Nontroglobitic Fishes in Bruffey-Hills Creek Cave, West Virginia, and Other Caves Worldwide
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Environmental Biology of Fishes 62: 73-83,2001. O 2001 Kluwer Academic Publishers. Printed in the Netherlands. Nontroglobitic fishes in Bruffey-Hills Creek Cave, West Virginia, and other caves worldwide William J. Poly Department of Zoology, Southern Illinois University, Carbondale, ZL 62901 -6501, U.S.A. (e-mail: [email protected]) Received 29 February 2000 Accepted 17 December 2000 Key words: neoteny, pigment, eye Synopsis Six species of fishes were tagged and released in September and November 1995 and on five dates between November 1998 and October 1999 inside Bruffey-Hills Creek Cave. Most of the tagged fishes were creek chub, Semotilus atromaculatus, and green sunfish, Lepomis cyanellus. The overall recapture rate was 2.6% as only three of 117 fishes were recaptured. Forty-nine days was the longest time of residence by two L. cyanellus and one S. atromaculatus, and movement of 83.4m was observed only for the creek chub. Tag loss was confirmed as one factor in the low recapture rate. Nine species of fishes were collected in the cave, including Phoxinus oreas and Pimephalespromelas, two species never collected in a cave. Most of the fishes were pigmented normally, but many creek chubs were depigmented and appeared white or pigmentless when first observed in the cave stream. Four species, Phoxinus oreas, Pimephales promelas, Ameiurus nebulosus, and Lepomis macrochirus, were collected for the first time in Bruffey-Hills Creek Cave and in the Bruffey and Hills creeks drainage. Although data from this study did not shed light on residence time of fishes in the cave, the consistent occurrence of epigean fishes in this cave was shown. A list of epigean fishes from caves worldwide was included along .vith a discussion of aspects of the ecology of epigean fishes in caves and of evolution of troglobitic fishes. Introduction species of troglobitic crayfish in Shelta Cave, Alabama, then made periodic observations and noted move- Epigean fishes inhabiting caves have been given lit- ments of individual crayfish within the cave (Cooper & tle attention in the past by speleologists and cave Cooper1). Hobbs (1978) performed a mark-recapture biologists (Proudlove 1979, 1982, Chapman 1993). study of the crayfish, Orconectes inermis inermis, in Poly & Boucher (1996) reviewed the occurrence of Pless Cave, Indiana. Means & Johnson (1995) marked epigean fishes in caves and discussed the relevance Ozark cavefish, Amblyopsis rosae, with alphanumeric of these fishes to cave ecosystems and to understand- tags to estimate population size and to document move- ing evolution of troglobitic organisms. Information on ment of fishes within a cave, and Trajano (1991, the ecology of nontroglobitic fishes in caves remains 1997) performed similar studies on Pimelodella kronei, scarce, and the present study was undertaken to help fill this void. Tagging and periodic monitoring of fishes would provide useful biological information on I Cooper, J.E. & M.R. Cooper. 1971. Studies of the aquatic epigean fishes in caves. Tagging and recapturing of ecology of Shelta Cave, Huntsville, Alabama. ASB Bull. 18: 30 (abstract). Cooper, J.E. & M.R. Cooper. 1976a. Marking crayfish individual organisms in caves has not been attempted for long-term ecological studies. ASB Bull. 23: 52 (abstract). often and most previous studies have dealt with troglo- Cooper, M.R. & J.E. Cooper. 1976b. Growth and longevity in bitic organisms. Cooper (1975) used ink to mark three cave crayfishes. ASB Bull. 23: 52 (abstract). P. transitoria (an epigean species), and Trichomycterus a hip chain. Distance sampled (m) on each date was itacarambiensis in Brazil. Bruffey-Hills Creek Cave (in chronological order): 479.0, 264.4, 305.2, 132.6, was chosen as a study site because several species of 136.0, 138.0, 129.2, 127.6, and 123.4m. Becauseof an fishes occur in the cave in fair abundance. Informa- impassable logjam, less than 140 m of the cave could be tion is available on the vertebrate fauna of Bruffey- sampled between November 1998 and December 1999. Hills Creek Cave (Garton et al. 1993, Poly & Boucher Each tagged fish was weighed to the nearest 0.1 g on 1996), but the invertebrate fauna has not been exam- an Acculab Pocket Pro 250-B portable electronic bal- ined (Holsinger et al. 1976). The goals of the study ance (a spring scale was used during 1995 - weights were to gather information on: (1) residence time of recorded to the nearest 1 g), then measured to the near- individual fishes in the cave, (2) movement of indi- est 1 mm standard length (SL) and total length (TL) viduals throughout the cave, (3) growth of individuals with dividers and a stainless steel ruler. Initial pigmen- living in the cave environment, (4) reproduction within tation when captured (pallid or 'normal') as well as any the cave, and (5) temporal changes in pigmentation of changes from pallid (light) to 'normal' pigmentation individual fishes. after exposure to light for several minutes were noted. That a relationship exists between pigmentation and Then fishes were tagged with visible implant tags uti- length of residence in a cave is suggested by the study lizing a syringe style injector and precut, alphanumeric of Rasquin (1947) and reports of depigmented fishes tags (Northwest Marine Technology, Inc.). Tags were in caves (Hora 1924, Kosswig 1937, Proudlove 1979). inserted on the top of the head of cyprinids, on the Some fishes that were depigmented when captured in lower jaw of centrarchids, and on the adipose eyelid a cave were able to regain some pigment after sev- of salmonids. Voucher specimens were preserved in eral months of exposure to light (presumably a nor- 10% formalin and deposited in the Ohio State Univer- mal photoperiod, Banister 1984), whereas a 'white' sity Museum of Biological Diversity Fish Collection fish that appeared depigmented regained a more normal (OSUM), Southern Illinois University at Carbondale coloration in 24 h (Proudlove 1979). Also, fishes held Fluid Vertebrate Collection (SIUC), and University of in aquaria often exhibit temporary pigment changes, Tennessee, Knoxville (UT). Fish nomenclature follows appearing pallid after exposure to darkness (overnight), Robins et al. (1991) and Eschmeyer (1998). then appearing normal following exposure to light (per- Following completion of the field work, 55 creek sonal observation). Such a temporary state of depig- chubs obtained from Big and Little creeks, Union mentation was observed in some epigean fishes in a County, Illinois (16 January 2000), were tagged on cave (Poly & Boucher 1996). 17 January 2000 and checked weekly for the presence of tags. Creek chubs were held in circular fiberglass tanks in a greenhouse and fed every other day with Methods commercial pellets; small minnows were kept with the large creek chubs as additional forage. Smaller fishes Fish surveys were conducted on 22 October 1994, 30 (n = 50, SL = 41-82mm, n = 48 measured) were September 1995, 18 November 1995, 14 November kept in a separate tank from the larger chubs (n = 5, 1998,6 February 1999,27 March 1999,27 July 1999, SL = 114-134 mm, n = 4 measured) to reduce loss of 2 October 1999, and 29 December 1999 in Bruffey- fishes through predation. The temperature varied from Hills Creek Cave, Pocahontas County, West Virginia. 1.5 to 19.7"C between 17 January and 5 March. Fishes were tagged beginning 30 September 1995 and ending 2 October 1999. The cave is approximately 4 km north of Droop, WV, and approximately 2 km southeast Results of Lobelia, WV, Little Levels Township, 38"07'2lUN, 80" 16'42 W. The spring indicated on the Droop WV One hundred and ninety-six individuals of nine fish USGS 7.5' quadrangle map is actually the inflow species were collected between 1994 and 1999, and and cave entrance; Bruffey-Hills Creek Cave contains most of the fishes were creek chub and green sunfish 2156m of surveyed passages (Storrick 1992). Fishes (Table 1). One hundred and seventeen individuals of six were collected with a battery-powered backpack elec- species were tagged and released inside the cave: 90 trofishing unit, except on 22 October 1994 when seines Semotilus atromaculatus, four Rhinichthys atratulus, and dipnets were used. Distance sampled and distance 20 Lepomis cyanellus, one Lepomis macrochirus, one of capture or observation were measured to 0.1 m with Ambloplites rupestris, and one Oncorhynchus mykiss. Table 1. Fish species, number collected, and distance of capture in Bruffey-Hills Creek Cave, West Virginia. Date Species (size range, catalog number) Number Distance in cave' 22 Oct 1994 Semotilus atromaculatus (26-146mm, n = 11, OSUM 77874) 46' 4.8-476.0 Lepomis cyanellus (61 & 69 mm, OSUM 77875) 42 45.8476.0 30 Sep 1995 Semotilus atromaculatus (39-167 mm, n = 18) 19 2.7-264.4 Lepomis cyanellus (54-1 15 mm, n = 14) 14 2.7-198.5 Ambloplites rupestris (68 & 104.6 mm, SIUC 24889) 2 2.7-187.6 Rhinichthys atratulus (73 mm) 1 35.8 Oncorhynchus mykiss (275 mm) 1 35.8 18 Nov 1995 Semotilus atromaculatus (53-159 mm, n = 7) Lepomis cyanellus (63-1 10 mm, n = 6) 14 Nov 1998 Semotilus atromaculatus (45-185 mm, n = 27) 27 2.9-132.6 Lepomis cyanellus (141 mm) 1 110.5 Lepomis macrochirus (85 mm) 1 38.8 Rhinichthys atratulus (40 & 42 mm) 2 14.8 Phoxinus oreas (42-53 mm, n = 3, SIUC 33140) 3 2.9 6 Feb 1999 Semotilus atromaculatus (43-95 mm, n = 3) Rhinichthys atratulus (51 mm) Phoxinus oreas (53 mm, UT 44.861 1) 27 Mar 1999 Semotilus atromaculatus (3249 mm, n = 6) 6 19.5-37.0 27 Jul 1999 Semotilus atromaculatus (38-150mm, n = 18) Phoxinus oreas (34 & 58 mm, UT 44.861 1) Ameiurus nebulosus (210 mm, UT 48.1006) Ambloplites rupestris (1 10 mm, UT 90.2636) 2 Oct 1999 Semotilus atromaculatus (44-96 mm, n = 12) Lepomis cyanellus (25 & 75 mm) Pimephales promelas (53 mm, UT 44.8609) Rhinichthys atratulus (size not recorded) 29 Dec 1999 Semotilus atromaculatus (36-152 mm, n = 6, UT 44.8612) 64 2.4-106.2 Lepomis cyanellus (20-69 mm, n = 7, UT 90.2628) 74 2.4-46.0 'A range of capture distances is given when numerous individuals were captured.