Carex juniperorum (), a New Species from Northeastern North America, with a Key to sect. Phyllostachys Author(s): Paul M. Catling, Anton A. Reznicek, William J. Crins Source: Systematic Botany, Vol. 18, No. 3 (Jul. - Sep., 1993), pp. 496-501 Published by: American Society of Taxonomists Stable URL: http://www.jstor.org/stable/2419421 . Accessed: 05/07/2011 10:07

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http://www.jstor.org SystematicBotany (1993), 18(3): pp. 496-501 ? Copyright 1993 by the American Society of Plant Taxonomists (Cyperaceae), a New Species fromNortheastern North America, with a Key to Carex sect. Phyllostachys

PAUL M. CATLING BiosystematicsResearch Centre, AgricultureCanada, Wm. Saunders Building, C.E.F., Ottawa, Ontario KlA 0C6, Canada

ANTON A. REZNICEK Universityof Michigan Herbarium, North UniversityBuilding, Ann Arbor, Michigan 48109

WILLIAM J. CRINS Ontario Ministryof Natural Resources, P.O. Box 9000, Huntsville, Ontario POA 1KO, Canada

ABSTRACT. Carex juniperorum, a distinctivenew species of sect. Phyllostachys,is described. Carex juniperorumoccurs in shallow, calcareous soil in open red-cedarwoodland and associated glades in Ontario, Ohio, and Kentucky.Section Phyllostachysis lectotypifiedand a key is provided forthe six known species. Carexjuniperorum is most closely related to C. jamesiiand C. willdenowiibut differs fromboth of these species by its shorterculms 2-6.5 cm tall, shorterperigynium beaks 1.4-2.1 mm long, and pistillate scales lacking hyaline margins.

Mackenzie (1935) noted that Carex L. sect. While studying the composition of the veg- Phyllostachys(J. Carey) L. H. Bailey (as sect. etation of red-cedar (Juniperusvirginiana L.) "Phyllostachyeae")consists of "four species of woodlands surrounding alvar openings on the dry woodlands, exclusively confined to North Napanee limestone plain of southeastern On- America,north of Mexico" and commentedfur- tario, several specimens of a cespitose sedge ther that it is "a very stronglymarked group." clearly belonging in Carex sect. Phyllostachys Mackenzie's comments hold true except that were found. These , however, differed fivespecies are now known; the fourMackenzie fromthe other species of the section present in treated,C. willdenowiiWilld., C. jamesiiSchwein., Ontario (C. backii,C. jamesii,and C. willdenowii), C. backiiBoott, and C. saximontanaMack., as well and also could not be placed with any other as C. latebracteataWaterfall (1954). known species of this section. The affinityof The section is certainly "strongly marked." these distinctive plants was clearly with the Striking diagnostic features such as the folia- group of species including C. jamesii and C. ceous pistillatescales and the few-flowered,sol- willdenowii,as definedby Crins (1990). Similarly itary,androgynous spikes on basal peduncles distinctplants were also found in Ohio by A. which are strongly dilated at the apex (Crins W. Cusick. Here we present the results of a 1990) imparta unique appearance to the group. morphological and distributionalstudy of these The section is not obviously closely related to distinctive plants resulting in the description any other group of Carex and is apparently co- of a new species. hesive and natural. In a phylogenetic analysis, Crins (1990) showed that the section could be MATERIALS AND METHODS divided into two groups of species: one, con- sisting of C. backii,C. saximontana,and C. late- More accurateinformation on the habitatand bracteatacharacterized by wide leaves, few sta- range was needed since the Napanee limestone minate flowers,wide pistillatescales, and thick, plain of eastern Ontario is not a region known clavate stigmas,the other,consisting of C. james- for endemics. Following the discovery of the ii and C. willdenowiicharacterized by narrow Ontario population and an indication that sim- leaves, numerous staminate flowers, narrow ilar plants occur in Ohio, we checked compa- pistillate scales, and thin, flexuous stigmas. rable habitats(edges of natural openings where

496 19931 CATLING ET AL.: CAREX 497 the soil was a shallow clay over limestone rock) and their distinctive features clearly justified in both areas and in the region between these recognitionof the new species proposed below. two occurrences,and to the north.In addition, The following key summarizes distinctions many herbaria were searched, including BKL, among the species in the section and is the first CAN, DAO, GH, HAM, MICH, MO, MU, NY, key to the entiresection since Mackenzie (1935). NYS, TRT, TRTE, UBC, US, and VDB. A key to the species of the sect. Phyllostachysand a dis- KEY AND TAXONOMIC TREATMENT tributionmap of the new taxon were prepared based on examination of these herbarium ma- Carex sect. Phyllostachys (J. Carey) L. H. Bailey, terials and specimens collected in Kentucky, Proc. Amer. Acad. Arts22: 128. 1886.- Lec- Ohio, and Ontario. totype,here designated: C. backiiBoott. 1. Widest leaves 6-15 mm wide, very glaucous RESULTS AND DISCUSSION ...... C. latebracteata 1. Widest leaves 2-5.5 mm wide, ? green. The originalsite in Ontario discoveredin 1989 2. Lowest pistillatescales 2.4-4.5 mmwide, much (the type locality) contained over 1000 plants. wider thanperigynia, appressed againstand Additional stations were discovered 0.2 km N essentially concealing them; stigmas of the type locality,where 15 plants occurred, straight or convolute, essentially smooth and 2 km W, where ca. 600 plants occurred.Four (papillae minute);staminate flowers 2-4 per stationswere discovered in southern Ohio and spike. fivewere discovered in northernKentucky. At 3. Perigynia 3.8-4.7 mm long; beaks 0.5-1 all of the Ohio and Kentuckystations, the plant mm long; anthers 0.9-1.2 mm long ... was abundant with populations of at least sev- ...... C. saximontana 3. eral hundred individuals. Extensive searching Perigynia 4.5-6.2 mm long; beaks 1.5-2.5 mm long; anthers 1.2-1.8 mm long ... in western New York on the limestones near ...... C. backii Watertownand in the Niagara Frontierregion 2. Lowest pistillate scales 1.2-2.5 (3) mm wide, did not reveal the plant, nor was it found in not more than 1.5 times as wide as peri- limestoneareas of northernOhio, southwestern gynia,spreading and not concealing them; and south-centralOntario, and southern Mich- stigmas flexuous,conspicuously papillate; igan. staminate flowers5-25 per spike. Despite the factthat only two relativelysmall 4. At least the uppermostpistillate scales with regions of occurrence were discovered, they broad, hyaline margins;tallest culms (6) were sufficientlyfar apart to encompass a broad 8-30 (40) cm; perigyniumbeaks (1.5) 2- 5.5 mm long. area. Withthe exception of the plants fromKen- 5. Perigynia(3) 4-9 per spike; achenes 1.5- tucky and Ohio averaging slightly larger than 2 times as long as wide; staminate those fromOntario, there was remarkablemor- scales obtuse to acute . . C. willdenowii phological uniformityamong the stations. 5. Perigynia 2-3 (4) per spike; achenes 1- The factthat the plant was not found in field 1.5 times as long as wide; staminate reconnaissance over a large geographical area, scales ? truncate ...... C. jamesii nor in herbarium material examined, suggests 4. All pistillate scales lacking hyaline mar- that it is a highly localized species, and prob- gins; tallest culms 2-6.5 (8.5) cm; ably quite rare. We have not found material of perigyniumbeaks (1.2) 1.4-2.1 mm long the new taxon filed with the species it most ...... C. juniperorum closely resemblessuch as various species in sect. AcrocystisDumort., nor other species in sect. Carex juniperorum Catling, Reznicek, and Phyllostachysincluding C. backii,C. jamesii,and Crins, sp. nov. (Fig. 1).-TYPE: Canada, On- C. willdenowii.We have not seen any collections tario,Hastings Co., Tyendinaga Twp., S side made prior to 1986. of Hwy. 401, E of Wyman Rd., 7 km NE of Although additional materialof the new tax- Shannonville, 9 June1991, Catling9100 (ho- on could not be found among the herbarium lotype: DAO; isotypes: B, BM, BUF, CM, F, materialexamined, the fieldreconnaissance did GENT, GH, K, KE, KNK, KY, MICH, MO, produce thousands of plants fromwidely sep- MSC, NY, NYS, OS, P, QK, TRT,TRTE, US, arated localities. The uniformityof the plants VSC). 498 SYSTEMATIC BOTANY [Volume 18

E

1mm 1mmi X 1mm X X

C 1mm~~~~~~~~~m

lmm ~ ~ 1m m

FIG. 1. Carex juniperorum. A. Habit. B. Ligule, and lower blade of outer leaf. C. Upper sheath. D. . E. Basal pistillate scale (right),upper pistillate scale (left). F. Perigynium. G. Perigynium dissected to show achene. H. Achene, side view (bottom), top view (top). I. Portion of staminate spike showing anthers. Drawing by Marcel Jomphefrom Catling 9100 (DAO). 1993] CATLING ET AL.: CAREX 499

Plantae valde cespitosae, culmi 2-6.5 (8.5) cm long, 0.8-1.2 mm wide, encircling rachis and alti; vaginae basales rubiginosae, glabrae. Folia connate basally, truncateto obtuse, sometimes 4-8; laminae 5-28 cm longae, 2-4 (5) mm latae; mucronate, whitened on erose-ciliate apex, vaginae 2-3.5 cm longae, glabrae, zonis interio- brown or reddish-brown with a green center ribus vaginarum hyalinis vel rubiginosimacu- below, essentially nerveless. Perigynia 3.8-5.1 latis. Inflorescentiaex 1-4 pedunculis basilibus mm long, 1.5-2 mm wide, ascending, nearly spicis androgynis solitariis foliis absconditis terete in cross-section and tightly investing constructa. Squamae pistillatae (7) 11-29 (38) achene, green,becoming whitish-green in fleshy mm longae, 1.2-2.5 (3) mm latae; squamae stam- basal portion,sometimes red-flecked,glabrous, inatae 0.1-0.5 mm longae, 0.8-1.2 mm latae. Per- sessile, 2-veined, contractedabruptly into beak; igynia 3.8-5.1 mm longa, 1.5-2 mm lata, sub- beak ? flattened,(1.2) 1.4-2.1 mm long, 0.6-1 teretia, viridia interdum rubiginosimaculata, mm wide near base and gradually tapering to glabra, in rostrumcontracta; rostra (1.2) 1.4-2.1 apex, serrulate-edged, reddish-brown below mm longa, 0.6-1 mm lata, serrulata.Achenium pale whitish-brownbidentate or flattip. Achene 1.9-2.5 mm longum, 1.5-2 mm latum. Stigmata 1.9-2.5 mm long, 1.5-2 mm wide, terete,dark 3. Antherae 3, 1-1.5 mm longae. brown at maturitywith a prominent,enlarged, pale brown stipitatebase ca. 0.3-0.5 mm long. Plants densely cespitose, with rhizomes up Styles thickened at base, articulated in a de- to 3 mm long between adjacent shoots; roots pression on top of achene; stigmas 3, elongate brown, the younger with a sparse to moderate and prominentlypapillose. Anthers3, 1-1.5 mm covering of hairs 0.16-0.48 mm long; culms 2- long. 6.5 (8.5) cm high, with 2-3 (4) glabrous reddish- The specific epithet "juniperorum,"("of the brown bladeless basal sheaths. Leaves 4-8; junipers") is given in recognition of this spe- blades 5-28 cm long, 2-4 (5) mm wide, plicate cies' close association with Juniperusvirginiana. to flat,leathery, glabrous, green, margins an- Distribution(Fig. 2). Eastern Ontario and trorselyserrulate distally; leaf sheaths 2-3.5 cm southernOhio and adjacent Kentucky,on clay- long, tightlyenveloping culm, glabrous, lower ey soils derived fromcrumbling limestone bed- reddish-brown on back, upper reddish-brown rock in open woodlands with virginianaand basally to brown, green, or whitish; inner band J. Quercusspp. surrounding natural glade, alvar, hyaline, varyingin color with the back but also or prairie openings. sometimesred-dotted near slightlyconcave, flat, Phenology. Flowering and fruitingin April or slightly convex, thickened apex; ligules of and May in the southern portion of the range, lower leaves truncateor even concave or emar- fruitingextending to June in the northernsta- ginate, those of inner leaves triangularand up tions. to 3 mm long. Vegetative shoots 8-30 cm tall; pseudoculms 1.5-6.5 cm high and 1-2.5 mm wide, with 2-4 dark reddish-brown bladeless Additionalspecimens examined. CANADA. Ontario: basal sheaths; leaves 5-8; blades up to 40 cm Hastings Co., Tyendinaga Twp., Salmon River Alvar, 7 km NE of Shannonville, 26 June 1989, Catling& long, 0.9-3 mm wide, otherwisesimilar to those Catlings.n. (MICH), 14 June 1990, Catling8205 (DAO, of fertileculms. Inflorescenceof 1-4 erectbasal UBC, herbariumW. J.Crins), N of quarry,W of Salm- peduncles of varying lengths, each with a sol- on River, 5.5 km NE of Shannonville, 11 June 1991, itaryandrogynous spike, arising centrallyfrom Catling9102 (DAO, MICH, TRTE), N side of Hwy. 401, a pseudoculm of the previous year and ? hid- E of Wyman Rd., ca. 7 km E of Shannonville, 9 June den among the leaves; peduncles 1.2-6.5 (8.5) 1991, Catling9101 (DAO). cm long, filiformnear base, 0.9-1.9 mm wide U.S.A. Kentucky: Bath Co., "Knob Lick," 1.5 miles in expanded distal portion,triquetrous, smooth W of Polksville on US 60, 7 May 1991, Campbells.n. except for antrorselyserrulate, winged angles. (MICH), "Blue Clays" on Fearing Road, 1 mi W of Polksville on US s.n. Spikes 5-13 mm long, with (2) 4-9 pistillate 60,31 May 1991,Campbell (MICH); Lewis Co., 1/4mi S of Crooked Creek Rd. & 1?/2mi NW flowers and (5) 7-15 (21) staminate flowers. of St. Rt. 57 at Covedale, 9 May 1990, Cusick 28813 Lower 11-29 mm pistillate scales (7) (38) long, (MICH), 7 May 1991,Reznicek et al. 8755 (DAO, GENT, 1.2-2.5 (3) mm wide, spreading and not con- KY, MICH, TEX), Hughes Knob, ?/4mi N of East Fork cealing perigynia, leaf-like, long-acute, serru- Church, offChalk Ridge Rd., 7 May 1991, Rezniceket late-margined,green, 6-8-veined, leathery;up- al. 8754 (DAO, KNK, MICH, US, VDB, VSC, WIS, herb. per much reduced. Staminate scales 0.1-0.5 mm C.T. Bryson,herb. V. McNeilus), Hymes Knob, N side 500 SYSTEMATIC BOTANY [Volume 18

830 800 45o

450 -

v S f \ \ \ 1 < ~~~~~~~~~LakeOntario

Lake HuroLakn 420 k@ 4,

1- 1 ;7 P~~~~~~~~~~~~~~ennsylvani>ag

420~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ :7 < ~~~OhioXX

390

39 0 West Virginia

200 km Kentucky

830 800

FIG. 2. Distribution of Carex juniperorum.

Chalk Ridge Rd. 11/4mi of TrinitySchool, 7 May 1991, OS), Adams Lake State Park, Adams Lake Prairie,W Rezniceket al. 8756 (MICH). Ohio: Adams Co., Lynx of State Rd. 41, 1.5 mi N of West Union, 6 May 1991, Prairie (no. 2), Lynx Prairie Preserve, 1/4mi SE of E Rezniceket al. 8742 (BUF, DAO, GH, KE, MICH, MO, LibertyChurch and 1.5 mi S of Lynx, 18 April 1986, NY, OS, PH, US, VDB, VPI, WARM, WIS, herb. C.T. Cusick35061 (MU, NY), 6 May 1991, Rezniceket al. 8748 Bryson, herb. V. McNeilus), Chaparral Prairie Pre- (CM, DAO, K, KNK, KY, LE, MICH, MIN, MO, NYS, serve, Hawk Hill Rd. 31/2mi NW of West Union, 6 1993] CATLING ET AL.: CAREX 501

May 1991, Rezniceket al. 8744 (DAO, MICH), Green ciliate. On the other hand, the perigyniumand Twp.,"Hanging Prairie" N of BlackRun Rd., 3 mi staminatescale morphologyare similarto those SW of Lynx, 6 May 1991, Rezniceket al. 8750 (MICH). of C. jamesiirather than those of C. willdenowii. Comments. Carex juniperorumis typically The lack of a hyaline border to the pistillate abundant in its habitatsand is one of the major scales of C. juniperorumis differentfrom both C. to the dominants of the ground layer. An associated jamesiiand C. willdenowii,although similar dominant at all sites was Danthoniaspicata (L.) scales of the C. backiigroup. The extremelyshort P. Beauv. At the type locality and nearby On- culms are unique, though approached in C. tario stations, the herb layer was additionally willdenowiivar. megarrhynchaF. J. Herm. Con- dominated by Astercordifolius L., C. siccataDew- sequently C. juniperorumis a very distincttaxon. ey, C. pensylvanicaLam., C. umbellataWilld., However, it is easily overlooked because its fruit Comandraumbellata (L.) Nutt.,Fragaria virginiana ripens very early and the culms are hidden at some- Mill. var. virginiana,Hieracium piloselloides Vill. ground level well below the leaves, and and Poa compressaL. Among the other species times within the moss mat. present were several with ratherrestricted dis- The recent discovery of this distinctivenew tributionsin easternOntario, including Calyste- species in a supposedly botanicallywell-known northeastern gia spithamea(L.) Pursh ssp. spithamea,Carex backii, area suggeststhat even the floraof com- C. richardsoniiR. Br., Geumtriflorum Pursh, Hed- North America is not as well-known as is monly supposed. This species has now been yotislongifolia (Gaertn.) Hook., Monardafistulosa of the L., Oenotheraperennis L., and Potentillacanadensis collected from12 sites. Fortunately,three L. At the Ontario sites, Juniperuscommunis L., Ohio sites, the Chaparral Prairie Preserve, Ad- Rhamnuscathartica L. and Rhus aromaticaAiton ams Lake State Park, and the Lynx Prairie Pre- are were frequentshrubs. The pH of the soil ranged serve (E. Lucy Braun Preserve), protected for the from 7.0 to 7.8, and the soil was almost com- natural areas, and thus the prospects good. pletely covered by the moss Thuidiumabietinum continued survival of this species appear (Hedw.) Bruch, Schimp. & Giimbel. with some ACKNOWLEDGMENTS. We are very grateful to Al- Brachytheciumsp. lisonCusick for invaluable help finding sites for Carex Locally, in the Ohio and Kentuckysites, Schiz- juniporumin Ohio and Kentucky,and forbringing his achyriumscoparium (Michx.) Nees, Carex meadii earlycollections to our attention.We also thankthe Dewey, Hypoxishirsuta (L.) Coville, Liatrisspp., Ohio Departmentof NaturalResources, Division of Silphiumterebinthinaceum Jacq., and Sisyrinchium NaturalAreas and Preserves,for cooperation with workon theirpreserves. Julian Campbell shared with albidumRaf. were frequentassociates. Soils were us hisdiscoveries of this species in BathCounty, Ken- calcareous, with pH 6.8 and 6.7 and Ca++ con- tucky.Julian Campbell, Vivian Catling, Ray Cranfill, centrations2080 and 1720 ppm respectivelyin Don Cuddy,Allison Cusick,Max Medley,Michael two sites sampled (vouchered by Reznicek8742 Oldham,and SusanReznicek helped with field work. and 8754, respectively).The habitat is moist in HowardCrum kindly identified the mosses from the early spring and becomes quite dry in summer. Ohio and Kentuckysites. Marcel Jomphe prepared As with the Ontario stations, there was a sig- the illustrationof C. juniperorum. nificantmoss layer, including Thuidiumabietin- um, T. delicatulum(Hedw.) Mitt., T. recognitum LITERATURECITED (Hedw.) Lindb., Bryoandersoniaillecebra (Hedw.) BOETTCHER,S. E. and P. J.KALLISZ. 1991. The prairies H. Rob., Campyliumchrysophyllum (Brid.) Lange, of the E. Lucy Braun Preserve,Adams Co., Ohio: and Fissidenscristatus Mitt. Recently,Boettcher A soil study. Ohio Journal of Science 91: 122- and Kallisz (1991) have published a detailed soil 128. and vegetation study of a characteristichabitat CRINS,W. J. 1990. Phylogenetic considerations be- of C. juniperorumin Ohio. low the sectional level in Carex.Canadian Journal With its 4-9 perigynia per spike, coriaceous of Botany 68: 1433-1440. leaves, and truncateto rounded, often brown- MACKENZIE,K. K. 1935. Tribe 1. Cariceae. North ish, thickened, entire ligules, C. juniperorumis American Flora 18(4): 174-177. T. 1954. A new species of Carex(sect. similar to C. willdenowii.Carex jamesii has fewer WATERFALL,U. Phyllostachyae)from Oklahoma. Rhodora 56: 21- perigynia per spike, softerleaves and the lig- 23. ules are hyaline, rounded to nearly acute, and