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A RE-EVALUATION OF CAREX SPECUICOLA AND THE CAREX PARRYANA COMPLEX (CYPERACEAE) PDF

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AND THE OF CAREX SPECUICOLA A RE-EVALUATION COMPLEX (CYPERACEAE) CAREXPARRYANA Murray AAReznicek D.F. RESUMEN INTRODUCTION taxonomic difficulty.Adding to this dif^ beenasource of Dewey have always Carexparr^ana Plants referred to and uncommon or Carex parryana its rare. widespread, are though some species are all even ficulty is that, have inflorescences that having ±elongated members Racemosae in from other of section close relatives differ com- mm not unispicate), spikes or ascending lateral (if 4-6 erect narrow 8 wide), cylindrical, relatively (2 mm Mackenzie recog (1935) and C. utahensis). long (-3.9 in C. specuicola perigynia 1.7-3.3 bined with small and par idahoa L.H. Bailey, C. E. Jones, C. halln Olney, C. complex, C. aborigii nized four species in this consisting of di species parryana variable Murray recognized C. ryana. (1969) ; Murray, par C. (Olney) D.F. parryana subsp. halln nd them subspecies; C. treated as obon^num, He parryana. did not treat C. and parryana subsp. Murray C. D.F. ryana subsp. idahoa (L.H. Bailey) q-K and parryana a Mackenzie (1935) was close to C. described in ‘Carex olwriginum- as realizing lhat much perig^m (4.W5- M endemic with larger which a local Idaho than nboriginum E. Jones, is ent plane C. Hermann recognized C. hath, mm W. Boon (Murray 2002). (1970) sermtodens 6.6 long very similar to C. described in “Carex aboriginum” as noted that M.E. Jones as spe< and aboriginum It idahoa, C. parryana, C. He treatedCnborynum of ME. typ^Mype. from Jones’s Maekenzie(19M)i; -completely different- C C. Herm. Murray (2002) treated C. tdahoa. linlln. ME. patrynnn brevisrparma FJ. not Jones, as C. var. zie, was not aboriginum- C. and noted that Mackenzie s “C. also and ohoriginum as species, parryana. C. Goodnch (m Welsh 2003) et al. M Finally, parryana brevisrfuoma. recognize C. var. but did not of E. Jones, m An. northern hanging gardens an endemic of by Howell (1949) as described concluded that C. specuicola, parryana in Utah. and recognized only C. parryana, within belonged C. zona, also ^ , pa^ana compt, a,there has fortu- Carex within the range of the activity decade of floristic in the last me w^ C many new speamens. This especially lor a great generate been enough collecting to nately Murray 2W2). North America, when was flora of treated for which lew specimens existed it specuLla, for we and new popuUuons, number of discovered a («e Roth 2004) Nation Navajo working with the Botanists 38 Journal of the Botanical Research Institute of Texas 7(1) now We have an excellent series of specimens to study. reviewed these new collections with several questions in mind. specukola a good Is C. species? Is recognition of C. hallii, C. idahoa, and C. parryana as species sup- ported by additional collections? Finally what is the identity of “Carex aborigimm” of Mackenzie (1935) not M.E. Jones (= C. parryana var. brevisquama FJ. Herm.), and is it a good species? We also provide illustrations of Plants referred to Carex parryana divide into two clear groups based on morphology and One geography. group, including C. specukola, has much strongly flattened perigynia than and larger loosely over the fitting achenes, an admixture of and 2-styled 3-styled and flowers, is essentially confined to Utah and Arizona. The other group, including C. hallii, C. idahoa, C. parryana, and “C. aboriginum” of Mackenzie not M.E. with Jones, pengyma the tightly enveloping the achene and entirely 3-styled flowers, is widespread, occurring from On- tario to Alaska, but south only to Colorado, Utah, Nevada, and California. The former group, with loosely fitting perigynia andamixof2-styledand3-styled two flowers consists of One species. is the lax hanging garden endemic Carex specukola with pale perigynia. In we addition, recognize anes^ntiallyalloF^tric and more widespread species fromadiversity of wetland habitats that isusually stiffly upright, wnhclearlypurple-tintedperigynia, which wedescribeasCut^^^^^^ In the latter group, with pe- ngyma tightly enveloping the achenes, the plants erroneously MackenL referred to Carex aboriginum by mm i^wKlest We 4-6.8 wuie. recognize this taxonatspecies rank and dcscribeithemnasC.l.olmgrento™m “ “ mm ">“> O n.rt r “ 3.6 3 ’ ' remaining Cara species of the parryana complex as recognized Murrav Qonil in W^rl""”"^^^^^^^ bestabuZhthe I mm spiL relatively broad, 3-5.5 C wide eral of LlJii anH Lat- clrrhlfliUndr and C. parryana show great variability in inflorescence k size number, ^P^^e and.2 , ity. Nevertheless, most culms spike sexual- of Carex k k hallii iftyqftnf lon^tlatem, C.pnr,,nm.,ttZr::^~W^ spike, in “ X “I”- ofient mfiTsTZo - 'rm f heightrrZ^ofZlXXtmfcZ L Whi and C. parryana do ^^ows not otZmfZ represent that C. emis X parryana group 2 3 4 5 6 number Spike C Fk;. 1. Distribution of spike numbers in Carexhallii and parryana. Achenes nnm 0.8-1 wide; .2 lateral Spikes (2-)3^-6), the 5. 1 times as long as the longest mm Loosely cespitose to slightly colonial from slender, ±elongate rhizomes 0.9-2.3 wide, rhizomes ca. 0.5-7 cm long between shoots, rhizome scales pale brown, rapidly disintegrating into fibers; culms (7.5-)19-70(- diately below the inflorescence; phyllopodic, cataphylls and basal sheaths pale to dark brown, youngest some- mm cm times tinged reddish-purple. Leaves 5-11, essentially basal; blades 1.8-38 long, 0.8-3.3(-4.5) wide. 1 5 ! group parryana Systematia of the Carex and Murray, Reznicek mm in cross section; the convex sides elliptic nattened-trigonous or occasionally lenticular 0.2-0.45 long, stipe brown, smooth; apex apicu ate achenes, trigonous in narrowly achenes, elliptic in lenticular nearly circular to stigm^ mostly mm deciduous; 2, formed by the peKislent style base; the style up 0.15 with an apiculus to mm mm 0.1-0.3 long. apiculus with a conspicuous ca. 30% Anthers 1.2-2.7 long, up sometimes to 3. 3, Care* spccuicota oc- “hanging gardens.’ sandstone cliffs— along shallow caves or alcoves Moist seeps in many meters of the many 400-300 plantssptead over culms as populations consisting of a few to as curs as local Utah. andimmediatelyadjacentsouthemmos. Arizona m.Endemic.o northern 1350-2323 Elevation cliff face. hanging gar ens species of Typical associated “hanging gardens." sandstone an endemic of Navajo This is Cirsnim micrantlm Eastw., vuginula Rydb., Aquilcgia Aniicica Adiantum cnpiUus-veneris L., region include & in ihe wtheam (Higgins S.L. caslwoodme Rydb., Plfltnnthera Mimulus Hook., Epipoctis gigniitcn rydbergii Petr., Though very local, this spe- & L.H. Bailey. Lndht and lew other Cnrex, especially C. hnssei 4lsh*Kartesz a remains constant some The arching febit lax, dominant cover in sites. and the can form a dense turf is cies cantedifficu^o .hough shownin Figure this growingonahorizontalsurface as 3, when are even the plants m 4 an noted Figure a yp and achenes are A sampling of perigynia, scales, herbarium material. discern in shown Figure 5B. specimen in from a herbarium is inflorescence cal May 5 2001, W Chelly, Canyon ie ARIZONA. Apache del Co.: specimens. Representative O’Kane, m 2276 (NAVA); Carson Mesa, 12Jun 2001, Jr. < canyonSWofIm narrow ASC); mi Walker Creek, (NAVA); 1 15 Wl^ev (NAVA); SW n 1340 Mesa,njul N Canyon Kayenta and ofBoot of (NAVA); Tseyi-hatsosi Aug Roth 1701 Mexican Water, 21 2003, 16W NE Rock 21 Aug 2003. Roth Dancing Rocks of Point, ( N Canyon 20 May 2002, Rinfe 1154 (ASC, BRY); of Giant trail, 9,Hevron214b(NAVA); N Canyon. 30 Sep End Canyon of Geshi W head Far ins (NAVA). Navajo Co.: of h & small spring, Roth Holiday 841; & Canyon. Rink 923 (NAVA); Jackass 1 3, Canyon. 2 Oct 2000. Roth Dowozhiebito watershed Canyon, 13 Sep 2000, end of Keet Seel sideofLongCanyon.SkeletonMesa,18May2004.Rinfe&Bungart3146(ASC.SJNM.NAVA). ; ,„,„v4ave p & Angel Arch Canyot UNITED STATES. Utah. San Juan Co.; Murray, r Reznicek D.F. sp. Type; Carex utahensis /. 2. 2Jun 1964, 5.500 andjuniper-pinyon cc slope, ca. ft. bottom 1 inyon mm cm between 1-5 long rhizomes 3-2.7 thick, ca. elongate rhizomes from 1. colonial Loosely cespilose to 22-55 cm, culms stiff, into coarse fibers; disintegrating brown purplish-brown, rhizLe to shoots scales camphylb mm below the inflomscence; wide immediately 0.5-1.4 papillose, trigonous, .erect,' ba« blades 3^ esKutmlly Leaves 6-12, reddish-purple. youngest often , and sheaths brown, the basal midnb smooth Enely an ror^ly and to mm margins much than culms, .folded, shorter wide, 1.8-4.5 long. ^mlong, gla ' sheaths 1.7-15 abaxially; leaf ±smooth papillose adaxially, scabrous distally, mm long 0.8-3.7 concave the apex; hgules ca. . shallowly at brown, hyaline, thin, pale ish or cm 2^ long, brown, inflorescences portion whitish to pale than wide, free shorter starninatgpUtillm^^^^ termlrmlgynaecandrous or occasionally t:X»ed.TLinalspikel2-31nimlongpistm^^^ mm How mm (0-)2()-65 wide, ca. 1.8-3.1 (0-)6-24.5 long, portion staminate widP f0-)L50-flowered ra mmlong,3.5-6nrm,^e,cyhndnc,^^e^^ 4-22.8 emd,'p^unchL8-lL5mmiong,Im.eralspikes mmwi mm long, 2.3-3.7 1. Staminate scales , ’ . 7-40 ascending perigynia. with ca rounded, obtuse or acute to especiaflydistally.apex hyahne margins, narrow brown purple with mm to awn up toO.3 long. Pistillate apiculus or short prolongedmtoa±scabrous-ciliate Lmerved center rarely margins inconspicuous, mm hyaline reddish-purple, mm reddish-brown to wide, 3-2.2 9-3 3 long 1 s 1 scabrous-cihate prolonged into a sometimes 1-3-nerved center green • ' ’ the - acute, i #1 ' 1 i ^ i V / i mm awn mm apiculus up mm or to 0.5 long. Perigynia 2.6-3.9 long, 1.6-2.5 wide, ±strongly flattened to con- m cavo-convex cross-section, ±obovate, not by much filled the smaller achene, herbaceous, papillose distally green to reddish-purpledistally,greenish brown to proximally,2-nervedand withafew frequently nervei faint Z ^ obscure to well defined beak cylindrical teaks01-0.4tnm long, and papillose sometimes setulose. green to purplish-brown, apex *erose obscurel, to sometimes lenticular in cross section; the convex sides narrowly cm ellinti gonousac r enes, ™^dfy elliptic in lenticular ^ achenes, brown, smooth apex usually apiculatewitl -"““y “p “> 'T'* %TATtto!3 3. %2.A„te3.1.8-2.6mn.lo„g,„ftenwia.aco„spc„o„sapiculmca.0.1-0.3mmbnfr . group parryana the Carex Systematics of and Murray, Reznicek FromthestateofUtahwherethetypecollectionisfrom. Etymology.— ^ Elevation bottoml,ands. meadows, and riparian wet and hanging gardens, wet alcoves slopes, Seeps, westernmost Colorado. Endemic Utah and adjacent 1400-3000 m. to nORfO mapped by Albeeaal. (1988) and WeUhetal. 2003) Goodrich Utahby (in PlantstreatedasCpanyana in below). and C.holmgreniorum (see species also are this Jt similar to C. Colorado Plateau ri northern portion of the be endemic to the Car« uMhcnsis appears to «ou^^ thus fitting perigynia lonely the much than the perigynia, smaller having achenes speemeda in darkerpistilbm^ with utahensisisarnorestimyerectplant, Bute, aeheneandnotfilledby it, “iri Cnrex specuicoln has a lax, purple tinged apically. perigynia distinctly and obovate green clearly A maturity. and iranslucent at whitish changing to perigynia and pale greenish, elliptic Sder scales Distuiate dCandatypicalinfloreseencefromaherbarium Figure Lphngotpengynia,scales,andachenesarenotedin shown Figure 5C. specimen in is rniOHDQ.Mo.ln,«C.iT47N,21)W.S13.L.S.ICi«kCy.l.5n.lEoiUul,-Ce...»a„K'iH„,Slanl994. . T.h,37M(COLO.UTC).OTAH.Ci«l«».Co.:TMSE17E.S17.3Ja„2«Xl,At,<,<»i25i72(Bm R.MSwdl.5Jun 1979,1 San (BIt«lBigE«aMt.,8A«gl9r7,Le»ii3125(IlRY); WWn&Mooir 8j™ S3 1979. May 4573 &. Yeatt Wash, 6 2001, Yeatfs Cottonwood San Rafael Reef, Jun Tuhy 3545; 13 1989, Canyon, Nebo peak from Pole 17 & Juab Mt. trail Co.: Evenden 24445 (BRY). Atwood 1999, (NAVA M with Slickhom Canyon, 1 jun 2005, Roth 1847 Co mi upstream from San Juan R. jet. 1 May ^16 (BRY). 2000, I & Canyon bottom, 14 22024 (BRY); John’s Welsh Chatterley 1983, (BRY) Canyon.3Jun2000.Atwood25765 Moonshine Canyon. Uintah Firewater 1900 (BRY). Co.: abon^num. Amer. Caric. pi. 421, as C. Mackenzie, K.K. 1940. N. 2, niustration.— mm rhizomes 0 5 ca. 2-3.5 thick, rhizomes ascending from short, Loosely cespitose cm 20-90 culms tall, into coarse fibers; brown, disintegrating rhizome scales shoots mm below the inflorescence; immediately wide 0.6-1.1 trTlous papillose, very finely phyllsandbasalsheathsbrown,theyounge3tusuanyreddlsh-purpk.Leavesca.M2^^m^l^^^^^^^^ midnb mooth an and to finely margins mm much than culms, ^folded, shorter cm 5-4 wide, 3 48 long 3 1 ™;!y^ltdis.aUy,Asm„othadaxial.y.papiUoseabaxiaUy,I^ mm long, acute to 0.8-5.5(-7.5 hgules apex; ca. concave the at brown, hyaline, thin, whitish or pale trally freeportionwhitishtopalebrownJn^^^^^^ than wide, the than to longer shorter rounded, slightly gynaecandrous a era s p sometimes staminate or , cm terminal with 3-5(-6) spikes, lone e7ai4e3-eriappingVersou.e.in.essepa«e.lowe7.wospikesa^^^^ mm cm and mm up long 1.5 to 7.5 lowest bracts ±bristle-like un 2 long; npHiinrle to 14 rriAtVi mm loug^^^^ Terminal spte(6.5-)11.5-27 raLss,up^rbractsabEupdy reduced. 10mmlong,4.5-7.2mmwide,ca.(0-)l-28-nowered,3tamina,eporuon4.5-2b3mmlo^ai^M^ mm ^ mm 4 2 ong, spikes Lateral 1.8-38 long. peduncle 10-65-flowered, ca. ' ^ 2 4 ^ Staminate scales ^ perigynia. 5-45 spreading-ascending with ca ' denselv flowered f’ , . mrjM"oL,e,reddI-brown\uhbroadbyaUne margins "dcen.er.Pis.illa.esca.esl.^2.3mm,ong,1.5-2.2rn^ ™ obovare^e— ^ broadly wwfr:2ied plumply biconvex, ro rrigonous br^n^x .ru^, green ro and so.ne.imes 2^.5 papillose mn. beaks 0 long, finely mm beak; mm mm L3-L6 wide, concavely Achenes L7-2.4 long, up long. 0.2 with teeth to Obscurely bidentulate , mm tapering to a thick stipe 0.1-0.2 long, flattened-trigonous in cross section; convex sides narrowly elliptic to mm obovate, brown, smooth; apex usually apiculate with an apiculus up to 0.2 formed by the persistent style mm mm base; style deciduous; stigmas 3. Anthers 3, 1.7-3 long, with an apiculus ca. 0.1 long. Etymology.—Carex holmgreniorum was chosen to honor Arthur H. Holmgren and Noel and Patricia Hol- who mgren, together have collected this species several times and whose scholarship has contributed so much our knowledge to of the Intermountain flora. Alkaline seeps, adjacent to springs, riparian zones, moist meadows, pasture land, calcareous and often saline soils, with species such as Juncus balticus Willd., Eleocharis and rosteUata (Torr.) Torr., Schoenoplectus pungens (Vahl) Palla. Elevation 1400-2000 m. UNITED STATES. Idaho, Nevada, Utah. Some Utah of the localities of C. parryana mapped by Albee presumably et al. (1988) refer to this species. Publication of Carex parryana Hermann brevisquama by var. (1968) lacked a citation of a type specimen was as required by the Code. Valid publication did not occur until 1970 when the name was again published by Hermann with Holmgren 8251 as the type for the name (the holotype, and apparently the only sheet seen by Hermann based on his annotation on the sheet). The epithet brevisquama was preoccupied rank the of at This is a distinctive species whose lack of was recognition due and to its relative rarity the confusion sur- rounding the misapplication name of the C. aboriginum. It is easily recognized by combination of its large pis- tillate lateral spikes, staminate or gynaecandrous terminal spike, broad achenes and perigynia, and pistillate scales distinctly shorter than the perigynia. an uncommon It is and local endemic of the northern Great Basin and western portion ol the Colorado A Plateau. sampling of perigynia, and scales, achenes are in Figure 4A 1 and a typical inflorescence from an herbarium specimen shown is in Figure 5A. Repre^ntauve specimens. IDAHO. Caribou Co.: Soda Springs, 9 Jun 1993, Moseley 2620 (BRY). NEVADA. Elko Co.: F National Wildlife Headquarters, 20 Jun IS (BRYM^ 8745 Im us,mnom.-Macltenzie. K.K. N. Amer. aric. pt CnmquUt, 11: 412; ^ A, etal, 1977. iutermountain Flc cm talUtiff, erect, trigonous, papillosewithltj::”^^^ finely mm wide ^ immediately antrorsely below scabrous, 0.5-1 the inflorescence- nhvllonorlir n . i, , usually reddish-purple.Uavesca. 6-12, ess^nttallylsaifcSS^'^^^^^^ shorter than culms, .folded, margins and sLth mXsI midrib L'” to smooth dislally, adaxially, n umrotscly scabro papillose abaxially ' ‘'"''V leaf sheaths 2 s brown, ^"”"i *'®^'^^^^ hyaUne, thin, cotmave theapexilJusItS^fl at lonpr than wide, the iX *' free portion whitish to pale brown pistuiate, upper *‘™"'«e, lateral spikes or rarely .mixed, .overlapping bwweerr som,H™ lateta sometimes separate, lower two spikes cm 0.3-5.2 distar

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