The same invertebrate fauna of 17 species was found in Devils Den Cave, Washington County, Arkansas, in 1969 and 1979. The fauna consists of 1 trogloxene, 14 troglophiles, and 2 troglobites, a spider, Porrhomma cavernicolum, and a collembolan, Pseudosinella dubia. Devils Den Cave has a well developed cavernicolous fauna, although it is in sandstone which generally supports a poor cave adapted fauna. The troglobites probably evolved in the vicinity of northwestern Arkansas in limestone caves or in deep forest soils of the Ozark region. They then dispersed overland, perhaps as recently as the late Wisconsinan, to occupy this sandstone cave

Peck, James H.

Peck, Stewart B.

English

UARK (University of Arkansas )

Journal of the Arkansas Academy of Science Volume 36 Article 15 1982 Invertebrate Fauna of Devils Den, a Sandstone Cave in Northwestern Arkansas Stewart B. Peck Carleton University James H. Peck University of Arkansas at Little Rock Follow this and additional works at: https://scholarworks.uark.edu/jaas  Part of the Terrestrial and Aquatic Ecology Commons, and the Zoology Commons Recommended Citation Peck, Stewart B. and Peck, James H. (1982) "Invertebrate Fauna of Devils Den, a Sandstone Cave in Northwestern Arkansas," Journal of the Arkansas Academy of Science: Vol. 36 , Article 15. Available at: https://scholarworks.uark.edu/jaas/vol36/iss1/15 This article is available for use under the Creative Commons license: Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0). Users are able to read, download, copy, print, distribute, search, link to the full texts of these articles, or use them for any other lawful purpose, without asking prior permission from the publisher or the author. This Article is brought to you for free and open access by ScholarWorks@UARK. It has been accepted for inclusion in Journal of the Arkansas Academy of Science by an authorized editor of ScholarWorks@UARK. For more information, please contact scholar@uark.edu. Arkansas Academy of Science Proceedings, Vol. XXXVI,198246INVERTEBRATE FAUNA OF DEVILSDEN,A SANDSTONE CAVE INNORTHWESTERN ARKANSASSTEWART B.PECKDepartment of BiologyCarleton UniversityOttawa, Ontario K1S 5B6, CanadaJAMES H.PECKDepartment of BiologyUniversity of Arkansas at Little RockLittle Rock, Arkansas 72204ABSTRACTThe same invertebrate fauna of 17 species was found in Devils Den Cave, Washington County,Arkansas, in 1969 and 1979. The fauna consists of 1 trogloxene, 14 troglophiles, and 2 troglobites, aspider, Porrhomma cavernicolum, and a collembolan, Pseudosinella dubia. Devils Den Cave has awell developed cavernicolous fauna, although it is in sandstone which generally supports a poor caveadapted fauna. The troglobites probably evolved in the vicinityof northwestern Arkansas in limestonecaves or in deep forest soils of the Ozark region. They then dispersed overland, perhaps as recentlyas the late Wisconsinan, to occupy this sandstone cave.INTRODUCTIONSurveys ofthe taxonomic composition ofthe invertebrate faunas ofNorthAmerican caves have progressed considerably since the checklistby Nicholas (1960) in which few records from Arkansas caves werereported. Considerable survey work has since been conducted inNorthAmerica (see Peck and Lewis, 1977, for a review), including Arkansas(McDaniel and Smith, 1976; McDaniel et al., 1979; Youngsteadt andYoungsteadt, 1978). These reports concerned faunas oflimestone caves,which generally received more attention than sandstone caves inArkansas and eastern North America. Arkansas has numerous sand-stone features approximating caves, such as the sandstone crevice cavesof Devils Den State Park, Washington County, in northwesternArkansas. The report ofa troglobitic collembolan from a cave inthepark (Christiansen, 1960) suggested to us that these sandstone cavesrequired a survey to assess their cavernicolous faunas.Devils Den State Park contains six crevice caves in Pennsylvaniansandstone (Harvey et al., 1981). These caves were formed byslippageofsandstone blocks. Widely spaced joints between blocks opened asthe blocks slipped on underlyingbeds of shales, which were probablylubricated by seepage water. The best developed cave is named DevilsDen, from which the park receives its name and is a relatively straightfracture-opening penetrating the hillside approximately 180 m and havingonlya small entrance. We report the results of surveys conducted in1969 and 1979 to assess the invertebrate fauna ofDevils Den Cave.METHODSSurveys were conducted in May of 1969 and 1979, as part ofa researchprogram on the evolution and distribution of cave-inhabiting in-vertebrates. Organisms were obtained by hand collection and at baitsof human dung or carrion suspended above pit traps withpreservativesolutions. A faunal list was prepared and annotated with status, lifehistory, or distributional information. The probable ecological adap-tation of each species is given, following the criteria developed by Barr(1963) and used by McDaniel and Smith (1976).ANNOTATEDFAUNAL LISTPHYLUMARTHROPODAClass ArachnidaOrder AcarinaFamily RhagidiidaeRhagida sp. Troglophile. Species of this and related genera of mitesare often found incool and wet habitats, such as caves. Elliottand Strandtmann (1971) report R. longisensilla Shiba fromDiamond Cave, Newton Co., AR. Known from Japan andAlaska. Other species undoubtably occur in Arkansas, since R.hilli Strandtmann, R. weyerensis (Packard) and R. whartoniStrandtmann are found in Missouri caves (J. L. Craig, 1977,and pers. comm.) Zacharda (1980) has revised the nomenclatureof this family.Family EupodidaeLinopodes sp. Troglophile. Predatory mites occasionally found incaves.Order AraneaFamily AraneidaeMeta menardi (Latreille).Troglophile. Widespread incaves and similarhabitats throughout much ofeastern United States. McDanielet al. (1979) report this spider from Needles Cave, Izard Co.Family LinyphiidaePorrhomma cavernicolum (Keyserling). Troglobite. A small, pale,eyeless species, widespread inthe Appalachian and Interior LowPlateau regions from Virginia to Oklahoma, but known onlyfrom caves. This population is well established, since 4 males,7 females, and 4 immatures were observed. McDaniel et al. (1979)report this species from Davis Pit, Searcy Co.Order OpilionesFamily IschyropsalidaeSabacon cavicolens (Packard). Troglophile. Immatures only. They maynotbe this species, which is widespread inforest habitats in theeastern U.S. (Shear, 1975). Reported from Roasting Ear Cave,Stone Co. (McDaniel et al., 1979).Family Phalangodidae46Journal of the Arkansas Academy of Science, Vol. 36 [1982], Art. 15Published by Arkansas Academy of Science, 1982Arkansas Academy of Science Proceedings, Vol. XXXVI,1982 47AStewart P. Peck and James H. PeckCrosbyella spinturnix (Crosby and Bishop). Troglophile. Reported inother Arkansas caves by Goodnight and Goodnight (1942),Youngsteadt and Youngsteadt (1978), and McDaniel et al. (1979).Class DiplopodaOrder ChordeumidaFamily ConotylidaeTrigenotyla parca Causey. Troglophile. Phylogenetically close to thecave restricted Scoterpes of caves throughout Ozark and Ap-palachian regions. Amonotypic genus restricted tonorthwesternArkansas (Shear, 1972).Class CollembolaOrder EntomobryomorphaFamily EntomobryidaePseudosinella dubia Christiansen. Troglobite. Known only from 4limestone and sandstone caves in Washington Co. (Christiansen,1960; Christiansen and Bellinger, 1980). Christiansen (pers.comm.) also records the collembolans Folsomia Candida Willem.,Tullbergia iowensis Mills, and Onychiurus pseudofimetarius(Absolon) from the cave in the 1950's.Class InsectaOrder OrthopteraFamily RhaphidophoridaeCeuthophilus sp. Trogloxene. Immature cave crickets were on theceiling, just inside the cave entrance.Order ColeopteraFamily CarabidaePlatynus tenuicollis LeConte. Troglophile. Widelydistributed incavesof the southeastern U. S. and Ozarks.Family LeiodidaePtomaphagus shapardi Sanderson. Troglophile. Found most often incaves, but also occurs in forest litter in Arkansas and Oklahoma.Its small eyes and winglessness suggest adaptation fordeep soilhabitats (Peck, 1973).Family StaphylinidaeAthetasp. Troglophile? Small, poorly known aleocharines. Abundantin caves.Order DipteraFamily SphaeroceridaeLeptocera sp. Troglophile. Common, scavenging flies found in cavesof the southeastern U. S.Family MycetophilidaeMacrocera nobilis Johnson. Troglophile. Larvae were observed inrunway-like webs under rock ledges (Peck and Russell, 1976).Adult fungus gnats were present, but avoided capture.Family TipulidaeUndetermined genus. Trogloxene. Observed on ceiling, just inside caveentrance.Family SciaridaeBradysia sp. Troglophile. Common, scavenging flies in caves throughoutsoutheastern U.S.Family PsychodidaeUndetermined genus. Troglophile. Scavenging moth flies of cavesthroughout southeastern U.S.continued exhaustive surveys ofArkansas caves. The relatively smallsandstone cave we surveyed, then, has an average number (Barnett, 1967)in its fauna. Biospeleologists tend to ignore caves similar toDevils Den.Recognition ofa significant cavernicolous fauna, includingtroglobites,froma sandstone cave suggests that additional surveys ofsimilar cavesare necessary.Terrestrial troglobites probably descended from forest litter-dwellingancestors which became genetically isolated in caves in conjunction withPleistocene climatic changes (Barr, 1968, 1973). After speciation, varioussubterranean avenues have been used for dispersion to other caves.However, deep, subterranean terrestrial dispersal from the Mississip-pian age limestone caves ofnorthwestern Arkansas through insoluableshales and sandstones into the Pennsylvanian age sandstones of the cavesat Devils Den State Park is unlikely.Amore likelyavenue of dispersalto these caves could have been overland within surficial litter, humus,and soil. Some troglobites have been suggested as being capable ofusing such a dispersal avenue ineastern North America (Peck, 1973,1978, 1980, 1981; Peck and Lewis, 1977) and inEurope (Juberthie etal., 1980). Inthe latter case, troglobites were found in deep soils ontopofnon-calcareous bedrock, precluding a subterranean dispersion.The occurrence of troglobites at DevilsDen State Park supports a sur-ficial,overland dispersal avenue, rather than the deep, subterraneandispersal avenue suggested by Barr (1968, 1973).Time and conditions ofoverland dispersal to Devils Den State Parkcannot be precisely pinpointed, since little is known concerning Ozarkforest litter invertebrates and their distributional and habitat re-quirements. It is reasonable to conclude that species favoring cool,moisthabitats would have found such habitats to be more prevalent inArkansas forests during the fullglacial climatic conditions which oc-curred most recently during the Wisconsinan glacialperiod from 80,000to 15,000 years before present (Watts, 1980). Inthe western Ozarks,King(1973) found the dominant vegetation tobe boreal spruce forestfrom at least 20,000 to 13,500 years before present. Such a forest mayhave provided conditions suitable foroverland dispersal by troglobitesfromMissouri or Arkansas limestone cave areas through moss and deeplitter into the vicinityof Devils Den State Park. Subsequent isolationof troglobites at Devils Den State Park represents an example ofclimaticallyinduced vicariance distributional pattern. Whether disper-sal to these caves occurred in earlier glacial epochs or during severalepochs can not be assessed. Additional surveys of the litter invertebratesnear cave entrances and in forest habitats between caves might clarifycomponents of the overland dispersal hypothesis.ACKNOWLEDGMENTSDrs. W. J. Gertsch, W. A.Shear, and K.Christiansenare thanked for providing determinations of spiders,millipeds, and collembolans, respectively. Drs. Shear,Christiansen, and V. R. McDaniel are thanked for review-ing the manuscript. Field work was supported byoperating grants from Canadian Natural Sciences andEngineering Research Council to S.B.P.DISCUSSIONSeventeen species were found to inhabit Devils Den Cave in 1969 and1979. Congruence of the faunas over the 10 year period suggests thatthese organisms are regular inhabitants of the cave and notaccidentaloccupants. The fauna consists of 1 trogloxene, 14 troglophiles, and 2troglobites. The trogloxene and troglophiles are not cave limited, anddo not reveal much about the dynamics of faunal dispersal, cave oc-cupation, and evolution. However, the troglobites (obligate cavemicoles)are significant in this respect. Arkansas limestone caves are occupiedby an average of 2 troglobitic invertebrates per cave (Barnett, 1970)based mainly on aquatic species. McDaniel and Gardner (1977)suggest that the average number is actually higher, but will requireLITERATURE CITEDBARNETT, D. E. 1970. An ecological investigation of cavernicolepopulations in Mansell cave, Randolph County, Arkansas. M.S.Thesis, Northw. Louisiana St. Univ., Natchitoches, LA 51p.BARR, T.C. 1963. Ecological classification of cavemicoles. Cave Notes5(2):9-12.BARR.T. C, JR. 1968. Cave ecology and the evolution of troglobites.Evol. Biol. 2:35-102.47Journal of the Arkansas Academy of Science, Vol. 36 [1982], Art. 15https://scholarworks.uark.edu/jaas/vol36/iss1/1548Arkansas Academy of Science Proceedings, Vol. XXXVI,1982InvGrtGbratG rauna of Uevils Den, A bandstone Uavs in Nortnwsstern ArKansasBARR, T.C., JR. 1973. Refugees of the Ice Age. Nat. Hist. 82:26-35,72-73.CHRISTIANSEN, K. 1960. The genus Pseudosinella (Collembola,Entomobryidae) in caves of the United States. Psyche 67:1-25.CHRISTIANSEN, K., and P. BELLINGER. 1980. The Collembolaof North America north of the Rio Grande. Part 3, FamilyEntomobryidae. Grinnell College, Grinnell, Iowa. pp. 785-1042.CRAIG, J. L. 1977. Invertebrate faunas ofcaves to be inundated bythe Meramec Park Lake ineastern Missouri. Bull.Natl. Speleol.Soc. 39:80-89.ELLIOTT,W. R., and R. W. STRANDTMANN. 1971. New localityrecords for Rhagidia from Mexican and American caves. J. Kan.Entomol. Soc. 44:468-475.GOODNIGHT, J., and M. L. GOODNIGHT. 1942. NewPhalangodidae (Phalangida) from the United States. Amer. Mus.Novitates No. 1188, 18p.HARVEY, M. J., J. J. CASSIDY, and G. G. O'HAGAN. 1981.Endangered bats of Arkansas: distribution, status, ecology, andmanagement. Ecological Research Center, Dept. Biology,Memphis St. Univ., Memphis, TN. 137p.JUBERTHIE, C, B. DELAY, and M. BOUILLON. 1980. Surl'existence d'un milieu souterrain superficiel en zone non calcaire.C.R. Acad. Sci. Paris 290 (ser. D):49-52.KING, J. E. 1973. Late Pleistocene palynology and biogeography ofthe western Missouri Ozarks. Ecol. Mono. 43:539-465.MCDANIEL,V. R., and K. L. SMITH. 1976. Cave fauna ofArkansas: selected invertebrate taxa. Proc. Ark. Acad. Sci.30:57-60.MCDANIEL,V. R., and J. E. GARDNER. 1977. Cave fauna ofArkansas: vertebrate taxa. Proc. Ark.Acad. Sci. 31:68-71.MCDANIEL,V.R., K. N. PAIGE,and C. R. TUMLISON.1979. Cavefauna of Arkansas: additional invertebrate and vertebraterecords. Proc. Ark. Acad. Sci. 33:83-84.NICHOLAS, BRO. G. 1960. Checklist of macroscopic troglobiticorganisms of the United States. Amer. Midi. Nat. 64:123-160.PECK, S. B. 1973. A systematic revision and the evolutionary biologyof the Ptomaphagus (Adelops) beetles of North America(Coleoptera: Leiodidae: Catopinae), with emphasis on cave-inhabiting species. Bull. Mus. Comp. Zool. 145:29-162.PECK, S. B. 1978. New montane Ptomaphagus beetles from NewMexico and zoogeography of southwestern caves (Coleoptera:Leiodidae: Catopinae). Southwestern Natur. 23:227-238.PECK, S. B. 1980. Climatic change and the evolution of caveinvertebrates in the Grand Canyon, Arizona. Natl. Speleol. Soc.Bull. 42:53-60.PECK, S. B. 1981. The geological, geographical, and environmentalsetting ofcave faunal evolution, pp.501-52, in B.F.Beck, Proc.Eighth Inter. Congr. Speleology, Dept. Geol., Georgia Southw.Coll., Americus, GA.PECK, S. B., and J. J. LEWIS. 1977. Zoogeography and evolutionof the subterranean invertebrate faunas of Illinois andsoutheastern Missouri. Natl. Speleol. Soc. Bull. 40:39-63.PECK, S. B., and D.R. RUSSELL. 1976. Life history of the fungusgnat Macrocera nobilis in American caves (Diptera:Mycetophilidae). Can. Ent. 108:1235-1241.SHEAR, W. A. 1972. Studies in the milliped order Chordeumida(Diplopoda): A revision of the family Cleidogonidae and areclassification of the order Chordeumida in the New World.Bull. Mus. Comp. Zool. 144:151-352.SHEAR, W. A.1975. The opilionidgenera Sabacon and Tomicomerusin America (Opiliones, Troguloidae, Ischyropsalidae). J.Arachnol. 3:5-29.WATTS., W. A. 1980. The late quaternary vegetation history of thesoutheastern United States. Ann. Rev. Ecol. Systs. 11:387-409.YOUNGSTEADT, N. W., and J.O. YOUNGSTEADT. 1978. Asurveyof some cave invertebrates fromNorthern Arkansas. Assoc. Ark.Cave Studies 1:1-13. (Box 682, Harding College, Searcy, AR72143).ZACHARDA, M. 1980. Soil mites of the family Rhagidiidae(Actinedida: Eupodoidea). Morphology, Systematics, Ecology.Acta Universitatis Carolinae (Prague), Biologica 1978(5-6):489-785.48Journal of the Arkansas Academy of Science, Vol. 36 [1982], Art. 15Published by Arkansas Academy of Science, 1982

Invertebrate Fauna of Devils Den, a Sandstone Cave in Northwestern Arkansas

ScholarWorks@UARK

Journal of the Arkansas Academy of ScienceVolume 36 Article 151982Invertebrate Fauna of Devils Den, a SandstoneCave in Northwestern ArkansasStewart B. PeckCarleton UniversityJames H. PeckUniversity of Arkansas at Little RockFollow this and additional works at: http://scholarworks.uark.edu/jaasPart of the Terrestrial and Aquatic Ecology Commons, and the Zoology CommonsThis article is available for use under the Creative Commons license: Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0). Users are able toread, download, copy, print, distribute, search, link to the full texts of these articles, or use them for any other lawful purpose, without asking priorpermission from the publisher or the author.This Article is brought to you for free and open access by ScholarWorks@UARK. It has been accepted for inclusion in Journal of the Arkansas Academyof Science by an authorized editor of ScholarWorks@UARK. For more information, please contact scholar@uark.edu, ccmiddle@uark.edu.Recommended CitationPeck, Stewart B. and Peck, James H. (1982) "Invertebrate Fauna of Devils Den, a Sandstone Cave in Northwestern Arkansas," Journalof the Arkansas Academy of Science: Vol. 36 , Article 15.Available at: http://scholarworks.uark.edu/jaas/vol36/iss1/15Arkansas Academy of Science Proceedings, Vol. XXXVI,198246INVERTEBRATE FAUNA OF DEVILSDEN,A SANDSTONE CAVE INNORTHWESTERN ARKANSASSTEWART B.PECKDepartment of BiologyCarleton UniversityOttawa, Ontario K1S 5B6, CanadaJAMES H.PECKDepartment of BiologyUniversity of Arkansas at Little RockLittle Rock, Arkansas 72204ABSTRACTThe same invertebrate fauna of 17 species was found in Devils Den Cave, Washington County,Arkansas, in 1969 and 1979. The fauna consists of 1 trogloxene, 14 troglophiles, and 2 troglobites, aspider, Porrhomma cavernicolum, and a collembolan, Pseudosinella dubia. Devils Den Cave has awell developed cavernicolous fauna, although it is in sandstone which generally supports a poor caveadapted fauna. The troglobites probably evolved in the vicinityof northwestern Arkansas in limestonecaves or in deep forest soils of the Ozark region. They then dispersed overland, perhaps as recentlyas the late Wisconsinan, to occupy this sandstone cave.INTRODUCTIONSurveys ofthe taxonomic composition ofthe invertebrate faunas ofNorthAmerican caves have progressed considerably since the checklistby Nicholas (1960) in which few records from Arkansas caves werereported. Considerable survey work has since been conducted inNorthAmerica (see Peck and Lewis, 1977, for a review), including Arkansas(McDaniel and Smith, 1976; McDaniel et al., 1979; Youngsteadt andYoungsteadt, 1978). These reports concerned faunas oflimestone caves,which generally received more attention than sandstone caves inArkansas and eastern North America. Arkansas has numerous sand-stone features approximating caves, such as the sandstone crevice cavesof Devils Den State Park, Washington County, in northwesternArkansas. The report ofa troglobitic collembolan from a cave inthepark (Christiansen, 1960) suggested to us that these sandstone cavesrequired a survey to assess their cavernicolous faunas.Devils Den State Park contains six crevice caves in Pennsylvaniansandstone (Harvey et al., 1981). These caves were formed byslippageofsandstone blocks. Widely spaced joints between blocks opened asthe blocks slipped on underlyingbeds of shales, which were probablylubricated by seepage water. The best developed cave is named DevilsDen, from which the park receives its name and is a relatively straightfracture-opening penetrating the hillside approximately 180 m and havingonlya small entrance. We report the results of surveys conducted in1969 and 1979 to assess the invertebrate fauna ofDevils Den Cave.METHODSSurveys were conducted in May of 1969 and 1979, as part ofa researchprogram on the evolution and distribution of cave-inhabiting in-vertebrates. Organisms were obtained by hand collection and at baitsof human dung or carrion suspended above pit traps withpreservativesolutions. A faunal list was prepared and annotated with status, lifehistory, or distributional information. The probable ecological adap-tation of each species is given, following the criteria developed by Barr(1963) and used by McDaniel and Smith (1976).ANNOTATEDFAUNAL LISTPHYLUMARTHROPODAClass ArachnidaOrder AcarinaFamily RhagidiidaeRhagida sp. Troglophile. Species of this and related genera of mitesare often found incool and wet habitats, such as caves. Elliottand Strandtmann (1971) report R. longisensilla Shiba fromDiamond Cave, Newton Co., AR. Known from Japan andAlaska. Other species undoubtably occur in Arkansas, since R.hilli Strandtmann, R. weyerensis (Packard) and R. whartoniStrandtmann are found in Missouri caves (J. L. Craig, 1977,and pers. comm.) Zacharda (1980) has revised the nomenclatureof this family.Family EupodidaeLinopodes sp. Troglophile. Predatory mites occasionally found incaves.Order AraneaFamily AraneidaeMeta menardi (Latreille).Troglophile. Widespread incaves and similarhabitats throughout much ofeastern United States. McDanielet al. (1979) report this spider from Needles Cave, Izard Co.Family LinyphiidaePorrhomma cavernicolum (Keyserling). Troglobite. A small, pale,eyeless species, widespread inthe Appalachian and Interior LowPlateau regions from Virginia to Oklahoma, but known onlyfrom caves. This population is well established, since 4 males,7 females, and 4 immatures were observed. McDaniel et al. (1979)report this species from Davis Pit, Searcy Co.Order OpilionesFamily IschyropsalidaeSabacon cavicolens (Packard). Troglophile. Immatures only. They maynotbe this species, which is widespread inforest habitats in theeastern U.S. (Shear, 1975). Reported from Roasting Ear Cave,Stone Co. (McDaniel et al., 1979).Family Phalangodidae46Journal of the Arkansas Academy of Science, Vol. 36 [1982], Art. 15Published by Arkansas Academy of Science, 1982Arkansas Academy of Science Proceedings, Vol. XXXVI,1982 47AStewart P. Peck and James H. PeckCrosbyella spinturnix (Crosby and Bishop). Troglophile. Reported inother Arkansas caves by Goodnight and Goodnight (1942),Youngsteadt and Youngsteadt (1978), and McDaniel et al. (1979).Class DiplopodaOrder ChordeumidaFamily ConotylidaeTrigenotyla parca Causey. Troglophile. Phylogenetically close to thecave restricted Scoterpes of caves throughout Ozark and Ap-palachian regions. Amonotypic genus restricted tonorthwesternArkansas (Shear, 1972).Class CollembolaOrder EntomobryomorphaFamily EntomobryidaePseudosinella dubia Christiansen. Troglobite. Known only from 4limestone and sandstone caves in Washington Co. (Christiansen,1960; Christiansen and Bellinger, 1980). Christiansen (pers.comm.) also records the collembolans Folsomia Candida Willem.,Tullbergia iowensis Mills, and Onychiurus pseudofimetarius(Absolon) from the cave in the 1950's.Class InsectaOrder OrthopteraFamily RhaphidophoridaeCeuthophilus sp. Trogloxene. Immature cave crickets were on theceiling, just inside the cave entrance.Order ColeopteraFamily CarabidaePlatynus tenuicollis LeConte. Troglophile. Widelydistributed incavesof the southeastern U. S. and Ozarks.Family LeiodidaePtomaphagus shapardi Sanderson. Troglophile. Found most often incaves, but also occurs in forest litter in Arkansas and Oklahoma.Its small eyes and winglessness suggest adaptation fordeep soilhabitats (Peck, 1973).Family StaphylinidaeAthetasp. Troglophile? Small, poorly known aleocharines. Abundantin caves.Order DipteraFamily SphaeroceridaeLeptocera sp. Troglophile. Common, scavenging flies found in cavesof the southeastern U. S.Family MycetophilidaeMacrocera nobilis Johnson. Troglophile. Larvae were observed inrunway-like webs under rock ledges (Peck and Russell, 1976).Adult fungus gnats were present, but avoided capture.Family TipulidaeUndetermined genus. Trogloxene. Observed on ceiling, just inside caveentrance.Family SciaridaeBradysia sp. Troglophile. Common, scavenging flies in caves throughoutsoutheastern U.S.Family PsychodidaeUndetermined genus. Troglophile. Scavenging moth flies of cavesthroughout southeastern U.S.continued exhaustive surveys ofArkansas caves. The relatively smallsandstone cave we surveyed, then, has an average number (Barnett, 1967)in its fauna. Biospeleologists tend to ignore caves similar toDevils Den.Recognition ofa significant cavernicolous fauna, includingtroglobites,froma sandstone cave suggests that additional surveys ofsimilar cavesare necessary.Terrestrial troglobites probably descended from forest litter-dwellingancestors which became genetically isolated in caves in conjunction withPleistocene climatic changes (Barr, 1968, 1973). After speciation, varioussubterranean avenues have been used for dispersion to other caves.However, deep, subterranean terrestrial dispersal from the Mississip-pian age limestone caves ofnorthwestern Arkansas through insoluableshales and sandstones into the Pennsylvanian age sandstones of the cavesat Devils Den State Park is unlikely.Amore likelyavenue of dispersalto these caves could have been overland within surficial litter, humus,and soil. Some troglobites have been suggested as being capable ofusing such a dispersal avenue ineastern North America (Peck, 1973,1978, 1980, 1981; Peck and Lewis, 1977) and inEurope (Juberthie etal., 1980). Inthe latter case, troglobites were found in deep soils ontopofnon-calcareous bedrock, precluding a subterranean dispersion.The occurrence of troglobites at DevilsDen State Park supports a sur-ficial,overland dispersal avenue, rather than the deep, subterraneandispersal avenue suggested by Barr (1968, 1973).Time and conditions ofoverland dispersal to Devils Den State Parkcannot be precisely pinpointed, since little is known concerning Ozarkforest litter invertebrates and their distributional and habitat re-quirements. It is reasonable to conclude that species favoring cool,moisthabitats would have found such habitats to be more prevalent inArkansas forests during the fullglacial climatic conditions which oc-curred most recently during the Wisconsinan glacialperiod from 80,000to 15,000 years before present (Watts, 1980). Inthe western Ozarks,King(1973) found the dominant vegetation tobe boreal spruce forestfrom at least 20,000 to 13,500 years before present. Such a forest mayhave provided conditions suitable foroverland dispersal by troglobitesfromMissouri or Arkansas limestone cave areas through moss and deeplitter into the vicinityof Devils Den State Park. Subsequent isolationof troglobites at Devils Den State Park represents an example ofclimaticallyinduced vicariance distributional pattern. Whether disper-sal to these caves occurred in earlier glacial epochs or during severalepochs can not be assessed. Additional surveys of the litter invertebratesnear cave entrances and in forest habitats between caves might clarifycomponents of the overland dispersal hypothesis.ACKNOWLEDGMENTSDrs. W. J. Gertsch, W. A.Shear, and K.Christiansenare thanked for providing determinations of spiders,millipeds, and collembolans, respectively. Drs. Shear,Christiansen, and V. R. McDaniel are thanked for review-ing the manuscript. Field work was supported byoperating grants from Canadian Natural Sciences andEngineering Research Council to S.B.P.DISCUSSIONSeventeen species were found to inhabit Devils Den Cave in 1969 and1979. Congruence of the faunas over the 10 year period suggests thatthese organisms are regular inhabitants of the cave and notaccidentaloccupants. The fauna consists of 1 trogloxene, 14 troglophiles, and 2troglobites. The trogloxene and troglophiles are not cave limited, anddo not reveal much about the dynamics of faunal dispersal, cave oc-cupation, and evolution. However, the troglobites (obligate cavemicoles)are significant in this respect. Arkansas limestone caves are occupiedby an average of 2 troglobitic invertebrates per cave (Barnett, 1970)based mainly on aquatic species. McDaniel and Gardner (1977)suggest that the average number is actually higher, but will requireLITERATURE CITEDBARNETT, D. E. 1970. An ecological investigation of cavernicolepopulations in Mansell cave, Randolph County, Arkansas. M.S.Thesis, Northw. Louisiana St. Univ., Natchitoches, LA 51p.BARR, T.C. 1963. Ecological classification of cavemicoles. Cave Notes5(2):9-12.BARR.T. C, JR. 1968. Cave ecology and the evolution of troglobites.Evol. Biol. 2:35-102.47Journal of the Arkansas Academy of Science, Vol. 36 [1982], Art. 15http://scholarworks.uark.edu/jaas/vol36/iss1/1548Arkansas Academy of Science Proceedings, Vol. XXXVI,1982InvGrtGbratG rauna of Uevils Den, A bandstone Uavs in Nortnwsstern ArKansasBARR, T.C., JR. 1973. Refugees of the Ice Age. Nat. Hist. 82:26-35,72-73.CHRISTIANSEN, K. 1960. The genus Pseudosinella (Collembola,Entomobryidae) in caves of the United States. Psyche 67:1-25.CHRISTIANSEN, K., and P. BELLINGER. 1980. The Collembolaof North America north of the Rio Grande. Part 3, FamilyEntomobryidae. Grinnell College, Grinnell, Iowa. pp. 785-1042.CRAIG, J. L. 1977. Invertebrate faunas ofcaves to be inundated bythe Meramec Park Lake ineastern Missouri. Bull.Natl. Speleol.Soc. 39:80-89.ELLIOTT,W. R., and R. W. STRANDTMANN. 1971. New localityrecords for Rhagidia from Mexican and American caves. J. Kan.Entomol. Soc. 44:468-475.GOODNIGHT, J., and M. L. GOODNIGHT. 1942. NewPhalangodidae (Phalangida) from the United States. Amer. Mus.Novitates No. 1188, 18p.HARVEY, M. J., J. J. CASSIDY, and G. G. O'HAGAN. 1981.Endangered bats of Arkansas: distribution, status, ecology, andmanagement. Ecological Research Center, Dept. Biology,Memphis St. Univ., Memphis, TN. 137p.JUBERTHIE, C, B. DELAY, and M. BOUILLON. 1980. Surl'existence d'un milieu souterrain superficiel en zone non calcaire.C.R. Acad. Sci. Paris 290 (ser. D):49-52.KING, J. E. 1973. Late Pleistocene palynology and biogeography ofthe western Missouri Ozarks. Ecol. Mono. 43:539-465.MCDANIEL,V. R., and K. L. SMITH. 1976. Cave fauna ofArkansas: selected invertebrate taxa. Proc. Ark. Acad. Sci.30:57-60.MCDANIEL,V. R., and J. E. GARDNER. 1977. Cave fauna ofArkansas: vertebrate taxa. Proc. Ark.Acad. Sci. 31:68-71.MCDANIEL,V.R., K. N. PAIGE,and C. R. TUMLISON.1979. Cavefauna of Arkansas: additional invertebrate and vertebraterecords. Proc. Ark. Acad. Sci. 33:83-84.NICHOLAS, BRO. G. 1960. Checklist of macroscopic troglobiticorganisms of the United States. Amer. Midi. Nat. 64:123-160.PECK, S. B. 1973. A systematic revision and the evolutionary biologyof the Ptomaphagus (Adelops) beetles of North America(Coleoptera: Leiodidae: Catopinae), with emphasis on cave-inhabiting species. Bull. Mus. Comp. Zool. 145:29-162.PECK, S. B. 1978. New montane Ptomaphagus beetles from NewMexico and zoogeography of southwestern caves (Coleoptera:Leiodidae: Catopinae). Southwestern Natur. 23:227-238.PECK, S. B. 1980. Climatic change and the evolution of caveinvertebrates in the Grand Canyon, Arizona. Natl. Speleol. Soc.Bull. 42:53-60.PECK, S. B. 1981. The geological, geographical, and environmentalsetting ofcave faunal evolution, pp.501-52, in B.F.Beck, Proc.Eighth Inter. Congr. Speleology, Dept. Geol., Georgia Southw.Coll., Americus, GA.PECK, S. B., and J. J. LEWIS. 1977. Zoogeography and evolutionof the subterranean invertebrate faunas of Illinois andsoutheastern Missouri. Natl. Speleol. Soc. Bull. 40:39-63.PECK, S. B., and D.R. RUSSELL. 1976. Life history of the fungusgnat Macrocera nobilis in American caves (Diptera:Mycetophilidae). Can. Ent. 108:1235-1241.SHEAR, W. A. 1972. Studies in the milliped order Chordeumida(Diplopoda): A revision of the family Cleidogonidae and areclassification of the order Chordeumida in the New World.Bull. Mus. Comp. Zool. 144:151-352.SHEAR, W. A.1975. The opilionidgenera Sabacon and Tomicomerusin America (Opiliones, Troguloidae, Ischyropsalidae). J.Arachnol. 3:5-29.WATTS., W. A. 1980. The late quaternary vegetation history of thesoutheastern United States. Ann. Rev. Ecol. Systs. 11:387-409.YOUNGSTEADT, N. W., and J.O. YOUNGSTEADT. 1978. Asurveyof some cave invertebrates fromNorthern Arkansas. Assoc. Ark.Cave Studies 1:1-13. (Box 682, Harding College, Searcy, AR72143).ZACHARDA, M. 1980. Soil mites of the family Rhagidiidae(Actinedida: Eupodoidea). Morphology, Systematics, Ecology.Acta Universitatis Carolinae (Prague), Biologica 1978(5-6):489-785.48Journal of the Arkansas Academy of Science, Vol. 36 [1982], Art. 15Published by Arkansas Academy of Science, 1982

https://scholarworks.uark.edu/cgi/viewcontent.cgi?article=2570&amp;context=jaas

Invertebrate Fauna of Devils Den, a Sandstone Cave in Northwestern Arkansas

Abstract

Similar works

Full text

Available Versions

UARK (University of Arkansas )

ScholarWorks@UARK