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Home , Anostraca, Bow Peak, Branchinecta

This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. JOURNAL OF BIOLOGY, 13(1): 184-189, 1993

PERSISTENCE OF PALUDOSA () IN SOUTHERN WYOMING, WITH NOTES ON ZOOGEOGRAPHY

James F. Saunders III, Denton Belk, and Richard Dufford

ABSTRACT The fairy Branchinectapaludosa is a persistentresident of aestival ponds at high elevation in the Medicine Bow Mountains of southernWyoming. These populationsare far removed from the Arctic tundrahabitat that typifiesthe distributionof the species, and appear to representthe southern margin of the range in . All of the records for the northernUnited States and southernCanada appear to lie along the CentralFlyway that is a major migrationroute for waterfowland shorebirdsthat nest in the Arctic. Passive dispersal probablyprovides for frequentcolonization of marginalhabitats and gene flow to established populations. The fairy shrimp Branchinectapaludosa have been deposited in the University of (Muller)is widely distributedin the circum- Colorado Museum (UCM 2192, 2193, polar tundra of the Holarctic region (Vek- 2194). The Snowy Range is an axial rem- hoff, 1990). In Europe, it occurs chiefly at nant which rises about 300 m above the latitudes above 60?N, but there are isolated surrounding Medicine Bow Mountains recordsfrom the High Tatra Mountains on (Houston and others, 1978). The ponds are the borderbetween Czechoslovakiaand Po- mainly in the upperTelephone Creek drain- land at about 49?N (Brtek, 1976). Records age at elevations of 3,200-3,350 m. Most for Russia are typically along the Arctic of the ponds are underlainby the Nash Fork margin, but include the southern tip of the formation (Houston and others, 1978), and Kamchatka Peninsula at 52?N (Linder, the characteristicmetadolomite is present 1932). In North America, it is found in in all but one of the ponds. The ponds sup- Alaskaand most of the provinces of port no and most tend to be low in ionic (Linder, 1932; Reed, 1963; Belk, 1975; Da- content (Table 1). born, 1978). There are also a few scattered Prior to our collections, B. paludosa had records from the contiguous states of the been reportedtwice from the Medicine Bow United States (Linder, 1941; Lynch, 1958; Mountains (Linder, 1941; Home, 1967). Belk, 1977; Saunders, 1981a). The occa- Linder'sreport is very likely from the same sional southern record notwithstanding, general area as the present records. The most known populations occur near the sample (USNM 78574) was collected by Ir- Arctic Ocean. ving H. Blakeon 4 July 1935 from the Med- Recent collections of B. paludosa from icine Bow Mountains, and contains rather the Medicine Bow Mountains of southern cryptic locality data: "G.C. no. 303-3 (10) Wyoming suggest that the species is both a. ch. Sta. E. Temporary pond. alt. 9,700 locally common and temporallypersistent. ft." Althoughthe 9,700 ft (2,957 m) contour Branchinectapaludosa was collected from traversesconsiderable territory in the Med- eight ponds in the Snowy Range in Albany icine Bow Mountains,the possible locations County, Wyoming (R79W, T16N, sections are drasticallyreduced by examining inter- 9, 10, 15, 16 on USGS 7.5 minute series sections of that contour with roads that ex- Medicine Bow Peak Quadrangle).This spe- isted in 1935. The most likely prospect is cies has been collected consistently since WyomingHighway 130 that skirtsthe Green 1987 in conjunctionwith studies at the Gla- Rock Campground(now a picnicarea) which cier Lake Ecosystem Experiment Site bearsthe United States Forest Service iden- (GLEES).These ponds are best character- tification number 303. A small temporary ized as aestival (Eng et al., 1990; pond exists nearthe picnic areaon the south Daborn and Clifford, 1974) because they side of the Nash Fork at an elevation of always contain water but may freeze to the approximately3,000 m (R79W, T16N, sec- bottom in winter. Specimensfrom two sites tion 13 on USGS 7.5 minute series Centen- 184 SAUNDERS ETAL.: BRANCHINECTA PALUDOSA IN WYOMING 185

Table 1. Major ion chemistryof three ponds inhab- The zoogeographicalsignificance of the ited by Branchinectapaludosa in the Medicine Bow Wyomingrecords is greaterthan recognized Mountains, Wyoming. previously.The Colorado-Wyomingborder Pond near is probablythe southernextent of the North Pond near Towner Pond American distribution of B. A Lost Lake Lake Road sampled by paludosa. 17 June 1991 08 June 1991 Hore (1967) previous record of B. paludosa from the pH 6.1 7.0 7.0 Kaibab Plateauin Arizona (Belk, 1977) has Ca, mg/l 0.8 2.8 2.0 proven to be a misidentification.A descrip- Mg, mg/l 0.2 0.7 3.0 tion of what is now known to be a new Na, mg/l 0.2 0.2 1.0 speciesof Branchinectais in preparation(M. K, mg/l 0.2 0.5 0.0 and D. There is also a C1,mg/l 0.2 0.2 3.0 Fugate Belk). pre- SO4,mg/l 0.9 0.5 2.0 vious record for a site near Salt Lake City, HCO3,mg/l 2.2 12.0 4.0 Utah (Maynard, 1976), but the validity of Conductance, the report has not been confirmed despite 29.0 35.0 umho/cm 9.6 effortsby Belk to locate voucher specimens. Salt Lake City lies just south of the latitude that separates Colorado and Wyoming. nial Quadrangle).At the time the pond was Confirmed records published for the con- located in early August 1991, it was nearly tiguous United States are thus restrictedto dry, but yielded one specimen of B. palu- Irish Canyon of extreme northwesternCol- dosa. This pond remains the most likely orado (Saunders,1981 a), the plains of north match for Blake's locality despite a small central Montana (Lynch, 1958), and the discrepancyin elevation. Medicine Bow Mountains of southern Wy- Horne (1967) reported that B. paludosa oming (Linder, 1941). Lynch's specimens occurred in alpine ponds in the Medicine from Montana are in an uncataloged col- Bow Mountains, but gave no specific local- lection in the USNM (Acc. No. 294523). ity data. On the basis of elevation alone The record of B. paludosa from north- (3,000-3,500 m), the pond must be very western Colorado is very close to the Wy- close to the ones that we sampled. Further- oming border and thus is little differentin more, the chemistry is similar (Table 1). latitude from the Wyoming collections, al- Horne recalls collecting B. paludosa from a beit at a lower elevation. The ability of this pond in the upper Telephone Creek drain- population to inhabit the same pond from age as well as from a relatively large lake year to year is not known, because no sub- (Bellamy, Marie, or Mirror) in the nearby sequentattempts have been made to sample South French Creek drainage (Francis this ephemeral pond. The same area was Home, personalcommunication to Denton sampledby Powell's surveymany yearsago, Belk, 30 September 1991). The lakes in the but B. paludosa was not reported(cf. Pack- South French Creek drainage are now ard, 1883; Saunders, 1981b). G. D. Lang- stocked with trout and contain no fairy staff collected B. paludosa from seven sites . in the vicinity of the Great Divide Basin, Several unpublishedcollections add sup- Wyoming (now in D. Belk's personal col- port to the view that B. paludosais common lection); these sites are about 240 km north in the Medicine Bow Mountains and the of Irish Lakes. populations persist over time (Table 2). The Medicine Bow Mountains are

Table 2. Unpublished distribution records for Branchinectapaludosa in the Medicine Bow Mountains of southeasternWyoming.

Locality data Collector Status Pond near Big BrooklynLake, Albany County, 24 August 1952 J. E. Lynch USNM Acc. 294523 NE side of road several miles west of University of Wyoming Sci- J. E. Lynch USNM Acc. 294523 ence SummerCamp, 8 August 1967 >40 miles west of Laramie J. E. Lynch USNM Acc. 294523 "Jon's Pond," 10,670 ft., 3.5 km NE of Snowy Range Pass, 3 Sep- G. Langstaff D. Belk, personalcol- tember 1989 lection 186 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 13, NO. 1, 1993 thousands of kilometers from the Arctic the distribution of B. paludosa (Daborn, tundra that is typical for B. palu- 1978; Anderson, 1971). Most of the ponds dosa. Nevertheless, the temporal persis- in question lie along the westernedge of the tence of these populationsargues that south- CentralFlyway which is a major migratory ern Wyoming is a stable part of the range route for birds that nest in the Arctic and for the species. The southern limit for B. winter much farthersouth (Fig. 1). The ma- paludosa in North America is at a lower jor routes displayed in Fig. 1 were derived latitude than the limit in Europe (41? versus chiefly from the movements of waterfowl 49?). Brtek (1976) considered the perma- (Lincoln, 1939). Links to Bear Lake and nent populations in Central Europe to be in Utah are especially in- typical glacialrelicts. By analogyit could be terestingin relation to those Colorado and inferred that the southern limit in North Wyoming recordsthat are west of the Con- Americais definedby relictpopulations. In- tinental Divide. These links also hint at va- deed, Belk (1977) once speculated that the lidity for the Utah recordof Maynard(1976). populations in Utah and Arizona were It is well known that waterfowlcan serve Pleistocene glacial relicts. The distribution as agents of passive dispersalfor the eggs of of B. paludosa for North America differs various ,including anostracans substantiallyfrom that recordedfor Europe (e.g., Malone, 1965; Proctor, 1964; Proctor in terms of the degreeto which the southern et al., 1967). Colonization by B. paludosa fringe is isolated from the Arctic tundra could proceed in small jumps as dissemi- populations. The Tatra populations are ap- nules of successfulcolonists are carriedfar- parently quite isolated; whereas, the Wyo- ther and fartherfrom the Arctic by succes- ming populationsrepresent the terminus of sive migrations along the Central Flyway. a chain of recordsextending throughMon- The geographicallynarrow pattern of dis- tana and (Lynch, 1958; Hartland- tributionin southernCanada and the north- Rowe, 1965; Anderson, 1974; Daborn, ern United States is consistent with passive 1978). The presenceof populationsbetween dispersalalong the flyway, and the stochas- Wyoming and the Arctic diminishes the tic nature of extinction and recolonization likelihood that the Wyoming populations could explain the erratic temporal persis- are glacial relicts. tence of populations in Alberta. Donald (1983) studied anostracansin a It is thought that disjunct distributions temporarypond near Calgary,Alberta, for for two other Arctic anostracansare also the 14 consecutiveyears and found that B. palu- result of passive dispersal.Artemiopsis ste- dosa was not present in all years. He con- fanssoni and Polyartemiella hazeni have cluded that the observed pattern might be been recordedfrom southernAlberta (Hart- explainedby extinction and recolonization, land-Rowe and Anderson, 1968; Daborn, or by a combination of long-term egg via- 1976, 1977). Both sites are alongthe Central bility and specific physiological require- Flyway. ments that were not always met. Daborn The Central Flyway does not terminate (1978) also believed that intermittent re- in Wyoming, however. Birds migratemuch introductionwas probablyimportant for the farther south to reach winter habitats, but maintenance of populations of B. paludosa the range of B. paludosa does not extend in Alberta. farther south. The apparent absence of B. The frequency of colonization is clearly paludosa in adjacent high mountain areas important for populations subject to ex- is puzzling. Physical habitat conditions in tinction. For example, Johansen (1921) re- the Rocky Mountains of Colorado cannot ported an isolated population near Montre- be vastly different from those in southern al, Canada(45.5?N), that was collected once Wyoming, and the chemistry of the ponds and failed to appear in subsequent years. appears to differ little from that found at He speculatedthat the foundingof this pop- high elevation in the ColoradoFront Range ulationwas a chancecolonization event soon (cf. Saunders and Kling, 1990). Further- followed by extinction. more, migration routes cross the Colorado Among the various potential agents of Rockies in two places (Fig. 1). Nevertheless, passive dispersal,waterfowl and shore birds high elevation sites in Colorado almost in- are the most likely to be of significancefor variably contain B. coloradensis(Saunders, SAUNDERS ETAL.: BRANCHINECTA PALUDOSA IN WYOMING 187

Fig. 1. Distribution of Branchinecta paludosa in southern Canada and northern United States. The gray border defines the boundary of the Central Flyway. Arrows within the flyway represent major migration routes as described by Lincoln (1939). 1-Anderson, 1974; 7 sites in , Alberta. 2-Anderson, 1974; 8 sites in or near , Alberta. 3-Anderson, 1974; 2 sites in Waterton Lakes National Park, Alberta. 4-Lynch, 1958; 2 sites near Browning, Montana. 5-Saunders, 1981a; Irish Canyon, Colorado. 6- Belk collection; 7 sites in Great Divide Basin, Wyoming. 7-Present study; Linder, 1941; Horne, 1967; at least 10 sites in the Medicine Bow Mountains, Wyoming. 8-Daborn, 1978; 2 sites, Alberta. 9-Hartland-Rowe, 1965; 4 sites near Kerrobert, Saskatchewan. 10-Hartland-Rowe, 1965; 3 sites in and near Calgary. Donald, 1983; 1 site in Calgary. 11 -Hartland-Rowe, 1965; 2 sites, Alberta. 12-Hartland-Rowe, 1965; 2 sites near Hanna, Alberta. 13--Hartland-Rowe, 1965; 1 site near Consort, Alberta. 188 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 13, NO. 1, 1993

1981 a). The relationship between B. colora- Brtek,J. 1976. Anostraca,, Conchostraca densis and B. is made more in- und Calanoidader Slowakei(1. Teil).-Acta Rerum paludosa NaturaliumMusei NationalisSlovaci Bratislava22: triguing by the occurrence of both species 19-91. in the Medicine Bow Mountains. Lynch Daborn, G. R. 1976. Occurrenceof an arctic fairy (1958 and USNM Acc. No. 294523) re- shrimp Polyartemiella hazeni (Murdoch) 1884 ported B. coloradensis from ponds in the (Crustacea:Anostraca) in Alberta and Yukon Ter- same where B. occurs. ritory.-CanadianJournal of Zoology54: 2026-2028. drainages paludosa 1977. On the distributionand biology of an The absence of cooccurrences of these two arcticfairy shrimp Artemiopsis stefanssoni Johansen, species, which can inhabit adjacent ponds 1921 (Crustacea:Anostraca).-Canadian Journal of in the Medicine Bow Mountains, points to Zoology 55: 280-287. a area of Com- .1978. Distribution and biology of some potentially interesting study. nearctic tundra well be an factor. pool phyllopods.-Verhandlungen petition may important InternationaleVereinigung fur theoretischeund an- The ranges for B. paludosa and B. colora- gewandteLimnologie 20: 2442-2451. densis overlap extensively north of Colo- ,and H. F. Clifford. 1974. Physicaland chem- rado, but apparently it is very unusual to ical features of an aestival pond in western Cana- find both in the same It re- da.-Hydrobiologia 44: 43-59. species pond. Donald, D. B. 1983. Erraticoccurrence of anostra- mains a mystery as to what factor or factors cans in a temporarypool: colonization and extinc- will tip the balance to one species or the tion or adaptationto variationsin annualweather?- other. CanadianJournal of Zoology 61: 1492-1498. Branchinecta has been recorded Eng, L. L., D. Belk, and C. H. Eriksen. 1990. Cali- paludosa forniaAnostraca: and status.- in the Medicine Bow Mountains for over distribution,habitat, Journalof CrustaceanBiology 10: 247-277. 50 years and appears to be a very persistent Hartland-Rowe,R. 1965. The Anostraca and No- resident of ponds in the Snowy Range. De- tostracaof Canadawith some new distributionre- spite the temporal persistence and geo- cords.-Canadian Field-Naturalist79: 185-189. of these it , and R. S. Anderson. 1968. Arcticfairy shrimp graphical extremity populations, (Artemiopsisstefanssoni Johanson 1921) in southern is problematic that they are glacial relicts. Alberta, with a note on the genus Artemiopsis.-- The opportunity for passive dispersal from CanadianJournal of Zoology 46: 423-425. more northern sites along the Central Fly- Hore, F. 1967. Effectsof physical-chemicalfactors way provides for gene flow where a popu- on the distributionand occurrenceof some south- easternWyoming phyllopods. -Ecology 48: 472-477. lation already occurs or colonization where Houston, R. S., and others. 1978. A regionalstudy one does not. of rocksof Precambrianage in that partof the Med- icine Bow Mountains lying in SoutheasternWyo- ACKNOWLEDGEMENTS ming-with a chapter on the relationshipbetween T. Bowmanprovided information on holdingsin the Precambrianand Laramidestructure. -Geological NationalMuseum of NaturalHistory, Smithsonian In- Surveyof Wyoming,Memoir No. 1: 1-167. stitution, and commented on an earlier draft of the Johansen,F. 1921. The largerfreshwater Crustacea manuscript.R. Musselman of the Rocky Mountain from Canada and Alaska.-Canadian Field-Natu- Forestand RangeExperiment Station (U.S.D.A. Forest ralist 35: 21-30. Service)provided logistical support in connectionwith Lincoln, F. C. 1939. The migrations of American the GLEES program.George D. Langstaffprovided birds.-Doubleday, Doranand Co., New York,New collections from Wyoming to Denton Belk, and F. York. Pp. 1-189. Hore and G. Daborn provided informationon dis- Linder,F. 1932. Die Branchiopodendes arktischen tribution. Gebietes.-Fauna Arctica 6: 183-204. 1941. Contributionsto the morphologyand LITERATURECITED of the BranchiopodaAnostraca.- Zoolo- fran 20: 101-302. Anderson, R. S. 1971. Crustaceanplankton of 146 giske Bidrag Uppsala alpine and subalpine lakes and ponds in western Lynch, J. E. 1958. Branchinectacornigera, a new Canada.-Journal of the Fisheries ResearchBoard species of crustaceanphyllopod from the state of of Canada28: 311-321. Washington.-Proceedingsof the United StatesNa- 1974. Crustaceanplankton communities of tional Museum 108: 25-37. 340 lakes and ponds in and near the National Parks Malone, C. R. 1965. Dispersal of plankton:rate of of the CanadianRocky Mountains.-Journal of the food passagein mallardducks.-Journal of Wildlife FisheriesResearch Board of Canada31: 855-869. Management29: 529-532. Belk, D. 1975. Key to the Anostraca(fairy shrimp) Maynard,S.S. 1976. Branchinectapaludosa (Miiller) of North America.-SouthwesternNaturalist 20: 90- (Crustacea:Anostraca) in northernUtah with some 103. notes on its ecology.-Southwestern Naturalist 20: . 1977. Zoogeographyof the Arizona fairy 582-585. shrimps(Crustacea: Anostraca).- Journalof the Ar- Packard,A. S. 1883. Monographof phyllopodCrus- izona Academy of Science 12: 70-78. tacea.-Twelfth AnnualReport of the United States SAUNDERS ETAL.: BRANCHINECTA PALUDOSA IN WYOMING 189

Geological and GeographicalSurvey of the Terri- valvia) in the Colorado Front Range: the role of tories, Part I: 295-592. abiotic factors.-Freshwater Biology 24: 275-285. Proctor, V. W. 1964. Viability of crustaceaneggs Vekhoff,N. V. 1990. Faunaand distributionof fairy recoveredfrom ducks.-Ecology 45: 656-658. shrimps(Anostraca) and tadpoleshrimps (Notostra- ,C. R. Malone, and V. L. DeVlaming. 1967. ca) in water reservoirsof northerncircumpolar re- Dispersalof aquaticorganisms: viability of dissem- gion.-Proceedings of the SiberianDivision of Sci- inules recoveredfrom the intestinaltract of captive ence USSR, Section of BiologicalSciences. Issue 3: kill deer.-Ecology 48: 672-676. 71-77 (Division Reprint),Novosibirsk, Publishing Reed, E. B. 1963. Records of freshwaterCrustacea House "Nauka,"Siberian Division. from arctic and subarcticCanada.-Bulletin of the National Museum of Canada 199: 29-62. RECEIVED:25 January 1992. Saunders,J. F., III. 1981a. Eubranchiopodaof Col- ACCEPTED:9 April 1992. orado. Part 2. Anostraca.-Natural History Inven- tory of Colorado6: 1-23. Addresses:(JFS and RD) Centerfor Limnology,De- 198 b. A redescriptionof Lepidurusbilo- partmentof Environmental,Population and Organis- batus Packard (Crustacea:Notostraca).-Transac- mic Biology, University of Colorado, Boulder, Colo- tions of the AmericanMicroscopical Society 99: 179- rado 80309-0334, U.S.A.; (DB) Biology Department, 186. Our Lady of the Lake University, 411 SW 24th St., ,and G. W. Kling. 1990. Speciesdistributions San Antonio, Texas 78207-4666, U.S.A. and shell characteristicsof Pisidium (Mollusca:Bi-