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The University of Notre Dame The University of Notre Dame Ecology of Cave Use by the Frog, Rana palustris Author(s): William J. Resetarits, Jr. Source: American Midland Naturalist, Vol. 116, No. 2 (Oct., 1986), pp. 256-266 Published by: The University of Notre Dame Stable URL: http://www.jstor.org/stable/2425733 . Accessed: 25/01/2011 16:41 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=notredame. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Notre Dame is collaborating with JSTOR to digitize, preserve and extend access to American Midland Naturalist. http://www.jstor.org Ecologyof Cave Use by the Frog,Rana palustris WILLIAM J. RESETARITS, JR.' DepartmentofBiology, Saint Louis University,St. Louis,Missouri 63103 ABSTRACT: A trogloxenepopulation of thepickerel frog, Rana palustris, was studied at Ralph's Ford BluffShelter (RFBS), a cave on thenorthern edge of the Ozark Pla- teau in centralMissouri. Males werepresent beginning in Julywith females first ob- servedin September.Overall numbersbegan to increasein Septemberwith maxi- mum numbers present fromlate November through early March. Wintering populationnumbered in the hundreds.Movement of frogswithin the cave corre- spondedto changesin surfaceand cave temperatures.Frogs became inactiveat cave temperaturesbelow 6 C and concentratedin largenumbers in a physicallyand ther- mallysheltered hibernaculum near thecenter of thecave. Entryand emergencefrom hibernationin the cave correspondedwith the seasonal reversalsof temperaturegra- dientbetween the cave interiorand the surface.Emergence date variedby 6 weeks over 4 years of study.Winter mortality was skewedtowards juveniles and concen- tratedin thelatter half of winter (January-March). Feeding was suspendedfrom mid- Novemberthrough early-March. Rana palustrisis an importantcomponent of cave ecosystemsin theOzarks. The originand importanceof caveutilization in thespecies is probably related to historicalfactors, and dependent upon physiologicalpre- adaptationsand certainecological advantages of trogloxenehabits. INTRODUCTION Rana palustris(LeConte) occupies a broad geographic range encompassing much of eastern North America (Schaaf and Smith, 1970, 1971). Its typical habitat varies geo- graphically,ranging from"black water" swamps in the Carolinas (Hardy, 1964) to cool clear springs and caves in the limestone karst areas of Arkansas, Missouri, Illinois, Kentucky,Tennessee, Mississippi and Alabama (Schaaf and Smith, 1970). The perva- siveness of this latter association prompted Schaaf and Smith (1970) to suggest that caves and cold springsmay be a necessary component of R. palustrishabitat in this re- gion. Cave-associated populations of Rana palustrisare perplexingin termsof the distribu- tion, taxonomy and ecology of the species (Schaaf and Smith, 1970). Many North American species of the genus Rana have been observed in caves, includingR. catesbeiana (Barr, 1953; Holman, 1958; Lee, 1969; Pingleton et al., 1975; this study), R. clamitans (Myers, 1958; this study), R. sylvatica(Barr, 1953), R. pipienscomplex (Rand, 1950; Blair, 1951; Holman, 1958; Myers, 1958), R. sphenocephala(this study) and R. palustris (Smith, 1948; Barr, 1953; Brown and Boschung, 1954; Brode, 1958; Myers, 1958; Schaaf and Smith, 1970; Black, 1971; Knight, 1972; Brown, 1984; this study). With the exception of R. palustris,these instances appear to be sporadic, unrelated events. The cave habits of R. palustrisare unique among North American Anura, and may be unique among temperatezone Anura. Rana palustrishas been classifiedas trogloxene(Black, 1971) based on its persistent association with caves. Trogloxenesare species which occur in caves but are incapable of completingtheir life cycles there (Barr, 1963, 1968). Cave use, even at the trogloxene level, is especially interestingin an order containing no troglobiticor troglophilicspe- cies (Vandel, 1965) and raises questions regarding the origin and adaptive nature of cave use in trogloxenes. The major objectives of this study were to document the extent and nature of cave use in a trogloxenepopulation of Rana palustrisand to explore the possible role of caves in the distributionand ecology of the species. 'Present address: Departmentof Zoology, Duke University,Durham, North Carolina 27706. 256 1986 RESETARITS: CAVE USE BY FROGS 257 STUDY SITE The site chosen was Ralph's Ford Bluff Shelter (RFBS), located in southeastern Boone Co., Missouri, on Cedar Creek, at an elevation of approximately200 m. RFBS (Fig. 1) consistsof approximately213 m of main passageway,beginning in a bluff-face entrance 2 m above the normal level of Cedar Creek and ending in a collapsed chamber (sinkhole) in wooded pasture '\18 m above the creek. A single side passage branches off the centralportion of the cave and is traversiblefor '\50 m. The floorof this passage is higher than the main passage, consistsof hard-packed clay, and is usually dry. No sur- face entrance to this passage is known. The main passage contains a small intermittentstream with moderatelydeep (30-60 cm), permanent pools connected by short shallow runs. Flow is maintained in all but the driest conditions and is fed by direct surface discharge into the sinkhole entrance (during wet periods), and subterranean flow enteringat several points along the cave stream. Flow varies froma trickleduring prolonged droughts,to flash floods which fill the main passage to the ceiling. The main passageway was divided into six zones forthe purposes of this study (Fig. 1). Zones 1 and 6 correspond roughly to the lower and upper twilightzones, respec- tively; 2 through5 divide the interiorinto approximatelyequal sections (Fig. 1). The most significantmicrohabitat within the cave with respectto Rana palustrisis the pool in the center of Zone 4 (hereafterreferred to as the pool) (Fig. 1A.). The pool is slab and gravelbottomed, and approximately1 m x 4 m and 40-50 cm at its greatestdepth. The hibernaculum is a partiallysubmerged, physicallyand thermallysheltered area offthe NW edge of the pool. It consists of a shallow, 5-15 cm high, shelf-likecave, approxi- mately 60 cm across at the entrance and of unknown depth. In midwinterthe hibernac- ulum site contains almost the entire overwinteringpopulation of R. palustrisat RFBS. Water in the hibernaculum remains approximately 1.5-2.0 C warmer than the pool proper during midwinteras a resultof the depth of the passage into the rock and the di- rect influxof groundwater. \~CREEK A (dry) B LENGTH= 213m B ~~~~VERT.RISE =2Cm 4 UPPER ( ZONES ENTRANCE 6~~~~~~~~~~~~ 7m Fig. 1. -Map of RFBS showingcave zones and significantfeatures. (A) indicateslocation of thepool, (B) thehibernaculum and (C) the DP (dividerpool) temperaturestation 258 THE AMERICAN MIDLAND NATURALIST 116(2) Ozark caves typicallymaintain a uniformtemperature throughout the year, approx- imated by the local mean annual surfacetemperature. RFBS temperaturesvary mark- edly fromexpected and closely parallel changes in external ambient temperature(Fig. 2). Lowest temperaturesmeasured at the pool were 0 C air and 5.2 C water and the highestwere 18.9 -C air and 13.2 C water.Air temperatureextremes were measured us- ing a Taylor Max-Min thermometerplaced over the pool on a rock ledge. Water tem- peratureextremes were recorded during cave visitsusing a SchultheisQuick-Read ther- mometer.Water in RFBS froze to beyond Zone 1 during the winterof 1980-1981, the furthestextent of ice observed. Water 10 m fromthe upper entrance was not frozen at thistime. MATERIALS AND METHODS The study site (RFBS) was visited on an irregularbasis fromAugust 1979 through March 1982. The entire main passage was surveyedon most cave visits. Location and estimatednumbers of frogswere recorded. Air and water temperaturedata were taken at six stationsusing SchultheisQuik-Read and Taylor Max-Min thermometers.Supple- mental visits to the cave continued throughApril 1984. A mark-recapturestudy was completed in the winterof 1979-1980. Frogs were cap- tured by hand, freeze-brandedwith a number on the venter(Daugherty, 1976), and re- leased at the point of capture. Nine frogsmarked and kept in the laboratoryshowed no evidence of deteriorationof the brands over several months. Frog #36, marked on 1 January 1980, carried the clearlylegible brand when collected and preservedon 14 Feb- ruary 1981. Population estimateswere made using the Schumacher method (Schumacher and Eschmeyer,1943). This method was chosen for its usefulnessin providinga single reli- able population estimatefrom a series of mark-recaptureepisodes. Seventy-fiveRana palustris(33
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