Poasecunda J. Presl (Poaceae): a modern summary of infraspecific taxonomy, chromosome numbers, related species and infrageneric placement based on DNA.

Poasecunda J. Presl. s.l. is a morphologically highly variable bunchgrass that is a valuable forage species in western North America. There has been much controversy as to whether multiple taxa should be recognised and at what rank in this taxonomically challenging apomictic complex. Here we propose an infraspecific classification for Poasecunda of six varieties within two subspecies, juncifolia and secunda. New combinations are P.secunda vars. ampla, gracillima, juncifolia, nevadensis and scabrella. Conflicting plastid and nrDNA phylogenies show that P.sect.Secundae is of ancient hybrid origin. Based on this and its distinct morphology, the section is raised to the rank of subgenus. A key is presented for P.secunda infraspecies and closely related non-arctic species. Suppl. materials are provided of chromosome counts for Secundae taxa and D.D. Keck specimen annotations of taxa here included in P.secunda.

Introduction

J. Presl. s.l. is a morphologically highly variable species found primarily in western North America. It is common to dominant in grasslands extending from Alaska to Northern Mexico and eastwards in the northern Great Plains and scattered PageBreakmore eastern locations to the Gaspé Peninsula in Quebec (Marsh 1952, Kellogg 1985a, Soreng 2007). The species is disjunct but not common in South America, occurring in Patagonian Argentina and Chile (presumed origin of the type collection of ). This perennial bunchgrass is a valuable forage species that greens up and flowers in early spring and is abundant across western grasslands, coastal chaparral, Great Basin steppe, uplands of the Mojave Desert and dry forests surrounding these and reaches into alpine meadows. It exhibits diverse ecotypes across this wide geographic and climatic range. s.l. is abundantly represented in herbaria across the United States; for example, the United States National Herbarium (US) has more than three very full herbarium cases of specimens for the United States and Canada. It is distinguished from most species by its obscurely keeled lemmas, calluses often with a minute crown of hairs surrounding the base of the lemma (in subsp. ) and elongated spikelets (mostly 3.5–5 × longer than wide).

Differing taxonomies of the J. Presl complex continually appear in the literature. There are some 45 formal names applied to s.l. Some taxonomists follow A.S. Hitchcock’s (1935, 1951) taxonomic revision in the Manual of the Grasses of the United States, in recognising his two informal groups of species, “Scabrellae” and “Nevadenses”. In the former group, Hitchcock included (Thurb.) Benth. ex Vasey, Vasey, and (Scribn.) Howell; in the latter group Vasey ex Scribn., Scribn., Scribn. and Merr. Most of these taxa are still often recognised as species, subspecies or varieties (e.g. Keck 1950, 1959, Hitchcock et al. 1969, Dorn 1977, 1988). Keck (1950, 1959) additionally recognised Scribn., a serpentine endemic of the Sierra Nevada foot hills and Scribn. & T.A. Williams, a western sub-alpine/alpine element, within the Scabrellae group. Keck (1950, 1959) furthermore treated North American as Vasey, restricting to South America and Hitchcock et al. (1969) accepted this split. Arnow (1981) concluded was the correct name for plants of the form. Hitchcock (1935, 1951) included and in . Marsh (1950, 1952) and Kellogg (1985a, b) quite independently lumped most of these taxa in without any infraspecies (followed by Lesica 2012, for Montana). Kellogg’s work, based on morphometric analyses, separated ; Marsh maintained that and also . Soreng (1991b) divided s.l. into two subspecies, subsp. and subsp. (Scribn.) Soreng, corresponding to the taxa in the Scabrellae and Nevadenses groups, respectively and accepted and . Dorn (1988) accepted as full species the two subspecies that Soreng recognised and established a few varieties within those. Skinner (2010) followed the subspecies split, but provided separate pages for each of the common names that Hitchcock (1951) applied to species aligned within the two subspecies: Sandberg bluegrass (), Sandberg bluegrass [ (Scribn. & T.A. Williams) Beetle], Pine bluegrass (), Canby bluegrass (), Nevada bluegrass (), Big bluegrass () and Alkali bluegrass (). Several cultivars correspond to these different forms (Alderson and Sharp 1995). Thus, there appears to be a consistent desire and effort to maintain some or all of the diversity of forms commonly recognised, at some taxonomic rank.

Molecular (Patterson et al. 2005) and cytogenetic (Hiesey and Nobs 1982) studies show that s.l. is a highly complex, apomictic species. Apomixis, although it is facultative to varying degrees (Clausen 1961, Kellogg 1987), is apparently the main mode of reproduction in (Kellogg 1987). This has led to the production of various “strains”, ecotypes and races that are fairly monomorphic, but also to much intermediacy. Taxonomy in apomictic complexes is notoriously difficult, as is evident in the case of s.l. However, we concur with Marsh (1952) and Kellogg (1985a, b) that the rank of species is not justifiable for most of the Scabrellae and Nevadenses taxa.

s.l. belongs to , a primarily North American section of about eight species (Soreng 1991a, b, 1998, 2007, Soreng et al. 2003a, b, Gillespie et al. 2007). Soreng (1991a) established sect. with two subsections to accommodate species that share similar, apparently plesiomorphic traits, along with a derived chloroplast type (Soreng 1990) and suggested the section was of hybrid origin (Soreng 1990, 1991a, b). Our phylogenetic studies, based on plastid and nuclear ribosomal DNA, demonstrate that sect. is reticulate in origin (Gillespie et al. 2008, Cabi et al. 2017, Soreng et al. 2017). species exhibit several atavistic traits (Soreng 1991a, Soreng et al. 2015) that are otherwise odd in , including; a crown of hairs around the base of the callus, upper culm leaf sheaths margins often free more than 80–90% of their length and lemmas that are often somewhat rounded on the back. These traits are common in genera outside of and have led to many names for taxa being published in , and . species are tufted (rhizomes occur in some putative hybrids, see Species hybrids involving in the Taxonomy section) and perfect flowered, with anthers ranging from 1.2–3 mm long. Soreng (1991a, 2007) recognised two subsections: and V.L. Marsh ex Soreng. In addition to , s.l. and , also includes two Arctic species, A.E. Porsild and Gand. (Soreng 1991b) and is characterised by elongate, weakly compressed spikelets and lemmas rounded on the back. Species of subsect. (including Beetle, Trin., Scribn. ex Vasey) commonly have distinctly keeled lemmas and sometimes a papilliate epidermis on pedicels and leaf blades (Soreng 1991a, b, 2007).

Our goals in this paper are: 1) to provide a current overview of the taxonomy of s.l. and present an up-to-date infraspecific classification including new combinations for the forms often recognised as species and 2) to document what we know of the relationships and hybrid origin of s.l. and sect. . We also provide a review of the cytology of s.l. and other species in sect. .

s.l.: a revised infraspecific classification

In the interest of facilitating land-managers, ecologists, plant breeders, seed-storage facilities and collections managers in maintaining the understanding of variations in morphological forms in s.l. that are often recognised as species, we here PageBreakprovide varietal names within s.l. subspecies. This would also maintain herbarium collections that are organised or understood along A. S. Hitchcock’s taxonomic concepts, in which the taxa can be viewed to correspond to ecotypes or ecologically adapted apomictic lineages within . There is extensive intermediacy between these taxa and, thus, the rank of species is viewed as untenable (Marsh 1950, 1952, Kellogg 1985a, b, Soreng 1991b, 1993, 2007, Soreng et al. 2003a, b). RJS’s revised classification of s.l. is presented in the Taxonomy section below.

As a whole, s.l. is relatively easily split into two subspecies ( and , Fig. 1), but finer distinctions are often inconstant and overlapping. Within subspecies , it is a futile exercise to attempt to consistently separate , , and forms and so these are all included in var. . The epithet is often misapplied to plants RJS would classify as var. . Both and are extreme forms that many botanists seem to think are worthy of taxonomic recognition at some level. Kellogg (1985a, b) concluded that the form was the most distinct element within s.l., but the difficulty noted in the literature (see Hitchcock et al. 1969) of separating that from subspecies s.s., leads RJS to treat it as a variety in the subspecies. Range maps of the two subspecies are provided in Fig. 2.

Figure 1.

habit and panicles: A (Soreng 9359) B (Soreng 9358) C Panicles of subsp. (left) and var. (right) (Photos. RJS, Deschutes River near Madras, Jefferson Co., Oregon).

Figure 2.

Distribution of subspecies in North America: subsp. secunda (yellow) ; subsp. (blue).

An exhaustive summary of the nomenclature, protologues and types for the more than 45 names included in s.l. is beyond the scope of the present paper, but names of synonyms are listed. Hitchcock’s and some other historical synonymies for the nominal taxa can be found in his widely available "Manual of the Grasses of the United PageBreakStates" (Hitchcock 1951) and on-line: http://tropicos.org. See also the "Catalogue of New World Grasses" (Soreng et al. 2003a, and on-line Soreng et al. 2003b: http://tropicos.org/NameSearch.aspx?projectid=10) for synonymy for the two subspecies accepted.

A compilation of chromosome numbers for s.l. and other species in sect. (, , , and ) is presented in Suppl. material 1: Table S1. This shows that s.l. and all other taxa in sect. (for which chromosome numbers are known) are polyploid with a hexaploid base chromosome number of 2n = 42. There are many higher numbers, particularly in (2n = 42 to 105 or 106; some 140 counts). Subsp. has a mode at 2n = 84, while subsp. has a mode at 2n = 63 (2n = 42 to ca. 100) PageBreak(Soreng 1991b). Maintenance of a dysploid - nanaploid mode of ca. 2n = 63 in subsp. suggests this subspecies is highly apomictic.

Suppl. material 2 provides Keck’s annotations for each of the nominal taxa he accepted that are here included in . David Daniels Keck (1903–1995) had a deep interest in s.l. and other western North American species of and, in 1986, he kindly gave RJS his manuscript on those, as well as his extensive specimen annotation lists for of North America. He worked at the Carnegie Institute of Washington, at Stanford University, with J. Clausen and W. Hiesey for more than two decades (up to 1950), on the nature of species. He ceased his work on the lists by 1958 when he retired from the New York Botanical Garden. His lists focused on western Continental United States species, but included some mainly non-arctic Alaskan and Canadian and Mexican (Baja California) records, along with representative records of eastern United States species. Keck’s annotations represent hundreds of historical collections widely distributed in herbaria as vouchers for infraspecies. His annotations are considered to be sound by RJS, although here we recognise the taxa at the rank of variety. Copies of the full lists are stored in the reprint files in the Grass Lab in the Department of Botany, Smithsonian Institution. The lists are reproduced here in a semi-searchable form.

Phylogenetic relationships of s.l. and sect.

Methods

Phylogenetic analyses were performed on 78 samples (73 and five outgroups) (Suppl. material 3: Table S2.) using sequences of three plastid markers (matK, rpoB-trnC and trnT-trnL-trnF) and two nuclear ribosomal DNA (nrDNA) markers (ITS and ETS). Methods follow Soreng et al. (2017) and clade designations follow Soreng et al. (2010, 2017) (in the text, bold capital letters represent plastid types, bold small capital letters represent nrDNA types). Bayesian and maximum parsimony (MP) analyses were performed on the separate nuclear and plastid datasets. Branches having MP bootstrap support (BS) > 85% and Bayesian posterior probability (pp) support > 95% were considered strongly supported. See the following publications for genotype notation in : Soreng et al. (2010, 2017), Nosov et al. (2015), Cabi et al. (2016, 2017).

Results

Separate plastid and nuclear Bayesian trees are presented in Fig. 3 (summary statistics are given in Suppl. material 4: Table S3). In the plastid analysis members form a strongly supported subclade (pp = 1, BS = 97%) within the N clade (pp = 1, BS = 61%), whereas in the nuclear analysis members are intermixed with species of sects. Nannf. ex Tzvelev, Asch. & Graebn. and Dumort. PageBreakin the strongly supported s clade (pp = 1, BS = 99%). The postulated hybrids R. Br. × (Soreng 6055-2, 6107) resolved with (pp = 1, BS = 98%) in the P clade (pp = 1, BS = 100%) in the plastid tree and with in the s clade in the nuclear tree (pp = 0.99, BS = 64%) (Soreng 6055-2 not included since sequence unreadable due to multiple sequence copies).

Figure 3.

Bayesian 50% majority rule consensus trees of based on plastid (trnT-trnL-trnF, rpoB-trnC, MatK) data (left) and nrDNA ITS and ETS data (right). Bayesian posterior probabilities are shown above branches, maximum parsimony bootstrap values below branches. Outgroups are not shown. Major clades are indicated by colour and capital letters. Taxa shown in blue belong to ; those in grey are other taxa of putative hybrid origin that belong to different major clades in plastid and nrDNA trees.

Relationships amongst taxa within are mostly poorly supported and not congruent between plastid and nuclear trees. subspecies and form a clade and these form a clade with both subspecies in the nuclear tree. In contrast in the plastid tree, (Piper) D.D. Keck and are sister taxa, whereas conspecific subspecies are not together. formed a moderately supported subclade (pp = 0.98, BS = 0.75) within the s clade in the nuclear tree, but one of four samples (Gillespie 5725) resolved outside the N clade, amongst Vahl samples in the S clade, in the plastid tree. Rydb., currently considered a synonym of (Soreng et al. 2003b, 2007), resolved in but did not resolve with the latter species in either tree.

Discussion

The whole is shown to be of reticulate origin (Ns plastid / nrDNA genotype combination) consistent with our previous results based on fewer samples (Gillespie et al. 2008, Cabi et al. 2017, Soreng et al. 2017). All sampled taxa of the section have the N plastid genotype marker that is otherwise known only from Europe and SW Asia (sect. J. R. Edm. and unclassified species) and the snrDNA genotype of subg. (Dumort.) Soreng & L.J. Gillespie. While it remains a mystery how the N plastid type came to North America, it is evident that the snrDNA type came from (Ss genotype combination), either from sect. or, as we predict, from the primarily western North American sect. (where was historically placed; see notes in Soreng 1991b) or possibly from the morphologically related Russian Prob. (still lacking DNA data). Since N genotypes are otherwise absent from South America and S genotypes are otherwise rare there, we believe (, subsp. and ) arrived there secondarily by long distance dispersal from North America.

This ancient hybrid origin, together with its unusual morphology, make rather difficult to place within To better accommodate within the infrageneric classification of , we here raise the section to the rank of subgenus (see Taxonomy section below).

Curiously, All. and Boiss. & Heldr. ( Asch. & Graebn.) also have an Ns genotype combination. This species group is confined to alpine habitats in central and southern Europe and Anatolian Turkey. The taxa are rhizomatous, with extravaginal branching, have a strongly keeled lemma and a dorsal tuft of cobwebby hairs (web) on the callus lemma. Morphological characteristics lead us to hypothesise that this group of species is derived from a cross between Nn taxa of Europe and Ss taxa from , whereas we predict originated from a cross with Ss taxa. The N and s genotypes are slightly different between and , but if they PageBreakprove to have a common origin, sect. may be better placed within subgenus than with species they are usually associated with in (Roshev.) Olonova (type: Kom.), which exhibit Px genotypes (e.g. , Trautv. & C.A. Mey., Roshev.).

, an Arctic species distributed from Wrangel Island across the North American Arctic to Svalbard, represents an example in of more recent secondary reticulation. was shown previously (and again here) to include individuals with two different plastid types (N and S), interpreted as a case of recent and localised chloroplast introgression from (Gillespie et al. 1997, 2007, Gillespie and Boles 2001). The sample with S plastid type, clearly has a nrDNA type.

Vasey, a rhizomatous species of the Great Plains and eastern slopes of the Rocky Mts., was suggested by Soreng (2007) to have arisen from hybridisation of and some element of . The species appears to have the typical Ns genotype; it is shown here to have an N plastid genotype, whereas its nrDNA type is likely to be the s type as cloned nuclear DNA genes of track with (Patterson et al. 2005). So, the postulated origin of rhizomes in deriving from may be wrong, but deeper genetic analyses would be more convincing. Two collections, Soreng & Soreng 6055 and 6107, from Alaska, are viviparous plants identified by RJS as × . Both form a clade with (sect. ) in the plastid tree. In the nuclear tree, while the single sample included (6107, from the Alaska Range) resolves amongst taxa, it forms a clade with and these with , rather than . Based on geography, the paternal parent could not be (coastal Washington to California) and is unlikely to be (occurs in SE Alaska, but not in the Alaska Range); the only co-occurring is . Further research is needed, including deeper sampling in to determine the precise origin of this common viviparous Alaskan form.

Given that is an apomictic, polyploid (often with high and odd sets of seven and further dysploid chromosome numbers), hybridising complex of reticulate origin, it is not surprising that the detected phylogenetic structure within the group is mostly not well resolved nor supported. Neither the plastid nor nrDNA trees support the current division into two subsections, and . Several clades that are weakly to moderately supported may be informative of possible changes to the taxonomy. One example is the status of . This taxon was considered as a southern variant of , subsequently tentatively treated as a separate species in (Soreng et al. 2003a) and then most recently as a synonym of (Soreng et al. 2003b, Soreng 2007). Our data suggest that it may be a distinct species in subg. . Such taxonomic changes may be considered in the future, but we feel making changes within , based on the current phylogenetic trees, to be premature, especially given the small sample set within s.l. and the nature of the beast.

Taxonomy

(V.L. Marsh ex Soreng) Soreng & L.J. Gillespie, comb. et stat nov.

urn:lsid:ipni.org:names:77191590-1

Basionym.

V.L. Marsh ex Soreng, Syst. Bot. 16(3): 511, 523. 1991a.

Type species.

J. Presl.

Species included.

, , , , , (including ?), , .

Notes.

The was originally suggested to belong to (Soreng 1991a), but, in subdividing the genus into more subgenera based on our plastid phylogeny (Gillespie et al. 2007), the section was moved to subg. (Dumort.) Soreng & L.J. Gillespie. We subsequently discovered that the set of plastid genotypes resolved in a clade with three European diploid species with Nn genotypes, but the nrDNA genotypes of were like those of subg. (Ss) (Cabi et al. 2017, Soreng et al. 2017 and results presented above). The apparently ancient reticulate origin of the group, along with its odd morphology within , now leads us to raise the section to the rank of subgenus.

J. Presl, Reliq. Haenk. 1(4–5): 271, 1830

Range.

CANADA: Alberta, British Columbia, Manitoba (sw), Northwest Territories (sw), Ontario (Manitoulin Island), Quebec (Gaspe Peninsula), Saskatchewan, Yukon (s). UNITED STATES: Alaska (se interior border), Arizona (n), California, Colorado, Idaho, Montana, North Dakota, Nevada, New Mexico (n), Oregon, South Dakota (w), Utah, Washington, Wyoming, with outlying populations in Nebraska (w), Oklahoma (panhandle), Michigan (Isle Royal) and sporadic in Illinois and Maine, Massachusetts.

The geographic range of in North America north of Mexico is mapped in Fig. 2. Subspecies occurs throughout the species range and also reaches into Baja California Norte, Mexico (Fig. 2A). The range of subspecies is almost completely within that of subsp. , except for several scattered localities in Arizona, New Mexico and Nebraska; it is absent from Mexico and mostly absent from the Pacific Coast and coastal mountains (Fig. 2B). The ranges of varieties are broadly overlapping within their subspecies. Both subspecies have disjunct populations in the Patagonian Andes of Argentina and Chile.

(Scribn.) Soreng, Phytologia 71(5): 401. 1991b [1992]

Scribn., Bull. Div. Agrostol., U.S.D.A. 11: 52, pl. 8, 1898.

(Merr.) Soreng, comb. et stat nov.

urn:lsid:ipni.org:names:77191591-1

Figs 1B, C , 3A

Merr., Rhodora 4(43): 145, 1902.

Synonyms.

Merr., Rydb., (Merr.) Dorn, T.A. Williams, Rydb.

Habitat and range.

Open upland forests, mountain steppe, generally in light, well-drained soils to somewhat heavy soils. Range of the subspecies (Fig. 2B), but mostly absent from west side of the Sierra Nevada and westwards. Minor outlying occurrences in Arizona and New Mexico of subsp. (Fig. 2B) mostly represent var. (Soreng 1985), likely introduced by seeding.

Chromosome numbers.

Numbers reported as : 2n = 61, 62(x3), 63 (x11), ≈ 63 (x3), 63–64 (x2), 64 (x6), ≈ 65, 70–71, ≈ 97, ≈ 100.

(Scribn.) Soreng, comb. et stat nov.

urn:lsid:ipni.org:names:77191594-1

Fig. 4B.

Figure 4.

infraspecies illustrations (reproduced from Hitchcock 1935): A panicle and floret B panicle and floret C panicle and floret D habit, panicle and floret E habit, panicles, spikelet and floret F, G habit, panicle and floret variations A, B, C = subsp. D, E, F, G = subsp. . Scale bar: 5 mm for florets, 10 mm for spikelet, 5 cm for habits and panicles (10 cm for E habit and panicles).

Scribn., Bull. Div. Agrostol., U.S.D.A. 11: 52, pl. 8, 1898.

Piper, (Scribn.) M.E. Jones, Scribn., , D.D. Keck (?), (Scribn.) Beetle.

The type and other material identified as by Keck combine pubescent lemmas with narrow panicles, firm blades and short ligules on lateral shoots and appear to RJS to be intermediate in form between varieties and ( form).

Pine forests and steppe, riparian and alkali meadows, in well-drained to poorly-drained, light to heavy, often alkaline or saline soils. Range of the subspecies (Fig. 2B), but infrequent in the California Floristic Province and south-western states.

Numbers reported as : 2n = 42, 60, 62, 63, 63–64, 78, 84. The one 2n = 42 count was originally published by Hartung (1946) as , but RJS re-determined the California voucher as .

(Vasey ex Scribn.) Soreng, comb. et stat nov.

urn:lsid:ipni.org:names:77191595-1

Fig. 4C

Vasey ex Scribn., Bull. Torrey Bot. Club 10: 66, 1883.

(Vasey ex Scribn.) Beal, Thurb., Kuntze, Vasey ex Scribn., (Thurb.) Benth. ex Vasey, Vasey ex Scribn. (nom. inval.), (Kuntze) Vasey, (Vasey ex Scribn.) Ponert.

Open forests and steppe, in light, well-drained to heavier soils. Range of the subspecies (Fig. 2B), but mostly absent from west side of the Sierra Nevada and westward. Collected once in New Mexico (Soreng 1985).

Numbers reported as : 2n = 62 (x2), 62–63, 63 (x7), 64, 64–66 (x2), ≈ 65, 70.

(Vasey) Soreng, comb. et stat nov.

urn:lsid:ipni.org:names:77191596-1

Fig. 4D

Basionym.

Vasey, Contr. U.S. Natl. Herb. 1(8): 272, 1893.

Synonyms.

, Vasey, Scribn. & T.A. Williams.

Habitat and range.

Open forests, moist cliffs and rocks and subalpine to alpine meadows, in well-drained acid soils that are consistently moist through the growing season of the variety. Pacific Northwest and northern California.

Chromosome numbers.

Numbers reported as : 2n = 81, ≈ 81, ≈ 84, 86, ≈ 91.

(Thurb.) Soreng, comb. et stat nov.

urn:lsid:ipni.org:names:77191598-1

Fig. 4E

Thurb., Bot. California 2: 310–311, 1880. (Thurb.) Benth. ex Vasey, Grass. U.S. 42, 1883.

Munro ex Thurb., Thurb., (Thurb.) Kuntze, Scribn., (Munro ex Thurb.) Scribn., Scribn., Scribn., Vasey, (Thurb.) Benth ex Vasey, (Thurb.) Ponert, Munro ex Benth.

Open pine forests, coastal scrub and coastal and Central Valley grasslands, in well-drained or heavier soils. Mainly in the California Floristic Province, but extending northwards in the Pacific North West and southeast into the Mojave Desert, where it is largely replaced by var. .

Numbers reported as : 2n = 44+f, 61–63, ≈ 62, 63 (x4), 64, ≈ 66, ≈ 68, 81 (x2), 82 (x3), 84 (x10, ≈ 84, 84+f, 86, ≈ 86 (x2), ≈ 88, ≈ 91, 104.

Figs 1A, C , 4F, G

Pursh, Spreng. ex B.D. Jacks., (Pursh) Roem. & Schult., (Scribn.) Beal, Beal, Beal, Thurb., Vasey ex Beal ( form), Vasey, Phil., Phil., Scribn., Kuntze, Nutt. ex S. Watson, Nash, , (Vasey) M.E. Jones, (Vasey) M.E. Jones, (Vasey ex Beal) M.E. Jones, (Scribn.) Piper, Trin., (Scribn. & T.A. Williams) Hack., Rydb., Scribn. & T. A. Williams, Scribn., Vasey, Scribn., Vasey, (Scribn.) M.E. Jones, (Scribn.) M.E. Jones, Vasey, Scribn. & T.A. Williams (toward form), (Vasey) Dorn (= form), (Scribn. & T.A. Williams) Beetle, (Vasey ex Beal) Beetle, (Vasey) B. Boivin, Buckley, Nutt. ex S. Watson, Vasey, (Vasey) Vasey, Scribn., (Scribn.) Ponert, (Beal) Ponert

Notes.

There are various variety names included in the autonymic var. . Variety is s.s.; vars. , , and represent the form; and vars. , , and represent the form. Varieties and are generally subalpine to alpine with slightly open panicles; Hitchcock (1951) included them within , but Keck (1959 and annotations) treated them as . Vars. and are intermediate to var. , but are too smooth to be included in the latter variety. Hitchcock (1951) equated with var. , but we agree with Keck (Suppl. material 2) in equating it with the form. Nutt. ex S. Watson (1871), nom. illeg., was described from heterogeneous original material (at US, distributed as S. Watson 1318): E. Humboldt Mts. = var. ; Virginia Mts. = var. ; Diamond Mts. = var. . However, Watson also cited “ Nutt., ms. In Herb.” The Nuttall specimen with that name is at PH (PH00020091) and Buckley (1862) is based on that specimen. Therefore, Watson’s taxon name is superfluous.

Open forests, steppe, and alpine, generally in light, well-drained soils. Range of the subspecies (Fig. 2B), but mostly replaced by var. on west side of the Sierra Nevada and westwards and in the Mojave Desert.

Numbers reported as and avowed synonyms: 2n = 42, 56, ≈ 68, 70 (x2), ≈ 70 (x2), ≈ 74, ≈ 78 (x2), 81, 81, 82 (x2), 84 (x2), 84–88+II, ≈ 84, 85–87, 86 (x2), ≈ 87, ≈ 99, 104. The one count of 2n = 42 was reported by Bowden (1961) originated from Alberta. Numbers originally reported as : 2n = 56, 70 (x4), ≈ 72, ≈ 82 (x2), ≈ 83, 84 (x4), ≈ 84 (x2), 85, ≈ 86. Keck (in Munz 1959) attributed higher numbers to ). Numbers secondarily reported as : 2n = 90, 93, 94, 99, 105–106.

1	Lemmas more or less crisp-puberulent on the lower half or basal portion (sometimes obscurely so in P.secundavar.scabrella); ligules of lower culm usually well developed and acute to acuminate (short in P.tenerrima); tillers strictly intravaginal, cataphylls absent, prophylls well developed, mostly over 1 cm long; leaf blades commonly withering early, long-cells all or mostly fusiform and smooth-walled	2
–	Lemmas glabrous, smooth or scabrous (except in “P.juncifoliasubsp.porteri” form, but then plants from the plains of the eastern slope of the Rocky Mts.); ligules of lower culm and lateral shoot leaves truncate to rounded (acute in var. nevadensis); tillers intravaginal and sometimes extravaginal, the latter with cataphylls and reduced prophylls (mostly less than 2 mm long); leaf blades more or less persisting in form, long-cells mostly rectangular and sinuous-walled	6
2	Leaf blades short (mostly 1–3 cm long), (1–) 1.5–3 mm wide, flat, with prominent white, cartilaginous margins; plants of serpentine rocks in the Wenatchee Range of the Cascade Mts., Washington State	P. curtifolia
–	Leaf blades of various lengths and widths, but not short and flat, without prominent cartilaginous margins	3
3	Sheaths scabrous, at least on the margins; ligules scabrous; panicle branches scabrous, often densely so; plants mainly of California Floristic Province and Mojave Desert	4
–	Sheaths smooth; ligules smooth or lightly scabrous; panicle branches smooth or scabrous; plants mostly of the eastern slope of the western Cordilleras and eastward	5
4	Ligules of culm leaves well developed (2–6 mm long), acute to acuminate; blades filiform or broader; panicles branches capillary or thicker, appressed to ascending (rarely spreading); plants widespread; chromosomes 2n = 63 and higher	P. secunda subsp. secunda var. scabrella
–	Ligules of culm leaves short [0.5–1.5 (–2.5) mm long], truncate to obtuse (acute); blades filiform; panicle branches capillary, widely spreading; plants of serpentine barrens in central foothills of west slope of the Sierra Nevada; chromosomes 2n = 42	P. tenerrima
5	Panicles open, the branches spreading to patent, divergent more than 45° at anthesis and remaining open with spikelets absent in the lower half; plants of moist often shady places	P. secunda subsp. secunda var. gracillima
–	Panicles usually loosely to tightly contracted at maturity, branches sometimes ascending but branches finally divergent by less than 45°, spikelets from near the base or lower 1/3rd; plants mostly of more open places	P. secunda subsp. secunda var. secunda
6	Sheaths scabridulous; ligules elongated, acute, decurrent	P. secunda subsp. juncifolia var. nevadensis
–	Sheaths smooth; ligules of lower culm and basal leaf short, obtuse to truncate, not decurrent	7
7	Blades involute; plants of open riparian and alkali or saline meadows	P. secunda subsp. juncifolia var. juncifolia
–	Blades flat; plants of mountain meadows and forests	P. secunda subsp. juncifolia var. ampla