 (HOLOTYPE: TEX! 271238); sub nomine P. oblongiloba DC. Det.: S. Zamudio 1989.
Herba perennis, stolonifera; stolones flagelliformis, cerasini, usque ad 8 cm longis. Rhizoma simplex brevis, radicibus adventitiis numerosis funiformibus. Folia radicalia rosulata, biformia; rosula “hiemis” numerosa - 36, crassa, ovata vel lanceolata, acuta, 4-7(-12) mm longa, 1-3 mm lata, facie concava; rosula “aestatis” 4-7, distincte petiolata, petiolo erecto, 11-18 mm longo, 2-3 mm lato, margine ciliato, lamina lanceolata vel anguste oblongo- ovata, acuta, basin versus angustata, margine provunde involuta, superne glandulis sessilibus et glandulis stipitatis dense vestita, 18-33 mm longa, 6-12 mm lata. Hibernacula nulla; gemmatae. Pedicelli 1-3 erecti, cerasini, apicem versus glandulis stipitatis disperse obsiti, 105-164 mm alti, uniflon. Flores 39-51 mm longi (calcari incluso). Calyx bilabiatus, extus glandulis stipitatis
114
Luhrs: New species of Pinguicula from México 115
obsitus; labium superum trlobum, lobis anguste ovatis, 3 mm longis, 2 mm latis; labium inferum usque ad dimidium longitudinis bilobum, lobis ovatis, 2.5 mm longis, 1 mm latis. Corolla ringens, profunde bilabiata, magentea, labio infero ad basi striata alba ornato, extus glandulis stipitatis vestita; labium superum bilobum, lobis oblongo-ovatis, 10-14 mm longis, 5-9 mm latis; labium inferum profunde tripartitum, lobis lateralibus oblongo-lanceolatis, apicem versus angustatis, 11-16 mm longis, 4-7 mm latis, lobo intermedio paulo major 15-19 mm longo, 45 mm lato. Tubus_brevissimus infundibuliformis, 3-4 mm longus, intus pilosus, pilis cylindrico-subulatis, sine palato. Calcar cylindrico-acuminatum, sinuatum, 18-26 mm longum, cerasinum. Ovarium subglobosum, glandulis stipitatis obsitum. Stigma bilabiatum, purpureum, labio infero maximo, suborbiculato, fimbriato.
Capsula ovoidea, + 4 mm _ longa, glandulis stipitatis parum_ obsita. Florescentia VIII-IX.
Perennial herb, stoloniferous; stolons whip-like, cherry-red, up to 8 cm long, bearing up to 4 non glandular leaves (2-4[-6] mm long) along its length. Stem short, with numerous adventitious fibrous roots. Leaves rosulate, dimorphic; the leaves of the winter rosette numerous -36, thick, ovate or lanceolate, acute, 4-7(-12) mm long, 1-3 mm wide, concave; the leaves of the summer rosette 4-7, with a distinct petiole, erect, 11-18 mm long, 2-3 mm wide, margin ciliate, lamina lanceolate or narrowly oblong-ovate, acute, narrowing towards the base, margin deeply involute, the upper surface densely covered with sessile and stipitate glands, 18-33 mm long, 6-12 mm wide. Hibernaculum absent; provided with gemma-like buds. Scapes 1-3, erect, cherry-red, the upper part dispersedly stipitate glandular, 105-164 mm all, 1- flowered. Flowers 39-51 mm long, including the spur. Calyx bilabiate, stipitate glandular; upper lip 3-lobed, the lobes narrowly ovate, 3 mm long, 2 mm wide; lower lip divided to the middle into 2 lobes, the lobes ovate, 2.5 mm long, 1 mm wide. Corolla deeply bilabiate, red-purple, the base of the lower lip marked with a white vertical streak, the outer surface stipitate glandular; upper lip 2-lobed, the lobes oblong-ovate, 10-14 mm long, 5-9 mm wide; lower lip deeply 3-lobed, the lateral lobes oblong-lanceolate, narrowing towards the apex, 11-16 mm long, 4-7 mm wide, the middle lobe slightly larger, 15-19 mm long, 4-5 mm wide. Tube extremely short, funnel-shaped, 3-4 mm long, with cylindnical-subulate hairs inside, palate absent. Spur cylindrical-acuminate, 18-26 mm long, cherry-red. Ovary subglobular, stipitate glandular. Stigma bilabiate, purple, the lower lip much larger, suborbiculate, margin
fimbriate. Capsule ovoid, + 4 mm long, slightly stipitate glandular. Florescence August-September.
ADDITIONAL MATERIAL EXAMINED: MEXICO. Oaxaca: Distr. Ixtlan,
Sierra de Judrez, ne. of C. Pelén, on a steep loamy bank, + 2700 m, 29 Sep. 1991, Luhrs et al. 9105 (Herb. Luhrs); In umbrosis Totontepeque, Hartweg 509 (L).
Pinguicula stolonifera belongs to the section Orcheosanthus because of the deeply bilabiate corolla, the extremely short funnel-shaped tube, and the very large spur. Within this it is placed in the subsect. Caudatopsis, together with P. macrophylla H.B.K. and P. oblongiloba DC., because of the lanceolate or broadly ovate, acute or acuminate winter leaves, and long petiolate summer leaves as defined in Casper’s monograph of the genus Pinguicula (1966a). It shows some affinity with P.
116 PHYTOLOGIA August 1995 volume 79(2): 114-122
oblongiloba, especially on behalf of the corolla lobes (Hinton et al. 14504 [TEX]). Although it has been identified with P. oblongiloba by S. Zamudio, it differs from the latter by having much shorter, narrower, and deeply involute summer leaves, and by forming gemma-like buds, putting forth long whip-like runners, budding at the end of its tip, a feature which is unique in the Mexican pinguiculas and is known (in a much shorter stolon-like manner in P. calyptrata H.B.K. from Ecuador, and P. vallisneriifolia Webb from Spain. Unfortunately these differences are not easily observed when the plants are dried, resulting in