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TAXONOMIC IDENTITY AND HISTORICAL ACCOUNTS OF DALEA CYLINDRICEPS (FABACEAE), A SPECIES OF CONSERVATION CONCERN IN THE GREAT PLAINS (U.S.A.) PDF

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TAXONOMIC AND IDENTITY HISTORICAL ACCOUNTS OF DALEA CYLINDRICEPS (FABACEAE), A SPECIES OF CONSERVATION CONCERN THE GREAT IN PLAINS (U.S.A.) James Locklear H. Omaha, Nebraska 68 08, U.S.A. 1 INTRODUCTION the western Great Plains of Daka herb 0 native to (Figs. cylindriceps Barneby (Fabaceae) 1 ] i: New Mexico, Okla- Kansas, Nebraska, distribution, D. cylindriceps extent of historical homa, South and Wyoming. Despite the large its Dakota, Texas, and As noted in the recent Flora of Ne- scattered local. has been and occurrences are collected infrequently throughout wide geographic range.” Dalea almost its rare braska (Kaul “This distinctive species is et 2011), al. which has been docu- concern in but two of the states in it all conservation cylindriceps tracked as a species of is G3G4 (vulnerable). mented, and ranked by NatureServe (2013) as is indicates this species range of D. cylindriceps northern part of the survey by the author in the Initial field searched the sites of 22 historical occur- In 2010, may population decline. 1 have experienced considerable Wyo- Co Nebraska, and orado, specimen locality data) in herbarium from rences of D. (derived cylindriceps be noted that D. cylindriceps is a lelatively large should 4 ming and found populations at only sites. It existing cm making flowering spikes, 18 long) dm elongate (up to with distinctively plant 3-6 (generally [up to 8] tall) it a noticeable object in the landscape. >sment by yielding historical in- conserv, informs Analysis of and ecological literature floristic In the case ot associations, etc. j formation on abundance, ecological distribution, C an cm taxon. of this history nomenclatural made complex sis by is difficult the conservation sta of assessment its more accurate for this species will allow for i-Bot - Res. Inst. Texas 7(2): 879 890. 2013 Locklear, Dalea cyl indriceps, a species of conservation c Fig. 2. Dalea cylindriceps, Sheridan County, Nebraska. Photo James Locklear. Bameby monograph on the genu The taxonomic was reviewed by (1977) in his history of this species n currendy accepted L., in which he determined that a new binomial was required and proposed the Bameby. cylindriceps 1827 from ribed in The name taxon Petalostemum macrostachyum Torr., earliest published for this is < America North Tor- material collected in 1820 by Edwinjames in present-day Nebraska (see below). In Flora of , binomial name (giving the full Petalostemon rey and Gray (1838, 309) altered the rendering of the generic to p. synonym. Gray later (1848) dis- omatum Douglas ex Hook, as a as and Petalostemon macrostachyum) listed P. omata now recognized as Dalea (Douglas) tinguished P omatum from macrostachyum, and the former is P. & This ear and northern Great Basin. Eaton Wright, a species of the Columbia Plateau plant of the Pacific P 19th century botanical literature a resulted in macrostachyum being treated in certain Northwest Coulter 1885). as well as the Great Plains (e.g.. P synonym under macrostachyum compacta Spreng. as a added Dalea Torrey and Gray 690) later (1840, p. ac- and .Texas occurs in eastern 097^0 species .hat ta, Bameby eompochr a separate as determined by is dong wkh misappUcatkm of (he speoffcepi- ampacta, of D. cent Arkansas and Oklahoma. Misinterprecation (Peterson 2000). surrounding D. cylindriceps confusion much nomenclatural thet, has been at the root of of the macrostachyum (cor Torrey’s Petalostemum Great Plains region (East- the botanical masculine appears in the early lit. gender of Petalostemon) & but subsided after pubhca- Coulter 1874), Porter wood Parry 1870; Engelmann Gray 1848, 1863; 1893; 1862; Journal of the Botanical Research Institute of Texas 7(2) (Doane College Natural compactus (Spreng.) Swezey in 1891 in Nebraska Flowering Plants tion of Petalostemon Doane Goodwin College in pamphlet written by D. Swezey, professor at Crete, History Studies No. a 17-page 1), synonym While proposed Nebraska. Swezey cited macrostachyus Torr.” as a replaced of his P. compactus. in “P. name apparently gained recognition and adoption thanks to the re- obscure publication, Swezey’s a relatively view pamphlet by Nathaniel Lord Britton (1891) published in the widely-circulated Bulletin of the Torrey of his Botanical Club. which Torrey and was new combination based on D. compacta (Sprengel 1826), compactus Swezey’s a P. macrostachyum (Torrey 1827) but which Swezey Gray 690) included in the concept of Petalostemon (1840, p. P on should be noted that Swezey applied the name compactus to specimens elevated based prior publication. It known One herbarium Do- Two in the of what such specimens are to the author. is of in fact D. cylindriceps. is Garden on 1890 near Lewellen in ane College in Crete, Nebraska, collected by E.E. Sprague (s.n.) 25 July NY and The County, Nebraska (annotated by Rolfsmeier [1987] as D. cylindriceps). other is at (1259224) ap- S. under an same and distributed pears be a duplicate of the Sprague collection (with collection date locality) to Wemple andD. compactum”). by [1961-1965] as “Herb. ofG.D. Swezey” label (annotated D. lsely [1958] “P. was Michx. and Dalea Further clouding the nomenclature of this taxon the reduction of Petalostemon L. Kuntze, under to the genus Kuhnistera Lam. by Kuntze (1891), in which he proposed K. compacta (Spreng.) “ added synonyms. which he placed D. compacta and Petalostemon macrostachyum” as Heller later (1896) “Pet- name what ostemon compactus Swezey” to the of synonyms under K. compacta. Use of the K. compacta for al list occurred in a number of botanical and ecological publications in Great Plains states clearly D. cylindriceps is & (Hitchcock 1896; Pound Clements 1900; Saunders 1899). The publications of Per Axel Rydberg were foundational to state and regional treatments of the flora of the Great Plains in the part of the 20th century. Regarding D. cylindriceps, Rydberg initially (1894, 1895) took first up K. compacta for the taxon, citing Petalostemon macrostachyus and D. compacta as synonyms, but subse- quently adopted Petalostemon compactus (Spreng.) Swezey, first in Flora of Colorado (1906), then in Flora of the Rocky Mountains and Adjacent Plains (1917) and Flora of the Prairies and Plains of Central North America (1932). Brown adopted During same period, Britton and (1897) treated this taxon as K. compacta but later (1913) this “ compactum Swezey’s name, rendering Petalostemum (Spreng.) Swezey.” it Subsequent treatments of this taxon in the flora of the Great Plains region followed Rydberg in using works Swezey’s Petalostemon compactus (often rendered P. compactum). These include floristic and ecological for the states of Colorado (Harrington 1954; Ramaley 1939), Kansas (Bare 1979; Barkley 1968; Gates 1940), & & New Nebraska Webber Mexico (Wooton Standley Martin Hutchins 1980), (Petersen 1923; 1892), 1915; & Wyoming Texas (Correll Johnston 1970; Turner 1959), and (Dorn 1977b), as well as other regional literature & (Barr 1983; Coulter Nelson 1909; Dorn 1977a; Rogers 1953) including Atlas of the Flora of the Great Plains (GPFA 1977). Swezey The taxonomic identity and nomenclatural problems associated with compactus (Spreng.) re- P. Wemple and ceived further consideration clarification in the 1970s. In his revision of the genus Petalstemon, “ (1970) recognized Petalostemon compactum (Spreng.) Swezey,” treating both P macrostachyum and D. com- monograph on from the pacta as synonyms. But, in his the genus Dalea, Barneby (1977) separated D. compacta P taxon originally described as macrostachyum, which excluded the type of D. compacta from his concept of P. macrostachyum, making the latter the oldest available name for the species. Since Barneby was reducing the mac- genus Petalostemon to Dalea, he needed to transfer P. macrostachyum to Dalea, but there was already a D. “unwelcome new was adopted in rostachya Moric., necessitating the epithet” of Dalea cylindriceps. cylindriceps Flora of the Great Plains (GPFA 1986) and has since been used in most treatments of the flora of the region & nomen- (Dorn 2001; Kaul et 2011; Turner et 2003; Van Bruggen 1985; Weber Wittmann 2012). Correct al. al. and synonymy clature for this species is provided below. New Dalea cylindriceps Barneby, Mem. York Bot. Gard. 27:227. 1977. Expedition. Dr. Type: United States: “Long's 1st South Platte River in Lincoln County, Nebraska, 22-23 Jun 1820. 883 Discovery and Type Locality The type specimen of what would eventually be recognized as D. cylindriceps was collected by Edwin James while traveling with the Stephen H. Long Expedition of 1820. In his enumeration of the botany of the expedi- tion, Torrey (1827) stated the locality ofJames’ collection of Petalostemum macrostachyum as “About the forks of NY No known the Platte.” date or locality information is provided on the holotype (the only specimen) at (Fig. only the statement, “Long’s Expedition. Dr. James.” James did not mention this species in either his pub- 1st 3), & (Goodman Lawson lished account of the expedition (James 1823) or in his personal diary 1995). The North and South Platte rivers (“the forks of the Platte”) join to form the Platte River in Lincoln Coun- From Nebraska, just east of the city of North Platte. here westward to Ogallala, Nebraska (a distance of ca. ty, 70 km), the main channels of the North and South Platte rivers run within a few (5-6) kilometers of each other km until they begin to diverge about 18 east of Ogallala, the North Platte to the northwest and the South Platte between North to the southwest. Thus, “About the forks of the Platte” could be a general description of the area & (Goodman Lawson Platte and Ogallala, which the expedition traversed 22-25 June 1820 1995). North But this phrase appears to refer to a more limited geographic area. After reaching the junction of the North and South on 22 June 1820, the expedition continued west along the north side of the Platte Platte Platte heading southwest, making camp on the north side of the South River a few miles then crossed the river for Long Goodman and Lawson Platte River. In their reconstruction of the route and itinerary of the Expedition, km) North camp 22 6-7 miles (10-12 west of the city of Platte, place the location of the ofJune at (1995, 15) p. community of Hershey, Nebraska. which would bVjust to the east of the present-day km) On the morning of June 23, the party traveled about two miles (3.3 upstream then crossed to the The would have been in the vicinity of Hershey. James (1823, p. 468) south South crossing side of the Platte. when our attention was arrested by the beautiful states that the party “had no sooner crossed the [South] Platte & N A. Gray). In a footnote to his discussion Torr. (« O. coronopifolia white primrose (Oenothera pinnatijida. .)” were collected “about the forks of the Platte.” other species that of the Oenothera, James several lists byJames as “about the forks of the Platte” is tied to the locality of Thus, appears that the area referred to it on The Nebraska 23 June 1820. type of Lincoln County, present-day South River in this crossing of the Platte and towns Hershey Sutherland somewhere between the of was along the river D. cylindriceps likely collected & (Goodman Lawson camp ofJune 23 km of the expedition’s being the vicinity (ca. 12 to the west), the latter made on 05 August 1989 (D. Suther- supported by a collection of D. cylindriceps 1995, p 16) This conclusion is south of Hershey. South Riverjust NEB, NY) along the Platte land 6802 with Rolfsmeier at S. ...... Accounts Historical . and ecological htera- the early flonstic history of D. cylindriceps, Armed nomenclatural with knowledge of the summary specie* The following is a of notable references to this was searched for tore of the Great Plains ^ ndl ”^ aooss the cc traveling many botanical collectors by of the early was encountered Aug^ Ihllea cylindriceps m Fendfa 1846 (Toney 18CT 1820 Edwin James in MOs. The includes Great Plains in the list and Harbour (with Charles C. and Elihu Hall (Engehnann 1862), J.P. Hayden 1857 (Gray Ferdinand in 1848), was typically ^^fretoatandance cylindriceps D. indicated, is & Ramaley Rogets 1953) Tins ts tn spue of fact that 1939; Clements 1900; Pound ized, etc (Hitchcock 1896; comments Pound and of the when as reflected in present, landscape the the object in species a noticeable is ccmpacm) K. as described D. cylindriceps (as where they Nebraska, Clements The Phytageogmphy of (1900) in : 885 most “the remarkable species of this genus in our stems, which flora. Its are densely leafy, are often a meter high and are very conspicuous on account of the long decimeter) nodding (a spikes of white or yellowish flowers .” may Dalea cylindriceps have been more abundant prior to settlement and agricultural development within “ Eastwood its range. (1893) included Petalostemon macrostachyus Torr” in A Popular Bora Denver, Colorado of , book aimed her at “students” and “beginners” and therefore focused on the showier or more common elements Eastwood of the flora. stated the species occurred in “North Denver.” There are a number of D. cylindriceps herbarium specimens collected in the late 1800s from what is today the Denver metropolitan area northward toward Fort but Collins, the native vegetation of this area has been largely displaced by suburban and ex-urban development. The following references provide early accounts of the ecological associations of D. cylindriceps Colorado: “Sand (Ramaley Prairie” 1939, p. 14 as P. compactus). In his ecological study of the sandhill vegeta- tion of northeastern Colorado, Ramaley recognized four plant communities occurring upland sandy in soils (Loose Sand and Blow-out; Sand-hills-Mixed; Sand-Sage; Sand he P compactus only Prairie); listed under Sand Prairie, where he placed it on the list of “Other species,” indicating relative infrequency. P Kansas: “Prairie” (Gray 1848, 33 as macrostachyum Gray enumerated plants collected by Augustus p. ). New Fendler in the vicinity of Santa Fe, Mexico, which he reached in 1846 after traveling from Louis St. along the Santa Fe Trail. Gray reported the locality of the collection as “18 miles west of Lower Springs, Wagon Cimarron [River].” Lower Spring (also called Bed Spring) was a site along the Cimarron Cutoff of A the Santa Fe Trail located on the Cimarron River in what is today Grant County, Kansas. location along the 18 miles west of Lower Spring would be in present-day Morton County, Kansas in the vicinity of trail number the Cimarron National Grasslands, where a of recent collections of D. cylindriceps have been Kansas: “Ulysses, Grant County... on sandy knolls along the South Fork of the Cimarron [River]. Rare.” Thompson on by (Hitchcock 1896, 543 as K. compacta Hitchcock reported the plants collected C.H. p. ). in southwestern Kansas in 1893. Nebraska: “Sand along the Loup fork and Niobrara [rivers]” (Engelmann 1862, p. 189 as P. macrostachy- hills ium Engelmann enumerated plants collected by Ferdinand Hayden in the Elkhom, Loup, Platte, [sic]). and Niobrara river valleys of present-day Nebraska during an expedition led by Lieutenant G.K. Warren of the U.S. Topographical Engineers in 1857. & county (Pound Clements 1900, 250 as K. Nebraska: “Localized in the sand-hills of Scotts Bluff (sic)” p. on herbarium material Pound and Clements based their account of this species col- compacta). likely NY NEB on 04 August 1259216, 1259217) 1891. by PA. Rydberg Rydberg 177085, 177088; lected 61; ( number “Petalostemon macrostachyus Torr.” in his journal (manuscript at Rydberg identified his 61 as NEB), noting the locality as “On the sandhills north of the Platte River, Scotts Bluff Co.” Kuhn Kaul Such early references, as well as those found in more recent literature (Hazlett 2004; et al. 2011; et & & Sutherland Rolfsmeier 1989), indi- Steinauer 2010; Rolfsmeier al. 2011; Neid et 2007; Reif et al. 2009; al. common sand and sand dunes are Eolian sheets with sandy habitat. cate a strong association of D. cylindriceps New Oklahoma, Mexico, and Nebraska, Colorado, Kansas, notably in landforms western Great Plains, in the by & dune are mostly stabilized vegetation. Texas (Muhs Holliday 1995). At present, these fields herbarium with examination of speci- coupled and ecological literature Review of Great Plains floristic com- with plant frequently associated mens indicate that D. cylindriceps is and survey, the author’s initial field Occur- a dominant or distinctive element. (Artemisia is munities which shrub sand sagebrush filifolia) in the and southwest Kansas are associated Colorado Nebraska, eastern (Fig. 4), southwest rences of D. cylindriceps in with interspersed tailor moderately dense layer oiA.filifolia with communities comprised ofasparse to steppe sod with geography, precipitation, texture, etc. varying component grass species mid-height the grasses, & Frequently Rolfsmeier Steinauer 2010). Ramaley 1939; Neid 2007; (Kuchler 1974 Lauver 1999; et al. et al communities occur on sands and loamy sands as- sometimes extensive these referred to as “sandsage prairie,” sociated with dune systems. eolian journal of the Botanical Research Institute of Texas 7(2) 886 Dalea cylindriceps has been collected (to a lesser exte deposits of streams and colluvium derived from sandstone outcrops and escarpments il jus plant communities that develop in such habitat occur as small patches or bands < The association of D. cylindriceps with sandsage prairie could be a factor in the apparent decline of this species. Throughout the Great Plains, extensive tracts of sandsage prairie have either been converted to irri- Sexson 1983). Future Research Kansas Ronald McGregor University of botanist L. (1986) asserted nearly 30 years ago that D. cylindriceps “needs more careful study ” and his statement holds true yet today. Additional research needed to more fully is assess the conservation status of D. cylindriceps. Survey of historical occurrences throughout the range of the species is needed to determine the number of existing occurrences and to develop a more detailed ecological As a profile that includes habitat requirements, edaphic factors, disturbance factors, associated species, etc. more precise understanding of the ecology of D. cylindriceps gained, workers can be more strategic in is field arch for additional occurrences. needed Research into the life history traits (phenology, reproductive ecology, etc.) of D. cylindriceps is t< how nine these shape demography and population trends. This species appears to be a short-lived or evei 887 monocarpic herbaceous (Bameby perennial 1977; Kaul et al. 2011). McGregor (1986) noted that on dunes and areas of loose sand, D. cylindriceps “sometimes flowers the first year and frequently expires at the end of the second or third season,” yet is often a longer-lived perennial in more stable sandy areas. Many plant species associated with sandy habitat are specialized to a particular ecological niche or stage of recovery related to natural disturbance, and research needed determine such is to if is the case for D. cylin- Sandsage dynamic driceps. prairie is a naturally plant community, with species composition, patterns of vege- tation, and percent canopy cover changing over time in response to fluctuations in precipitation (Collins et al 1987; Farrar 1993b; Kelso et al. 2007; Ramaley 1939; Rondeau 2003, 2013). Such recurring natural disturbance could result in fluctuations in the presence and abundance of D. cylindriceps over time. Monitoring studies of existing populations of D. cylindriceps are needed to answer these and other ques- tions of history, demography, and population trends. Further study of D. cylindriceps would be of value not life only for the conservation and management of this species, but also holds promise for a better understanding of the ecology and dynamics of sandsage prairie, a plant community that is of conservation concern throughout most of its distribution in the western Great Plains (NatureServe 2013). Given its close association, D. cylindri- ceps could serve as an indicator species of high quality occurrences of sandsage prairie and of habitat integrity and health. A Common Name Bameby Dalea for cylindriceps and The misleading common name “Andean prairie-clover” has been applied to D. cylindriceps is currently in use by NatureServe and other databases listing plants of conservation concern. Dalea cylindriceps is not (2013) and common name may have had origin in the binomial D. macrostachya Moric., a plant of the Andes, this its now which macrostachyum Tore, the validly published name for what is recognized as D. is similar to P. first & Dalea macrostachya a replaced synonym for D. coerulea (L.f.) Shinz Thell. var. longispicata cylindriceps. is Bameby, which occurs in the Andean region of South America (Bameby 1977). (Ulbr.) common names have been adopted for D. cylindriceps that point to the dense, elongate spike- Several other These include “dense-flowered morphological feature of this species. which like inflorescence a distinctive is (GPFA and & “massive spike prairie-clover” 1986), Brown Saunders 1899), 1897; prairie clover” (Britton & Conard The two names reflect Bar- Stubbendieck 1989). latter (Kaul 2011; “large-spike prairie-clover” et al. massive spike.” neby’s (1977) etymology of cylindriceps: “of the proposed here based on the close association of D. cylin- The common name “sandsage prairie-clover” is communities in which sand sagebrush (A. filifolia) is a domi- driceps with sandsage prairie and similar plant nant or element. distinctive acknowledgments David Sutherland, profes- who provided assistance with this research. would individuals I like to think several genus forthcoming treatment of the Omaha and author of the Nebraska sor emeritus the University of at at my information pertinent and brought to attention manuscript reviewed the Dalea for Flora North America, of ami professor en.ert.us, respec- professor Rob Wikel, associate and NEB. Brad Elder in the Rydberg diary at Goodwm Swezey on publications of D. die information pnrvkfed Doane Nebraska), tively, College (Crete, at and research professor with the Kaul, curator Robert Doane College. and access to his botanical collections at ^1891 photocopy of provide Nebraska, Unhersity of Charles Bessey Hetbarium (NEB) of the E. «SOfl Hams Herbanum rihgh. of the director Rolfsmeier, i Steven pamphlet, Nebraska Flowering Plants. and on die ecology of D. cylmdnccps also observations shared his Nebraska), Chadron (Chadron, State College reviewed the manuscript. Lawrence, Kansas. of Kansas, s Press . . Journal of the Botanical Research Institute of Texas 7(2) Minnesota Minneapolis, Minnesota. Barr, C.A. 1983. Jewels of the Plains. University of Press, Mem. New -891 977. Daleae imagines. York Bot. Gard. 27:1 Barneby, R.C. 1 Club18:195-196. 1891. [Review of] Nebraska flowering plants. Bull.Torrey Bot. Britton, N.L. Britton, N.L. and A. Brown. 1897. An illustrated flora of the northern United States, Canada, and the British possessions, New New Charles Scriber's Sons, York, York. vol. 2. Britton, N.L and A. Brown. 1913. An illustrated flora of the northern United States, Canada, and the British possessions, New New 2nd ed, vol. 2. Charles Scriber's Sons, York, York. Collins, S.L., JA Bradford, and P.L Sims. 1987. Succession and fluctuation in Artemisia dominated grassland. Vegetatio 73:89-99. Correll, D.C. AND M.C. Johnston. 1970. Manual of the vascular plants ofTexas. Texas Research Foundation, Renner, Texas. New Manual the botany (Phaenogamia and Pteridophyta) of the Rocky Mountain region, from 1885. of Coulter, J.M. New New Mexico to the boundary. American Book Company, York, York. British Coulter, J.M. and A. Nelson. 909. New manual of the botany of the central Rocky Mountains (vascular plants). American 1 New New Book Company, York, York. Wyoming. Dorn, R.D. 1977a. Flora of the Black Hills. 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