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STOR ® Robert J Systematics of the "Epiles" Group of Poa (Poaceae) STOR ® Robert J. Soreng Systematic Botany, Vol. 16, No. 3. (Jul. - Sep., 1991), pp. 507-528. Stable URL: http://links.jstor.org/sici?sici=0363-6445%28199107%2F09%2916%3A3%3C507%3ASOT%22GO%3E2.0.CO%3B2-T Systematic Botany is currently published by American Society of Plant Taxonomists. Your use of the JSTOR archive indicates your acceptance of JSTOR' s Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http ://w w w .j stor. org/j ournals/aspt .html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is an independent not-for-profit organization dedicated to creating and preserving a digital archive of scholarly journals. For more information regarding JSTOR, please contact [email protected]. http: //www .j stor. org/ ThuOct 19 16:13:38 2006 Systematic Botany (1991), 16(3): pp. 507-528 © Copyright 1991 by the American Society of Plant Taxonomists Systematics of the "Epiles" Group of Poa (Poaceae) ROBERT J. SORENG Biology Department, New Mexico State University, Las Cruces, New Mexico 88003 Present address: L. H. Bailey Hortorium, Cornell University, Ithaca, New York 14853 ABSTRACT. The taxonomy of the western North American "Epiles" group of Poa is revised in the light of new information on morphology, anatomy, breeding systems, and chloroplast DNA restriction sites. The artificial group, "Epiles" s.L, of A. S. Hitchcock is divided into three homo- geneous parts: sect. Madropoa subsect. Epiles, sect. Secundae subsect. Halophytae (all formalized here), and sect. Abbreviatae. These three sections belong to three larger chloroplast groups in Poa. A new subspecies, P. cusickii subsp. pallida, is proposed, and hybrids between this and P. fendleriana are distinguished as P. x nematophylla. From the Sierra Nevada, P. stebbinsii is proposed as a new name for P. hansenii sensu auct., and a new species, P. keckii, is segregated from P. suksdorfii. A descriptive key to western North American "Epiles" group s.L, distributional maps, and illustrations of species of the subsect. Epiles s. str., and species newly referred to sect. Abbreviatae are provided. New and additional chromosome counts are reported for: P. cusickii subsp. pallida, In = 56+11; P. secunda (P. juncifolia form, previous citation as P. cusickii), 2M = 42; P. napensis, In = 42; P. strictiramea, In = 28+1; and P. unilateralis, In = 84. Much confusion has surrounded the species proposed. The two species have been well col- included within the "Epiles" group of Poa L. as lected by botanists in the high Sierra Nevada, proposed by A. S. Hitchcock (1935). The group, but had not been examined in detail. Because as originally defined, is clearly artificial, in- good descriptions and illustrations are already cluding all perennial species of North America available for the other taxa in the group (Marsh with a cespitose habit and keeled glabrous lem- 1952; Hitchcock 1969), only a descriptive key is mas. Loss of lemma pubescence has occurred in presented. many diverse groups of Poa around the world, even within species. The fact that several MATERIALS AND METHODS "Epiles" taxa sometimes have pubescent lem- mas has led to frequent misidentifications and Leaf epidermal peals were prepared by soft- confusion about relationships among the spe- ening tissues with Pohl's softener, scraping away cies. Difficulties have been compounded by in- the opposing epidermis and mesophyll with a terspecific hybridization and apomixis in some razor, mounting the exposed epidermis on a species, and by a lack of detailed circumscrip- slide, staining it with safranin, and clearing the tion and biological understanding of the taxa tissue with glycerol. Chromosome counts were involved. Overly simplified keys have also hin- made by fixing bud material in Carnoy's Fixa- dered identifications in this very complex group. tive and staining pollen mother cells in aceto- The objectives of the present paper are to re- carmine. Breeding system phenotypes were de- define the limits of the "Epiles" group, formally termined by surveying herbarium collections describe subgeneric taxa and two new species, and population samples for intraspecific pat- and present a biological interpretation of the terns of deviation from hermaphroditism, in- species based on new knowledge of their chlo- dicating gynodioecism or dioecism, or, when roplast origins (Soreng, in press), anatomy, all individuals are pistillate, apomixis (Soreng morphology, cytology, breeding systems, and 1986). Chloroplast DNA methods, results, and geography of the apomicts (Soreng 1986). Pop- phylogenetic analysis are reported by Soreng ulations from the Sierra Nevada previously re- (in press). ferred to as Poa hansenii Scribner and P. suksdorfii Specimens examined in this study were bor- (Beal) Vasey ex Piper represent new species. In rowed from ARIZ, ASU, CAN, CAS, DS, GH, recognition of two renowned Californian bot- ID, MICH, MSC, MO, NMC, NY, ORE, OSC, anists, the names P. keckii and P. stebbinsii are TAES, TEX, UBC, UC, US, V, WIS, WSU, WTU, 507 508 SYSTEMATIC BOTANY [Volume 16 WYAC, or collected in situ. Lists of exsiccatae to 1000s of years), and are treated taxonomically will be deposited at MO. as if they comprised subsets of populations, or races, within sexual species, but with inter- breeding events occurring infrequently and RESULTS AND DISCUSSION when extended over long periods of time, re- Concepts of Taxa. The species concept I ap- sulting in groups of lineages with inheritance ply is similar to that used in Antennaria by Bayer and variation patterns similar to those found and Stebbins (1983), but additionally incorpo- within polymorphic sexual species. rates elements of the phylogenetic species con- Subgeneric Classification. Three groups are cept as modified by Nixon and Wheeler (in distinguished from what has been called the press): A species is defined as a population or "Epiles" group by Hitchcock (1935). All three smallest set of populations that are coherent in are considered to be members of subg. Poa in biology, and that share a unique combination the broad sense (Soreng, in press). The sect. of fixed morphological characteristics, which Madropoa and Secundae and subsect. Epiles and consistently distinguish the component popu- Halophytae are described below in the "Taxon- lations from other sets of populations. I view omy" section. The following classification in- such characteristics as an indication of the ac- cludes all the known species within each tax- tual reproductive isolation of the species from onomic rank. other sets of biologically coherent populations. I. Sect. nov. Madropoa R. J. Soreng subsect. I have applied the rank of subspecies to sets of nov. Epiles: P. cusickii Vasey (including P. populations that are geographically distinctive, epilis Scribner), P. leibergii Vasey (includ- but distinguishable only by characteristics of a ing P. vaseyochloa Scribner), P. x netna- type that vary within populations across the tophylla Rydb. (pro sp.), P. porsildii whole species. A variety may occur sympatri- Gjaerevoll, P. pringlei Scribner, P. stebbin- cally or form distinct populations within the sii R. J. Soreng, sp. nov. Western North range of a subspecies as is common among America (figs. 1, 2). apomictic lineages. Two kinds of variation II. Sect. Abbreviatae Nannf. ex Tzvelev: P. occur: 1) Intergradient characteristics indicat- abbreviata R. Br., P. hartzii Gand., P. keckii ing incomplete segregation of incipient species R. J. Soreng, sp. nov., P. kolymensis Tzve- or broad secondary contact between former sis- lev, P. lettermanii Vasey (including P. ter species; and 2) intergradient characteris- montevansii Kelso), P. pattersonii Vasey, P. tics indicating contact between more distantly pseudoabbreviata Rosch. (including P. related species. The latter kind is more difficult br achy anther a Hulten), P. suksdorfii (Beal) to deal with. Populations with characteristics of Vasey ex Piper, P. vrangelica Tzvelev. Cir- this kind are treated as subspecific variants cumpolar, especially Beringia. within the most similar parent species, notho- III. Sect. nov. Secundae V. Marsh ex R. J. So- taxa, or species depending on their degree of reng subsect. nov. Halophytae V. Marsh distinctness, or stability of the new character ex R. J. Soreng: P. unilateralis Scribner in combinations. Because occasional hybrids occur Vasey, P. napensis Beetle. Western North between many species where apomixis is in- America. volved, I have formally recognized only those that occur over a substantial geographic range The above taxonomic arrangement treats the and are, therefore, likely to be encountered. 10 species of "Epiles" group (sensu Hitchcock Local hybrid populations of note are discussed 1935, emend. Hitchcock and Chase 1951) except under the species to which they key. I have for Poa strictiramea A. Hitchc. (=P. involuta A. attempted to recognize taxa as sets of biologi- Hitchc). That species, of arid mountains in and cally and morphologically coherent sexual or around the Chihuahuan Desert, shows only asexual populations, but I separate the sexual weak affinities to other "Epiles," and cannot be populations from the apomicts only when they satisfactorily placed in any North American can be morphologically distinguished. Facul- group at this time. The fact that a possible near- tatively apomictic clones or lineages (microspe- est relative of P. strictiramea, P. ruprechtii Peyr. cies of other authors) are considered to be in- of Mexico, has sheaths closed over more of their dividuals in an extended time frame (living 100s length, pubescent lemmas, is partially gyno- 1991] SORENG: POA 509 |1;° k d • |12Q e • 39- A 39- NV P J vs* •?v m y^\ ^ GA \ CA • \ ••*» } rf FIG.
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