BUllETIN OF THE NATIONAL SPELEOLOGICAL SOCIETY

VOLUME 33 NUMBER 1

Contents

NORTH AMERICAN TROGLOBITIC SALAMANDERS

INVERTEBRATE CAVE FAUNA OF GEORGIA

SOUTHERN CAVEFISH AT SOUTHEASTERN EDGE OF RANGE

CoMPARISON oF CAVE AND DEEP SEA ORGANISMS

CLASSIFICATION OF CAVERNICOLES

REsUME: 1970 BIOLOGY SHORT CoURSE

REVIEW: OBSERVATIONs ON THE EcoLOGY oF CAvEs

REVIEW: CAVE EcoLOGY AND TROGLOBITE EvoLUTION

JANUARY 1971 NATIONAL SPELEOLOGICAL SOCIETY The National Speleological Society is a non-profit organization devoted to the study of caves, karst and allied phenomena. It was founded in 1940 and is incorporated under the laws of the District of Columbia. The Society is affiliated with the American Associ­ ation for the Advancement of Science. The Society serves as a central agency for the collection, preservation, and dissemi­ nation of information relating to speleology. It also seeks the preservation of the unique faunas, geological and mineralogical features, and natural beauty of caverns through an active conservation program. The affairs of the Society are controlled by an elected Board of Governors, which appoints National Officers. Technical affairs of the Society are administered by specialists in the fields that relate to speleology through the Society's Biology Section, Section on Cave Geology and Geography, and Research Advisory Committee. Publications of the Society include the quarterly BuLLETIN, the monthly NEws, OccASIONAL PAPERS, and MISCELLANEous PUBLICATIONS. Members in all categories except Family Dependent receive the BULLETIN and NEws. A Library on speleological subjects is maintained by the Society at 21 William Street, Closter, New Jersey 07624. Material is available to Society members at a nominal fee to defray the cost of handling, and to others through inter-library loan. An extensive file of information on caves of the United States is maintained by the Society and is currently housed in Lawrence, Kansas.

OFFICERS 1970-71 RANE L. CURL, President WAYNE FooTE, Adm. Vice President JOHN E. CooPER, Executive Vice President DoNALD N. CoURNOYER, Treasurer

DffiECTORS JoHN R. HoLSINGER RussELL H. GURNEE ALAN E. HILL LARRY D. MATTHEWS WILLIAM R. HALLIDAY DAVID IRVING JoHN M. RUTHERFORD G. P. "JIM" HIXSON CHARLES v. LARSON EuGENE VEHSLAGE KENNETH N. LAIDLAw DAviD R. McCLURG

RESEARCH ADVISORY COMMITTEE DR. JoHN R. HoLSINGER Dept. of Biology Old Dominion University Norfolk, Va. 23508

BIOLOGY SECTION SECTION ON CAVE GEOLOGY AND GEOGRAPHY DR. RICHARD GRAHAM Dept. of Biology DR. WILLIAM WmTE Upsala College Materials Research Laboratory East Orange, N. J. 07019 210 Engineering Science Building The Pennsylvania State University University Park, Pa. 16802 BULLETIN of the NATIONAL SPELEOLOGICAL SOCIETY VOLUME 33, NUMBER 1 JANUARY 1971

BIOLOGY ISSUE John E. Cooper and Thomas L. Poulson, Guest Editors

CONTENTS NORTH AMERICAN TROGLOBITIC SALAMANDERS: SoME AsPECTS OF MoDIFICATION IN CAVE HABITATs, WITH SPECIAL REFERENCE TO Gyrinophilis Palleucus ...... Ronald A. Brandon 1 THE INVERTEBRATE CAvE FAUNA OF GEORGIA .. John R. Holsinger and Stewart B. Peck 23 THE SouTHERN CAVEFISH, Typhlichthys Subterraneus, AT THE SouTH- EASTERN PERIPHERY OF ITS RANGE ...... John E. Cooper and Anthony lies 45 THE BIOLOGY OF CAVE AND DEEP SEA ORGANISMS: A CoMPARISON .. Thomas L. Poulson 51 THE CLASSIFICATION OF CAVERNICOLES ...... Elery Hamilton-Smith 63 RESUME OF THE 1970 SHORT CouRSE IN CAvE BIOLOGY ...... John R. Holsinger 67 CAvEs As MoDEL SYSTEMS: A REviEw oF "OBsERVATioNs oN THE EcoLoGY oF CAvEs" ...... David C. Culver 68 A REviEW oF "CAvE EcoLoGY AND THE EvoLUTION oF TROGLOBITES" ...... Kenneth Christiansen 7 0

The BuLLETIN is published quarterly. The subscription rate in effect January 1, 1971: $6.00. Office Address: NATIONAL SPELEOLOGICAL SOCIETY 2318 N. KENMORE ST. ARLINGTON, VIRGINIA 22201 Discussion of papers published in the BuLLETIN is invited. Discussions should be 2,000 words or less in length with not more than 3 illustrations. Discussions should be forwarded to the appropriate editor within three months of publication of the original paper.

MANAGING EDITOR DAVID IRVING E. I. DUPONT, Savannah River Lab. Aiken, S. C. 29801 Biology Editor Earth Sciences Editor DR. DAVID c. CULVER DR. WILLIAM B. WHITE Dept. of Biology Materials Research Laboratory University of Chicago 210 Engineering Science Building Chicago, Ill. 60637 The Pennsylvania State University University Park, Pa. 16802

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The BuLLETIN is published quarterly in January, April, July, and October. Material to be included in a given number must be submitted at least 90 days prior to the first of the month of publication. North American Troglobitic Salamanders: Some Aspects of Modification in Cave Habitats, with Special Reference to Gyrinophilus palleucus

By Ronald A. Brandon ll<

ABSTRACT Seven of the eight known species of troglobitic salamanders are found in Korth America. Of these seven, all closely related within the family Plethodonti­ dae, only Typhlotriton spelaeus normally undergoes metamorphosis. There is ample evidence that the others, Eurycea troglodytes, Gyrinophilus palleucus, Haideotriton wallacei, Typhlomolge rathbuni, Typhlomolge tridentifera, and prob­ ably Eurycea latitans, are paedogenetic and reproduce while retaining larval body morphology. Of the last six, only G. palleucus is known to undergo major metamorphic change readily in response to experimentally administered thyroxin. The more highly specialized species tend to have more rigid control of paedo­ genesis, increased numbers of teeth, fewer trunk vertebrae, more reduced eyes, more reduced integumentary pigmentation, broader heads, flatter snouts, and more elongate and attenuate limbs. All of these features, except number of trunk vertebrae, are considered selectively advantageous to salamanders living under cave conditions; they probably evolved in response to food requirements and food availability and result in more efficient energy utilization by the population. Probable mechanisms in the evolution of these features are general neotenic trends and allometric growth changes. The number of trunk vertebrae seems related not to degree of specialization to cave habitats, but rather to the ancestry of the species. Gyrinophilus palleucus, in several ways, seems to be the least specialized troglobitic salamander. It responds readily to thyroxin, its eyes are not greatly reduced, members of some populations are quite heavily pigmented, and in most regards it is similar to larvae of the epigean species Gyrinophilus porphyriticus. It is highly variable geographically, and has a range which is on the periphery of that of G. porphyriticus, from which species it may have arisen under changing climatic conditions of the Pleistocene.

INTRODUCTION (Brame, 1967), only eight species are tro­ Of the more than 300 known species of globites (sensu Barr, 1960). These are obli­ salamanders, arranged in eight families gative cave-dwellers and can complete their life cycles only in caves. Larvae of one 0 Department of Zoology, Southern Illinois Uni­ species, Typhlotriton spelaeus, can be found versity, Carbondale, Illinois. in springs and spring-fed surface streams in

VoLU:\IE 33, NuMBER 1, jANUARY 1971 1 0.....___ 200 Miles

o T tridentifero

tJ. Typhlomolge rothbu..Q!

• Eurycea troglodytes .

€2) Gyrinophilus •

® Typhlotriton spetaeus

([) Profel!S anguinus

~ t1oideotriton wollacei Figure 1. Distribution of troglobitic salamanders. the vicinity of caves (and for this reason is and adjacent Florida; Typhlotriton spelaeus not considered a troglobite by Vande!, 1964, Stejneger, 1892, in the ozarkian Springfield or Poulson, 1964), but there is no evidence and Salem plateaus; and Gyrinophilus pal­ that they can transform outside of caves, and leucus McCrady, 1954, in Tennessee, north­ adults have not been found outside of caves. ern Alabama, and northwestern Georgia. Except in rare instances when they may be These seven species are all placed in the washed out in flood waters, individuals of family Plethodontidae, the largest (contain­ the other troglobitic species are never found ing about 180 species in 24 genera) and in epigean situations. most diverse salamander family. The family Only one troglobitic salamander, the pro­ Plethodontidae is thought to be indigenous teid Proteus anguinus Laurenti, 1768, is to the Appalachian Mountains of eastern known in the Old World. It occurs in karst North America (Dunn, 1926; Wake, 1966 ), regions along the eastern Adriatic coast, in and, except for two species of Hydromantes Trieste and Yugoslavia (Figure 1). The in southern Europe, it is presently restricted remaining seven troglobitic species are all to the New World. The only salamanders found in eastern United States; they are known to have invaded the tropics south Typhlomolge rathbuni Stejneger, 1896; of the equator are members of this family Typhlomolge tridentifera (Mitchell and (the genus Bolitoglossa in South America ) . Reddell, 1965); Eurycea latitans Smith and Of all the troglobitic salamanders, only Potter, 1946; and Eurycea troglodytes Baker, Typhlotriton spelaeus undergoes the com­ 1957, all in the Edwards Plateau of Texas; plete metamorphosis common to the life Haideotriton wallacei Carr, 1939, in the cycles of most salamanders. In all other Daugherty Plain of southwestern Georgia troglobitic species sexual maturity, mating,

2 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN and deposition of eggs apparently occur troglobitic invertebrates and in troglobitic while the adult retains larval morphology. fishes is also evident among the troglobitic Natural metamorphosis does not occur and salamanders. Similarly convergent, but more they are paedogenetic. Different amounts cryptic, features of physiology and ecology of metamorphic change can be induced certainly exist, but have barely been investi­ experimentally in these salamanders, rang­ gated in troglobitic salamanders ( cf. Poul­ ing from nearly complete metamorphosis in son, 1964). There is a general trend toward G. palleucus to only a very few metamor­ more rigidly controlled paedogenesis, in­ phic changes in H. wallacei. The paedoge­ creased number of teeth, reduced number netic adults of North American troglobitic of trunk vertebrae, reduced eyes, reduced salamanders are definitely of plethodontid body pigmentation, broadening of the head larval morphology. They lack lungs (as do and flattening of the snout, and elongation all members of this family and certain and attenuation of the limbs. The remainder stream-dwelling members of three other of this paper deals in detail only with the families), have no ypsiloid cartilage, have North American species (Table 1). three gill slits, and generally resemble ple­ Paedogenesis thodontid larvae externally. The paedoge­ netic species of Eurycea and Gyrinophilus Data indicating the paedogenetic nature closely resemble larvae of epigean members of some of the troglobitic salamanders are of the same genera. Typhlomolge mthbuni, circumstantial only, and perhaps the ques­ T. tridentifera and Haideotriton wallacei are tion of whether they are actually paedoge­ more similar in structure to larvae of Gyrin­ netic should be renewed and reviewed ophilus, Pseudotriton, Eurycea and Typhlo­ briefly here. Misinterpretations have occur­ triton than to those of other plethodontids. red in the past when paedogenesis was Thus, not only are all North American assumed but not documented. Gyrinophilus troglobitic salamanders placed in the same lutescens (see Rafinesque, 1832; Mittleman, family, but within the family they group 1942; Brandon, 1966a) and Typhlotriton closely and are probably closely related nereus (see Bishop, 1944; Brandon, 1966b) (cf. \Vake, 1966). are good examples. T. nereus, described as a paedogenetic species, was based on lar­ CHARACTERISTICS val, immature specimens of T. spelaeus. OF TROGLOBITIC SALAMANDERS The name G. lutescens, as used by Mittle­ The striking convergence of morphology, man for a paedogenetic species, was applied eye reduction, and depigmentation in many to immature larvae of G. porphyriticus.

TABLE I. Sample of specimens examined, and known maximum snout-vent length of each species (in this, and subsequent tables and figures, all measurements are in mm).

Maximum Size Range Number of Known Body of Specimens Specimens Species Length Examined Examined

Typhlomolge rathbuni ...... 76 28-60 44 Typhlomolge tridentifera ...... 37 19-34 20 Haideotriton wallacei ...... 44 13-35 25 Eurycea troglodytes ...... 40 22-40 13 Eurycea latitans ...... 53 39-46 2 Larval Typhlotriton spelaeus ... 56 13-56 hundreds Gyrinophilus p. gulolineatus .. 136 66-136 11 Gyrinophilus p. necturoides ...... 105 48-105 19 Gyrinophilus p. palleucus ...... 100 47-94 20

VoLC:'\IE 33, NuMBER 1, }ANUARY 1971 3 Following is a summary of information, described. Stejneger ( 1896, p. 620) com­ some from the literature and some original, mented that one living female specimen indicating paedogenesis in troglobitic sala­ ". . . expelled three eggs after being caught manders found in North America. . . .", and that the body of another female Gyri1wphilus palleucus.-Courtship, laid contained large eggs. Dunn ( 1926) exam­ eggs, and newly hatched young have not ined a female specimen whose spermatheca been reported. Lazell & Brandon ( 1962) was packed with sperm. Both of Emerson's reported a male whose cloaca held a sperm­ ( 1905) specimens were females, and her atophore. Dent & Kirby-Smith ( 1963) found figures (fig. D & E on p. 69) and her tex­ that males of G. p. palleucus begin to pro­ tual description show that one was mature. duce sperm when they are 70 to 84 mm Its oviducts were enlarged and convoluted, long (snout to vent), and that a female and its ovaries contained large ( 1.5 mm 100 mm long contained large ( 3.2 mm diameter), yolked ova. Enlarged and con­ diameter) ovarian eggs. The testes of three voluted oviducts indicate sexual maturity in male G. p. gulolineatus bore black pigment Plethodon cinereus (Saylor, 1966) and in (Brandon, 1965b), a sign of sexual maturity several species of Desmognathus (Organ, in G. porphyriticus and many other pletho­ 1961), and probably in T. rathbuni as well. dontids. These data, plus the nature of their I have examined the reproductive organs of response to metamorphic agents (Dent, et 19 female specimens ranging from .33 to 60 al., 1955; Dent & Kirby-Smith, 1963) make mm snout to vent. Oviducts were enlarged it clear that G. palleucus is normally neo­ and convoluted in the two largest specimens tenic, even though specimens readily and (58 & 60 mm), and these specimens con­ consistently undergo much metamorphic tained yolked ovarian eggs 0.7 to 0.9 111m change in response to thyroxin experiment­ in diameter. In five specimens, considered ally administered. immature ( 39, 42, 45, 45, 51 111m) the H aideotriton wallacei.-Courtship and oviducts were small, straight, and medially laid eggs have not been observed. The bolo­ placed; ovaries contained only small ( 0.2 type, the largest specimen known ( 44 mm to 0.4 mm diameter), unyolked ova. The snout to vent) is a gravid female as Carr's gonads of two smaller specimens ( 3.3 & 37 ( 1939) photographs clearly show. It was mm) were not sufficiently differentiated for preserved late in May. Valentine ( 1964) detem1ination of sex by gross examination. measured its ovarian eggs at 2 to 2.2 mm Sexual maturity probably occurs at 40 to in diameter; these are probably nearly full 50 mm snout-vent length in female T. size judging by size of mature eggs in other rathbuni. This compares with an estimate salamanders of comparable size-e.g., 2.5 to of 40 mm for males based on testicular 3 mm in Eurycea bislineata (Bishop, 1941). lobulation. I have examined the reproductive organs of 14 specimens ranging in snout-vent length Humphrey ( 1922) showed, in Desmog­ from 13 to 28 mm. On the basis of their nathus fuscus, that a positive correlation relatively undifferentiated gonads and repro­ exists between number of testicular lobes ductive tracts all are considered immature. and age of mature males. The manner in Response of several subadults to treatment which the spermatogenic wave passes with thyroxin in Dundee's experiments through the testis causes a progressive in­ ( 1962) (only slight metamorphic changes crease in number of lobes throughout the in the integument, loss of the coronoid bone life of the male. Organ ( 1961) used the in the lower jaw, and resultant death within number of lobes to recognize year classes 25 days) strongly suggests that H. wallacei of several species of Desmognathus, and a does not transform in nature and will not similar relationship between number of lobes under experimental conditions. and specimen length has been observed in Typhlomolge rathbuni.-Courtship has some other salamanders: for example, in not been observed nor have laid eggs been Phaeognathus hubrichti by Brandon ( 1965a)

4 THE NATIONAL SPELEOLOGICAL SociETY BcLLETIN and Typhlotriton spelaeus and others in­ the testes were black. Cloacal lips were cluded below. swollen conspicuously and appeared to be The number of lobes in males of T. rath­ glandular on the two largest males, were less huni ranged from none to four in seven distinctly swollen on two males, and were specimens examined, and relates well to not swollen at all on three. I would judge specimen length (Figure 2). It is probable the 34, 32, 30, and 26-mm males to be that individuals with 3 or 4 lobes have adult and sexually active. The 29-mm male produced sperm 4 to 7 times previously looked adult but sexually inactive, one 25- (see Organ, 1961, for method of estimat­ mm male and the 22-mm one seemed to be ing). In sexually mature T. rathbuni the approaching maturity, and another 25-mm testicular lobes are finely reticulated with male looked immature. Sexual maturity, melanophores and interlobular portions of therefore, seems to be reached in males 22 the testes and the vasa deferentia are solid to 25 mm long. Females also appear to black. Black pigment on these structures mature at about 25 mm. Immature females indicates sexual maturity in many other ( 20, 22, 23, 25 mm long) had small, medi­ plethodontid salamanders and certainly does ally placed oviducts and tiny ( 0.15 to 0.2 in T. rathbuni as well. mm diameter), unyolked ovarian ova. Fe­ males with enlarged and convoluted oviducts A. were 27, 28 ( 3), and 30 mm long, and two of these contained enlarging ( 0.6 to 1.5 mm • • • diameter), yolked ovarian ova. • • .. Eurycea latitans.-Courtship and laid (/) w ~I ., . eggs are unknown, and extremely little m 0 information of any kind is available on this ...J species. The maximum known snout-vent "'- 4 • 0 B. length is about 53 mm (Brown, 1967). a: 3 w • Smith & Potter ( 1946) included some infor­ m :I 2 mation on reproductive structures in their :::::1 • z description of this species. For example, they •• mentioned ( p. 107) that two of their female specimens under 53 mm snout-vent « ••• possess very large eggs presumably nearly 15 20 30 40 50 60 ready to be laid", and that "The males, SNOUT-VENT LENGTH !MM) regardless of date of collection, are rather Figure 2. Correlation between snout-vent well marked by the swollen glandular area length and number of testicular lobes in about the anus." Brown ( 1967) noted that Typhlomolge tridentifera (A) and Ty­ a swollen glandular vent is a feature of phlomolge rathbuni ( B ) . mature males. It should be pointed out that the holotype, USNM 123594, identified ori­ Typhlomolge tridentifera.-Courtship and ginally as an adult male, actually is a female laid eggs are unknown. Most specimens in whose oviducts are slightly enlarged and the type-series were identified to sex by whose ovaries contain numerous small ( 0.5 Mitchell & Reddell ( 1965) but no criteria mm diameter) ova. The specimen listed by for recognizing mature individuals were Smith and Potter ( 1946) as TCWC 1195, given. I have examined reproductive organs a paratype and the only additional specimen of 18 specimens: eight males, nine females, able to borow for exam­ and one undetermined juvenile. The num­ which I have been ber of testicular lobes ranges from none to ination, is a mature male whose darkly pig­ two in specimens 22 to 34 mm long (Figure mented testes are composed of one large and 2). Lobes of the two-lobe males were heav­ one small lobe each. ily reticulated with black pigment and the Eurycea troglodytes.-Nothing is reported vasa deferentia and interlobular spaces of about courtship or laid eggs. Known speci-

VOLUME 33, NUMBER 1, }ANUARY 1971 5 mens range from 23 to 40 mm snout-vent. correlated with dwelling in spring and cave Baker ( 1957) identified the 38-mm halo­ habitats. type as a sexually mature female, apparently Paedogenesis is probably of adaptive value on the basis of its folded cloacal walls. He to troglobitic salamanders and certainly is did not comment on the size at sexual not merely a phenotypic response to cave maturity or criteria for determining matur­ climatic conditions. In some way the rather ity; the body cavities of neither the holotype elaborate changes which take place in the nor twelve paratypes which I have seen had usual pattern of salamander metamorphosis been opened. I have examined the reproduc­ are avoided. P. anguinus has an active thy­ tive organs of the holotype (a female) and roid gland, but its tissues do not respond to 6 paratypes ( 3 males, 3 females). Preserva­ thyroxin (Schreiber, 1933). H. wallacei and tion of internal organs is poor, but all speci­ T. rathbuni respond only slightly to large mens over 23 mm snout-vent appear mature. experimental doses of thyroxin, even though Four of the females ( 32 to 40 mm) have their thyroid follicles secrete and release straight, medially placed, slightly enlarged hormone (Dundee, 1957, 1962; Gorbman, oviducts and numerous small ( 0.2 to 0.6 mm 1957); the treatment usually is fatal. G. diameter) ovarian ova. The poor preserva­ palleucus readily undergoes almost complete tion makes it impossible to determine if the metamorphosis during thyroxin treatment, largest ova are yolked; the specimens appear but the thyroid gland normally maintains a mature but sexually inactive. The males ( 32 low level of activity (Dent and Kirby-Smith, to 39 mm long) all have black vasa defer­ 1963). No metamorphosed individual of G. entia and darkly reticulated testes. The palleucus has ever been found in nature, but number of testicular lobes ranges from 1 to a few have transformed "'spontaneously" in 3. All of the males, thus, are sexually ma­ the laboratory (Dent & Kirby-Smith, 1963). ture. Sexual maturity probably occurs at Since at least three paedogenetic species of around 25 to 30 mm snout-vent in this Eurycea (E. tynerensis, E. nana, and E. species. neotenes) are known to undergo metamor­ In summary, there is ample evidence of phic changes after thyroxin treatment sexually mature, branchiate individuals of ( Kezer, 1952; Potter & Rabb, 1959), it is all of the North American troglobitic sala­ likely that E. troglodytes and E. latitans manders except Typhlotriton spelaeus and may give similar responses when tested. Eurycea latitans. Indications are that the Dundee and Gorbman ( 1960) have shown latter actually is paedogenetic as well. that E. tynerensis is paedogenetic probably In all instances in the family Plethodon­ due to a low level of thyroxin synthesis. tidae, paedogenesis is correlated with eco­ There seem to be at least two different logical restriction to cave or spring habitats. mechanisms resulting in paedogenesis in The only non-cavernicolous paedogenetic troglobitic salamanders: ( 1) loss of target plethodontids are Eurycea tynerensis and tissue sensitivity, and ( 2) low production some populations of E. multiplicata which of thyroxin. Paedogenesis seems to have inhabit springs and spring-fed streams along survival value regardless of the mechanism the southwestern edge of the Ozark Plateau, by which it is obtained. In more specialized and several more species of Eurycea (E. troglobites resistance to metamorphosis is nana, E. neotenes, E. pterophila) in springs more rigidly controlled; i. e., environmental along the Balcones Escarpment in Texas. variables and experimental treatments will All other paedogenetic plethodontids are not induce a successful metamorphosis. G. troglobites, and all of the troglobites, except palleucus, a relatively unspecialized troglo­ T. spelaeus, are paedogenetic. It should be bite morphologically, does not "'resist" en­ pointed out that paedogenesis and neoteny vironmentally or experimentally induced do occur in other salamander families, but metamorphosis, although metamorphosis in in them (with the exception of Proteus nature seems to be extremely rare if it occurs anguinus in the family Proteidae) is not at all.

6 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN Adults of Typhlotriton spelaeus have no ments, and would be important in nonvisual paedogenetic features, with the possible ex­ feeding behavior. As Poulson has also ception of a delayed metamorphosis of the pointed out, it is likely that some apparently premaxillary bone (Wake, 1966). Accord­ neotenic features are really secondary, allo­ ingly, Vandel (1964) and Poulson ( 1964) metric effects of selection for lower develop­ do not consider T. spelaeus a troglobite at mental and growth rates in response to food all since, in addition, its larvae are some­ scarcity. times common in surface waters. Barr Number of Teeth ( 1963) and Brandon, on the other hand, do consider T. spelaeus a troglobite. Adult T. Tooth counts most commonly made on salamander spelaeus are restricted ecologically to cave larvae are of those on the pre­ maxillary, habitats, and apparently can complete their prevomerine (vomerine of some authors), life cycle only there. In addition they are and pterygoid (palatine or palata­ pterygoid distinguished morphologically by reduced of some authors) bones. Data on number body pigmentation and reduced eyes. A of teeth on the lower jaw (on the dentary recurrent idea in the literature (e. g., Noble, and coronoid bones) have seldom been 1927; Mittleman, 1950; Conant, 1958; Van­ given, and I have not counted these del, 1964) is that larvae of T. spelaeus live either. Seldom are counts of samples through­ primarily in cave entranceways and surface out the size range of a species given in the streams and that as metamorphosis ap­ literature. Statements of average tooth num­ proaches they retreat into caves. Actually, bers are seldom comparable because of onto­ there is no evidence that this is the usual genetic changes in number. Counts of pre­ sequence of events, or that significant num­ maxillary teeth from T. tridentifera of known bers of surface-dwelling larvae actually do body size, however, are given by Mitchell return to caves. It is probable that subse­ & Reddel ( 1965). quent generations are produced mainly by I have counted teeth on a series of speci­ larvae that never leave the caves at all, and mens of each troglobitic species, except E. latitans, using that rather high mortality prevails among a binocular microscope at 15-20x, on specimens whose jaws were cut epigean larvae. Such large numbers of larvae to reveal the entire roof of the mouth. Data in above-ground situations, however, does on larval T. spelaeus can be found elsewhere suggest that epigean dispersion may be (Brandon, 1966b), as can some information important in this species. T. spelaeus has the on T. tridentifera (Mitchel & Reddell, largest geographic range of any troglobitic 1965), E. latitans (Smith & Potter, 1946), salamander and yet varies little geographi­ and H. wallacei (Valentine, 1964). cally. In each species examined, except G. Avoidance of metamorphosis in troglobitic palleucus, there is a distinct ontogenetic salamanders is probably related to food increase in number of premaxillary teeth requirements and food availability. Most (Figures 3 and 4). T. rathbuni and H. available food items deep within caves are wallacei have the most and larval T. spel­ aquatic, either aquatic cavernicolous inver­ aeus and E. latitans have the fewest. Data tebrates or organisms washed in from out­ on the latter, however, are available from side. Even transformed T. spelaeus, which only two specimens. G. palleucus, at all are essentially terrestrial, can feed heavily sizes, has more than larvae of its epigean on aquatic animals (Smith, 1948). Food in congener G. porphyriticus. T. tridentifera caves is usually scarce (Poulson, 1964). and E. troglodytes have an intermediate Avoidance of metamorphosis greatly reduces number and do not differ from each other. the energy requirements of the individual The number of prevomerine teeth in­ and of the population. In addition, neuro­ creases distinctly with ontogeny in H. wall­ masts typically degenerate when larval ple­ acei, T. tridentifera and larval T. spelaeus, thodontid salamanders transform. These slightly in G. palleucus, but not in T. rath­ sense organs are sensitive to water move- bttni or E. troglodytes (Figures 5 and 6).

VoLUME 33, NuMBER 1, JANUARY 1971 7 60 An ontogenetic change from a broadly

0 elongate patch of pterygoid teeth to fewer

0 teeth arranged in a single row occurs about 50 0 AO 0 half-way through larval life in T. spelaeus X A 0 1- (Brandon, 1966b ), perhaps in E. troglo­ ILl ILl oA dytes, and in at least two epigean species 1- 0 40 0 >- of Eurycea (E. lucifuga and E. multipli­ a: <( A cata). Even in the smallest specimens of ....J ....J the other troglobites examined, these teeth x<( 30 :::E A were arranged in a single row, as they were a:ILl • in larval G. porphyriticus. Interestingly, H . a.. A • • A A X cj XX I X wallacei has generally fewer pterygoid teeth z 20 XX XX X X • • than all other species examined, except eX large, larval T. spelaeus and larval G. 1101'­ phyriticus (Figures 7 and 8). Of those 10 20 30 40 50 species with these teeth in a single row throughout SNOUT-VENT LENGTH (MMl ontogeny, T. rathbuni has the most, followed by Figure 3. Number of premaxillary teeth T. tridentifera, E. troglo­ dytes and versus snout-vent length in Haideotriton E. latitans, G. palleucus, and H. wallacei. wallacei (triangles), Typhlomolge rath­ The buni (open circles), Typhlomolge tri­ correlation between increased num­ bers of teeth dentifera (X) and Eurycea troglodytes and specialization to cave habitats has (solid circles). been pointed out previously (Baker, 1957), at least for paedogenetic forms inhabiting the Edwards Plateau in The number is reported to be low in E. Texas. The increased numbers of premaxil­ latitans (Smith & Potter, 1946). Among lary and prevomerine teeth in more larger specimens, H. wallacei, T. rathbuni special­ ized troglobites and T. tridentifera have the most prevome­ is probably correlated with rine teeth, whereas E. troglodytes, E. lati­ increased efficiency in catching and holding tans and G. palleucus have fewer, although prey in an environment where the popula­ data are not comparable because of the tion density of prey organisms is low. difference in ontogenetic increase. Again, G. Reduction of Eyes palleucus generally has more than larvae of The detailed histological structure of the its relative G. porphyriticus. eyes of several of the troglobitic salamanders

40 I 1- w w 1- >- a: 30 <( 0 • ....J 0 • • • • x....J 0 • • • • • 0 • • • • ::::E 0 "'w 20 0 • •0 • a: 0 0000 00 )( a.. 0 ' 0 z0

10 20 30 40 50 60 70 80 90 100 SNOUT-VENT LENGTH !MMl Figure 4. Number of premaxillary teeth versus snout-vent length in larval Typhlotriton spelaeus (open circles), Eurycea latitans (X), and Gyrinophilus. palleucus (solid circles ) .

8 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN :I: .,_ XA L&J 0 w 40 0 t- 0 0 w A X 0 0 z A 0 0 0:: •eooo 0 lLJ XA XX oe ~ 30 0 0 0 )( • > ~ • l&J X 0:: • • Q. A A • 0 A z 20

10 20 30 40 50 60 ·SNOUT- VENT LENGTH (MM) Figure 5. Number of prevomerine teeth versus snout-vent length in Haideotriton wallacei (triangles), Typhlomolge rathbuni (open circles), Typhlomolge tridentifera (X), and Eurycea troglodytes (solid circles).

:r: 1- UJ 40 UJ 1- 0 0 w 0 z 0 0 • • • • • ~ 0 0 I • UJ 30 • 00 • ••• • • • ~ • • 0 0 • > 0 o UJ 0 08 0 X• a: X a. 0 0 0 0 20 z 0 0

10 20 ~0 40 50 60 70 eo 90 100 SNOUT-VENT LENGTH (MM) Figure 6. Number of prevomerine teeth versus snout-vent length in larval Typhlotriton spelaeus (open circles), Eurycea latitans (X), and Gyrinophilus palleucus (solid circles). has been investigated: e. g., of T. rathbuni Baker ( 1957) included information on eye ( Eigenmann, 1900 ), T. spelaeus (Eigen­ diameter in series of E. latitans, E. troglo­ mann & Denny, 1900; Alt, 1910; Stone, dytes and T. rathbuni of known body length, 1964), G. palleucus ( Lazell & Brandon, and showed a progressive decrease in eye 1962; Dent & Kirby-Smith, 1963 ), and H. size through this series. Mitchell & Reddell wallacei (Brandon, 1968). For other species ( 1965) listed eye diameters of several speci­ only general comments or information on mens of T. tridentifera of known body gross eye size are available. For example, lengths and noted that about one-half of the

YoLU:\IE :3:3, NuMBEH 1, jANUARY 1971 9 they apparently are functional under lighted ::t: 0 1- conditions ( Besharse and Brandon, unpub­ 0 0 ~ 30 0 0 lished) and are not degenerate histologi­ 1- • 0 0 0 cally (Lazell and Brandon, 1962: Dent and 5 X X X 0 00 0 (!) oX Kirby-Smith, 1963). ~ 20 • w A • Ontogeny of eye structure in T. spelaeus 1- • • 0.. At. • • represents a special case since the eyes may • z0 A 6. be functional in larvae which live under lighted conditions in springs and spring 10 20 30 40 50 runs, but are not functional in the strictly SNOUT- VENT LENGTH (t.1t.1l cavernicolous adults. The differing selective Figure 7. Number of pterygoid teeth ver­ sus snout-vent length in Haideotriton wal­ pressures on larvae and adults, therefore, lacei (triangles), Typhlomolge rathbuni have resulted in larval eyes which are ( open circles), Typhlomolge tridentifera relatively large, histologically complete and, (X), and Eurycea troglodytes ( solid in all but the largest larvae, perceptive. circles). During and after metamorphosis tissue de­ generation sufficient to cause blindness oc­ eyes lacked lenses; they further noted that curs (Stone, 1964), but the adult eyes still in a series of E. troglodytes all had lenses. are not as degenerate as those of T. rath­ Eye reduction involves both decrease in buni, H. wallacei or P. anguinus. Stone's eye size and histological simplification. ( 1964) observation that some large larvae Among those species living in the Edwards with histologically normal eyes behaved as Plateau of Texas, the progression from larg­ though they were blind suggests that the est to smallest eyes runs from E. latitans initiation of blindness may be extraoptic. to E. troglodytes to T. tridentifera to T. Eye degeneration at metamorphosis in T. rathbuni, but the progression of histologi­ cal degeneration and maximum eye degen­ spelaeus includes changes in the ganglionic eration in this series are not known. and bacillary layers, loss of the outer plexi­ The eyes of G. palleucus are small and form layer, atrophy of the optic nerve, re­ comparable in size to those of E. troglo­ cession of the eyeball, and fusion of the dytes. Although smaller than those of G. eyelids over the eye with accompanying in­ porphyriticus of comparable body length, vasion of the cornea by integumentary cells.

0 0 40 ::t: 0 1-w 0 w 1- 0 30 5 (!) >-a: 0 w 1- xo 0.. 20 0 ••• • ' .. 0 X • • z ' • • •• 0 0 • 0 10 20 30 40 50 60 70 80 90 100 SNOUT- VENT LENGTH

10 THE NATIONAL SPELEOLOGICAL SociETY BcLLETIN Histological studies of the tremendously licts in caves. According to this idea the reduced eyes of P. anguinus, T. rathbuni "regressive" features of troglobites result and H. wallacei suggest arrested develop­ not from subterranean life, but rather from ment of the secondary optic vesicle fol­ great phyletic age and resultant species lowed, possibly, by some degeneration senescence. ( Schlampp, 1892; Hawes, 1946; Brandon, Curiously, reduction of eye structure 1968). These three species seem to differ through the ordinary mechanism of evolu­ somewhat in degree of differentiation prior tionary change, that is, natural selection, to arrest, but ontogenetic changes in eye has generally been considered unimportant, structure have been studied well only in with the notable exception of Barr's ( 1968) P. anguinus. Of these three species H. wal­ analysis. The often elaborate alternative lacei has the smallest, most compact, least explanations have been based on the as­ differentiated eyes (Brandon, 1968), and sumption that well formed eyes are selec­ P. anguinus the largest and most highly tively neutral in a cave-dwelling salaman­ differentiated. The eyes in these salaman­ der. There actually is no reason to assume ders certainly are incapable of image per­ such a thing, and there is considerable rea­ ception and probably are not particularly son to think that such is not true. Mayr sensitive to light intensity either (see ( 1963) convincingly doubted that any gene Hawes, 1946; Pylka & Warren, 1958). Com­ can remain selectively neutral since gene parative examinations of embryonic and actions may be intimately interrelated and young larval eyes to determine the maxi­ far reaching. It seems highly unlikely that mum extent of eye differentiation should be the complex genetic base required for eye made on the most highly specialized troglo­ differentiation and function would have no bitic species. pleiotropic influences on phenotypic suc­ There have been numerous attempts to cess. Rosen ( 1967) argued essentially the outline mechanisms which would explain same point from the view of optimality satisfactorily present-day eye structure in theory. It seems likely that selection would troglobitic salamanders, and in troglobitic operate to reduce eyes in the interest of animals generally. The importance of pre­ energy economy. adaptation usually has been stressed with Eye structure, as well as general body ample justification (see references listed in form, of the most specialized troglobitic Poulson, 1964). Adaptation to some kinds salamanders is largely the result of neotenic of epigean habitats would lead populations trends. Energy conservation resulting from into cave-like habitats and into caves them­ avoidance of metamorphosis may also in­ selves. fluence retention in adults of other body Early explanations of eye reduction in features characteristic of early ontogeny. troglobitic animals were couched in La­ This would be true particularly of struc­ marckian terms of degeneration of unused tures, such as eyes, which apparently have organs. A similar, but more sophisticated no important function in cave habitats, but idea, that eyes in troglobites are selectively whose presence could be detrimental by re­ neutral and in the absence of stabilizing quiring energy for differentiation and by selection are lost through accumulation providing susceptible sites for mechanical of random mutations, has been popular and infectious IDJury. Natural selection (Hubbs, 1938; Poulson, 1964; Deamer, would cause eyes to become less and less 1964). Vandel (1964) has recently sup­ differentiated in adults and cranial features ported the curious idea that reduced eyes to become more embryonic. Hawes, for are a feature of "senile" species which can example ( 1946), has pointed out adult no longer exist among competitors in epi­ chondrocrania} features in P. anguinus gean habitats, but can survive as re- which are embryonic in other salamanders.

VoLUME 33, NuMBER 1, }ANUARY 1971 11 Wake (1966) noted that the orbitosphe­ ally those exposed to sunlight, may be noid, which ossifies in larvae of other ple­ intensely pigmented, whereas larger, cav­ thodontid salamanders and is ossified in ernicolous larvae and postmetamorphic in­ adults of other paedogenetic species, never dividuals are uniformly covered with mela­ does ossify in H. wallacei, T. rathbuni or nophores and the body appears grayish. T. tridentifera. The bodies of E. troglodytes and E. latitans Body Pigmentation both bear faint patterns, the latter more distinctly, but both apparently are less Contrary to even recent statements in the heavily pigmented than larvae of epigean literature that certain troglobitic salaman­ species of Eurycea. ders lack pigment (e. g., Deamer, 1964; There is a positive correlation between Vandel, 1964; Wake, 1966), all known degree of depigmentation and degree of species do have integumentary melano­ specialization to cave habitats reflected in phores and retinal melanin. Integumentary other body features such as neoteny, eye xanthophores can be seen on living H. wal­ reduction, tooth numbers, and increasing lacei, and other species probably possess head size. Poulson ( 1964) attempted to ex­ these as well (see Mohr and Poulson, 1966, plain depigmentation as a secondary, adap­ p. 131). In addition, the testes and vasa tively neutral, result of altered allometric deferentia of adult male G. palleucus, T. growth patterns resulting from selection for rathbuni, T. tridentifera and E. troglodytes slower growth rates. He also commented are marked with melanophores. Peritoneal that as in the case of eyes pigment may be melanophores are present in adult T. spela­ reduced by selection through energy econ­ eus, G. palleucus, T. rathbuni, T. tridenti­ omy. In my opinion the latter is the simpler fera, and E. troglodytes which I have seen, and more likely explanation. but adult specimens of H. wallacei and E. latitans were not available for study. Number of Trunk Vertebrae As a general observation, pigment on the The most highly specialized North Ameri­ body surface seems to have been reduced can troglobitic species have among the more than pigment in the peritoneum. Some fewest trunk vertebrae in the family Ple­ adult T. rathbuni and T. spelaeus, for exam­ thodontidae (Wake, 1966). It also happens ple, are more heavily pigmented internally that among the troglobitic salamanders the than externally. number is directly correlated with degree Of the troglobitic salamanders, some of specialization as judged by other criteria populations of G. palleucus are the most (Table 2). This has led to the conclusion, intensely pigmented externally and are probably unwarranted, that a reduction in marked with dark spots on a moderately number of vertebrae results from troglobitic dark background. Considerable geographic specialization (Baker, 1957; Mitchell & variation exists in this species, and some Reddell, 1965 ) . Wake ( 1966) has pointed populations are fairly pale. At the other out that in the family Plethodontidae a low extreme, the bodies of H. wallacei, T. number of trunk vertebrae is probably a rathbuni and T. tridentifera are covered primitive feature, and that the tendency by relatively few, scattered melanophores toward trunk elongation in a genus such as which form no particular pattern; as a re­ Eurycea represents specialization. In the sult, the skin has a pale, almost translucent genus Eurycea the number of vertebrae appearance. Hemoglobin is especially visi­ ranges from 14 to 22. The fewest are found ble in blood vessels of the gills and limbs in the eastern species E. bislineata, E. of these species (see Mohr and Poulson, aquatica, E. longicauda and E. lucifuga. 1966, pp. 99, 131). Ontogenetic regression The most are found in the neotenic E. of body pigmentation as well as eye struc­ tynerensis which inhabits small, spring-fed ture occurs in T. spelaeus. Larvae, especi- streams in the ozarkian Springfield Plateau,

12 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN TABLE 2. Number of trunk vertebrae in North American troglobitic salamanders and some additional species of Eurycea.

Number of Vertebrae Species 13 14 15 16 17 18 19 20 21 22

Typhlomolge rathbuni 35 tridentifera 3 11 H aideotriton wallacei ...... 18 2 Eurycea troglodytes 2 6 4 latitans ~ 2 1 neotenes ~~ X X pterophila ~ ~ X X nana ~~ X X tynerensis ~ ~ ~ X X X multiplicata ~ ~ ~ X X Typhlotriton spelaeus 2 85 60 2 Gyrinophilus palleucus 25 45 2

0 Some data taken from Smith & Potter, 1946. 00 Data taken from Baker, 1957. 0 0 0 Data taken from Dundee, 1965 a, b. and in E. multiplicata which has a similar The low number of vertebrae may be rel­ but broader range and neotenic tendencies. ictual rather than specialized. The non-troglobitic, paedogenetic species of The two troglobites with the broadest Eurycea in the Edwards Plateau of Texas geographic ranges, T. spelaeus and G. pal­ are intermediate in number between the leucus, show geographic variation in num­ ozarkian species and the Edwards Plateau ber of trunk vertebrae. The range of num­ troglobites. ber in G. palleucus is the same as in its It may be that the low number of verte­ epigean congener G. porphyriticus (Bran­ brae in T. rathbuni, T. tridentifera and H. don, 1966a). Geographic variation in T. wallacei [the latter is definitely similar to spelaeus is clinal ( Brandon, 1966b ) , and species of Eurycea and closely related to seems to be clinal also in G. palleucus them (Wake, 1966)] has nothing to do (Brandon, 1965b and unpublished data) with adaptation to cave habitats, but rather although some populations may be isolated. indicates that they arose from a primitive Broad, Flattened Heads Eurycea stock. Some of the troglobitic salamanders char­ The number of trunk vertebrae in the acteristically have proportionally larger series from T. rathbuni to T. spelaeus in heads than most salamanders have, and in Table 2 may indicate the relative time se­ addition the snouts are elongate, flattened, quence of isolation and differentiation of broadened and truncate. The most extreme the various forms from an ancestral stock example of this is in T. rathbuni, and to a gradually increasing in vertebral number. progressively decreasing degree it occurs

VoLUME 33, NuMBER 1, }ANUARY 1971 13 in T. tridentifera, G. palleucus, H. wallacei, involves altered allometric growth patterns. T. spelaeus, E. latitans and E. troglodytes There are differences in relative growth (see profile sketches in Mitchell & Reddell, rate of the head in different troglobitic sala­ 1965, and Brandon, 1966a). Flattening and manders, as indicated by differences in broadening of the head are not restricted slopes of regression lines fitted to plots of among salamanders to cave-adapted species, head width and length versus body length but are found also in such bottom dwellers (Table 3). Generally, the head increases as the mudpuppy (Necturus spp.) and the in size relatively faster ontogenetically in hellbender ( Cryptobranchus alleganiensis). the more highly specialized species. Thus, Among fishes and amphibians living as bot­ the head appears particularly large in large, tom dwellers in still or slowly moving adult individuals of T. rathbuni and G. p. waters, broadening and flattening of the gulolineatus. There are rather great differ­ head are rather common. ences in relative growth rate of the head The altered head shape, by providing a in the subspecies of G. palleucus. broader gape, may result in more efficient Attenuation of Limbs feeding movements. In addition, Walters One of the most striking morphological and Walters ( 1965), extrapolating from the features of T. rathbuni is its long, slender relation between head size and lateral line legs. Comparative data on leg attenuation development in amblyopsid cave fishes and elongation in other troglobitic sala­ (Poulson, 1963), have suggested that the manders, however, are not easily obtained. broad, blunt snout in cavernicolous popu­ Usually elongation has been expressed as lations of the poeciliid fish Poecilia sphe­ the ratio between body length and leg nops is a structural adaptation to facilitate length either by direct measurements or by nonvisual navigation. Broad heads of trog­ lobitic salamanders probably also result in overlapping of appressed legs. Such infor­ more efficient functioning of neuromasts. mation, however, could be used just as well The importance of neuromasts in locating to show shortening of the body rather than aquatic prey under conditions of total dark­ elongation of the legs. Since troglobitic ness may also have been a strong induce­ salamanders differ in number of trunk ver­ ment in selection for neoteny. tebrae from 13 to 20, and therefore differ Poulson (1963) has shown that the in body elongation, some sort of adjustment mechanism bringing about a proportional ought to be made when comparing leg increase in head size in troglobitic fishes lengths between species. If 13 trunk verte-

TABLE 3. Slopes of regression lines, calculated by the least squares method, for plots of head width and head length against body length in North American troglobitic salamanders.

Head Head Width Length Species b b ----- 4 Typhlomolge tridentifera ...... 0.244 0.356 Typhlomolge rathbuni ...... 0.203 0.319 Gyrinophilus p. gulolineatus ...... 0.200 0.258 H aideotriton wallacei ...... 0.190 0.229 Gyrinophilus p. necturoides ...... 0.161 0.235 Typhlotriton spelaeus ...... 0.147 0.244 Gyrinophilus p. palleucus ...... 0.134 0.229 Eurycea troglodytes ...... 0.118 0.138

0 Measurements taken directly from Mitchell & Reddell, 1965.

14 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN brae are considered a standard body length, The legs of H. wallacei, T. tridentifera then leg length can be compared more and T. rathbuni have the appearance of be­ meaningfully among the species as the ratio ing rather elongated partly because their of leg length to the "adjusted" body length, bodies are short, but also because the legs because variation in length of vertebrae is are attenuated. At small body sizes E. trog­ considerably less than variation in body lodytes actually has proportionally longer length. "Adjusted body length" is the por­ legs than the more highly specialized T. tion of the total snout-vent measurement tridentifera; but those of the latter are more which results from the head and 13 trunk slender. The differences between the vari­ vertebrae. ous troglobitic species in relative leg length Even when comparisons are based on ad­ are most striking when large specimens are justed body lengths, T. rathbuni, at all size compared, because of allometric growth classes, has the longest legs (Figure 9). differences (Table 4). The remarkably Those of G. p. gulolineatus are second elongated legs of T. rathbuni, especially of longest and are considerably longer than larger specimens, result from a high rela­ those of G. porphyriticus and other subspe­ tive growth rate; they appear even more cies of G. palleucus. These data, and others remarkable because they are attenuate. not elaborated here, suggest that G. p. gulo­ Legs of H. wallacei and T. tridentifera are lineattts may actually be a separate species. also noticeably attenuate. While there is a big difference between G. p. gulolineatus and the other nominal A. subspecies of G. pal­ leucus in relative leg growth, as well as in B. actual leg length, those of G. p. gulolinea­ C. tus are not attenuated. E. latitans is puz­ zling; its legs are short, but they show D. surprisingly high positive allometry. E. The most likely function of elongated F. limbs is to raise the body, and particularly G. the head, above the substrate (Poulson, 1964; Wake, 1966). The excellent photo­ H. graphs in Herald ( 1952), Mitchell & Red­ dell ( 1965) and Mohr & Poulson ( 1966) 5 5 10 5 10 15 AT 20MM ABL AT 30MM ABL AT 40MM ABL show how T. rathbuni and T. tridentifera stand with their heads lifted high above HIND LEG LENGTH the substrate. (MM) The slow, intermittent loco­ Figure 9. Comparison of hind leg length motion, with body raised, permits greater at three different body lengths in North efficiency in detection of moving prey by American troglobitic salamanders. ABL the neuromast system. Poulson ( 1964) also refers to "adjusted body length" as de­ pointed out that longer limbs permit the fined in the text. Values of leg length are salamanders to search a larger area per taken from regression lines fitted to plots unit of energy expended. of hind leg length versus "adjusted body length". Typhlomolge rathbuni (A), ORIGIN OF Gyrinophilus Palleucus Gyrinophilus palleucus gulolineatus (B), The primitive eastern North American Typhlomolge tridentifera (C), H aideo­ genus Gyrinophilus contains two polytypic triton wallacei (D), Typhlotriton spelaeus species, the epigean G. porphyriticus and ( E ) , Eurycea troglodytes (F), Gyrinophi­ the troglobitic G. palleucus. Eurycea is the lus palleucus palleucus and G. p. nectur­ only other genus of salamanders known to oides (G), and Eurycea latitans (H). contain both epigean and troglobitic species.

YoLUME 33, NuMBER 1, }ANUARY 1971 15 TABLE 4. Slopes of regression lines, calculated by the least squares method, for plots of hind leg length against "adjusted" body length in North American troglobitic salamanders.

Hind Leg Length Species b

Typhlomolge rathbuni ...... 0.359 H aideotriton wallacei ...... 0.329 0 Eurycea latitans ...... 0.291 0 0 Typhlomolge tridentifera ...... •.. 0.287 Typhlotriton spelaeus (larvae) ...... 0.215 Gyrinophilus p. gulolineatus ...... 0.211 Eurycea troglodytes ...... 0.190 Gyrinophilus p. palleucus and G. p. necturoides ...... 0.155

0 Data from Baker, 1957. 00 Some data taken from Mitchell & Reddell, 1965.

G. porphyriticus is a metamorphosing spe­ from two localities in the Ridge and Valley cies ranging from eastern Quebec and Province of McMinn and Roane counties, western Maine through the Appalachian Tennessee; and several unidentified popula­ uplift to the Fall Line in Georgia, Alabama, tions are known from northwestern Georgia and Mississippi (Figure 10). It shows con­ (Cooper, 1968), northern Alabama, and siderable geographic variation in various central Tennessee. The full range of this traits, most noticeably in body pigmenta­ species is not well known, and several tion, and four subspecies are currently rec­ workers continue to discover new localities. ognized. Its variation and systematics re­ Gyrinophilus palleucus differs from adult cently have been studied in some detail G. porphyriticus by being paedogenetic, and (Brandon, 1966a). A second neotenic from its larvae by having smaller eyes, a species, G. lutescens ( Rafinesque), was re­ broader head, a more spatulate snout, a ported by Mittleman ( 1942 ) ; it is not a different pattern of body pigmentation and distinct species, however, but was based on slightly higher tooth counts. In other gen­ larvae of one subspecies of G. porphyriticus eral body features it is similar to larval G. (Newcomer, 1960; Brandon, 19'63). porphyriticus. Specimens which have under­ Although the available sample is rela­ gone nearly complete metamorphosis during tively small, considerable geographic varia­ thyroxin treatment would not be confused tion is evident in G. palleucus, and three with adult G. porphyriticus; body pign1en­ subspecies are currently recognized (Bran­ tation, head proportions, and certain cranial don, 1966a; 1967b). They differ in body features remain different. For this reason pigmentation, head width, leg length, eye some workers have considered placing pal­ size, and modal number of trunk vertebrae. leucus in a separate genus, but there seems About 125 specimens of G. p. palleucus are little question that palleucus and porphyri­ known to me from five caves in the Crow ticus are closely related. G. palleucus un­ Creek drainage of Franklin County, Ten­ questionably is modified for permanent nessee; 25 of G. p. necturoides from one cave dwelling, and is considered taxonomic­ cave in the Elk River drainage of Grundy ally distinct from G. porphyriticus. There County, Tennessee; 17 intergrades between is little information bearing on the degree the first two subspecies from three caves in of reproductive isolation between the two the North Sauty Creek drainage of Jackson species, but such isolation has been as­ County, Alabama; 13 of G. p. gulolineatus sumed to exist. Detailed observations by

16 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN ·--·-\_ ---- ;--

/ j ·.J

EZ> ~ polleucus 0 Q.porphyriticus MILES 50 0 100 200

Figure 10. Distribution of Gyrinophilus porphyriticus and Gyrinophilus palleucus.

Cooper anJ Cooper ( 1968) on ranges and known about the reproductive ecology of habitats of the two species in Alabama either species. show them not to be sympatric there, and Mechanisms involved in the origin of leave open the possibility that reproductive troglobitic species have long been subject isolation results only from geographic or to discussion by biologists. G. palleucus habitat isolation. No tests of hybridization and its epigean relative are sufficiently well have been attempted yet, and little is known to allow consideration of some fac-

YoLU;'.IE :33, NuMBER 1, }ANUARY 1971 17 tors involved in the ongm and modification size, played an important part in the inva­ of a cave dwelling form. sion of caves by G. palleucus. Little is known of the evolution of ple­ Neoteny is not known to occur in any thodontid salamanders by direct evidence; population of G. porphyriticus, even though fossil remains are few, and there are none this species is a common part of the cave of Gyrinophilus. Those fossils which are entrance fauna in many parts of its range. known do not provide sufficient material for Adults, and especially larvae, are found phylogenetic considerations (but see Estes, deep within caves, but they differ in no 1965; Wake, 1966). Wake thoroughly way from individuals collected in epigean discussed the evolution of plethodontid situations in the same areas. The larval genera, primarily on the basis of osteologi­ period in G. porphyriticus, probably three cal evidence. years (Bishop, 1941 ) , is the longest of any It seems likely that G. palleucus evolved plethodontid salamander. It would thus from an epigean, metamorphosing ancestor, lend itself readily to development of neo­ possibly from G. porphyriticus. It is impos­ teny by selection for slower developmental sible to judge the age of G. palleucus as a and growth rates in response to a generally species, but compared with other troglobitic low food supply in caves. Comparisons of salamanders it seems to be a relatively re­ developmental growth rates of these two cent entrant into cave habitats. The present species, however, are not yet available. In­ cave systems in Tennessee are estimated to deed, extremely little information of this be of Pliocene to Pleistocene age (Barr, nature is available for either species. The 1961 ). most obvious biotic difference between epi­ The known range of G. palleucus lies at gean stream and spring habitats where G. the periphery of the present-day range of porphyriticus occurs and G. palleucus cave G. porphyriticus. Climatic conditions dur­ streams is in abundance of potential food. ing the Pleistocene may have resulted in the G. porphyriticus is aggressively carnivorous, physical isolation of peripheral populations even cannibalistic (Bishop, 1941). Many of G. porphyriticus and their subsequent specimens examined by radiography have adaptation to habitats in and around caves, salamanders in their stomachs. Larvae and and finally in restriction to cave habitats as adults seem to feed primarily on a diverse climatic conditions changed. fauna of aquatic invertebrates supple­ The present-day ranges of G. palleucus mented by smaller salamanders and some and G. porphyriticus appear to overlap in terrestrial invertebrates. Stomachs of G. eastern Tennessee, but the two species have palleucus contain at most a few isopods and never been found in the same cave, or even few other invertebrates (Brandon, 1967a). in the same vicinity. In northern Alabama The aquatic invertebrate fauna in their the two species appear to be parapatric caves is neither diverse nor abundant, and (Cooper and Cooper, 1968). Cooper and stomachs of many specimens are empty. Cooper ( 1968) recorded ecological differ­ Cooper and Cooper's ( 1968) observa­ ences between G. palleucus caves and G. tions of differences between feeding habits porphyriticus caves and suggested that the of a larval G. porphyriticus and a G. pal­ first species would have a competetive ad­ leucus under laboratory conditions are prob­ vantage over the second in cave habitats. ably not as significant as they seem. Feed­ The geographic distribution of both species ing behavior in young salamanders is fairly in Tennessee and northwestern Georgia is labile, and the reactions of captive speci­ still rather poorly known, but information mens to food probably reflect their histories is accumulating (Cooper, 1968). of conditioning to natural sources of food. It is certain that preadaptation, especi­ Striking examples of conditioning to unu­ ally in regard to neoteny, ecology, and eye sual foods or methods of introduction have

18 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN been observed in laboratory colonies of width, leg length, number of trunk verte­ other salamanders (Brandon, pers. observ.). brae, eye diameter, and perhaps in adult Of 12 live G. palleucus kept in my labora­ body size could be the result of more recent tory for a year, some fed immediately on local selective pressures, or of genetic beef liver and earthworms, whereas others drift in partially or completely isolated even now feed on these foods only spar­ populations. ingly and hesitatingly. In general, individ­ This article is based on a paper presented uals of G. palleucus eat less (and grab it at the meetings of the American Society of less voraciously) than do larval G. porphy­ Zoologists in Knoxville, Tennessee, Decem­ riticus and perhaps have a lower metabolic ber, 1964. rate (note Cooper and Cooper's comparison For the loan of specimens, I am grateful of heart-beat rates-1968, p. 23). to Gerald G. Raun and Charles L. Douglas There is considerable geographic varia­ (Texas Natural History Collection), James tion in eye size in both G. palleucus and G. A. Peters (United States National Mu­ porphyriticus, but the eyes of the former seum), Hemy M. Stevenson (Florida State are consistently smaller (Brandon, 1966a). University), Ernest E. Williams (Museum Within G. porphyriticus, Canadian larvae of Comparative Zoology), Robert F. Inger have the largest eyes, while those from the (Field Museum of Natural History), Ho­ southern Appalachian mountains and Pied­ bart M. Smith ( University of Illinois mont have the smallest. If G. porphyriticus Museum of Natural History), Richard G. is the parent species, G. palleucus would Zweifel (American Museum of Natural His­ have developed from these southern popu­ tory) and Laurence M. Hardy (University lations which have the smallest eyes. Sub­ of Kansas Museum of Natural History ) . sequent selection for further eye reduction Some specimens and suggestions were is likely. kindly provided by Thomas L. Poulson, Differences among the nominal subspecies Ronald G. Altig, Enoch H. Albert, Thomas of G. palleucus in body pigmentation, head C. Barr, and John E. Cooper.

LITERATURE CITED Alt. A. 1910. On the histology of the eye 1944. A new neotenic plethodont of Typhlotriton spelaeus from Marble salamander, with notes on related species. Cave, Mo. Trans. Acad. Sci. St. Louis, Copeia, 1944(1):1-5. 19:83-96. Brame, A. H., Jr. 1967. A list of the Baker, J. K. 1957. Eurycea troglodytes: a world's recent and fossil salamanders. new blind cave salamander from Texas. Herpeton, 2 ( 1) : 1-26. Texas Jour. Sci., 9( 3) :328-336. Brandon, R. A. 1963. Triturus (Gyrino­ Barr, T. C., Jr. 1960. Introduction To: philus) lutescens Rafinesque, 1832 (Am­ Symposium: Speciation and raciation in phibia, caudata) : proposed suppression cavernicoles. Amer. Midi. Natur., 64:1-9. under the plenary powers. Bull. Zool. 1961. Caves of Tennessee. Bull. Nomencl., 20:210-211. 64, Tennessee Div. Geol., 567 pp. 1963. Ecological classification of 1965a. Morphological variation cavernicoles. Cave Notes, 5 ( 2) :9-12. and ecology of the salamander Phaeo­ 1968. Cave ecology and the evo­ gnathus hubrichti. Copeia, 1965(1):67-71. lution of troglobites. In Evolutionary 1965b. A new race of the neo­ Biology, vol. 2:35-102. Appleton-Century­ tenic salamander Gyrinophilus palleucus. Crofts, N. Y. Copeia, 1965 ( 3) :346-352. Bishop, S. C. 1941. The salamanders of 1966a. Systematics of the sala­ New York. New York State Mus. Bull. mander genus Gyrinophilus. Illinois Bioi. 324, 365 pp. Monogr., 35:1-86.

VoLUME 33, NuMBER 1, JANUARY 1971 19 1966b. A reevaluation of the sta­ metamorphic agent. Science, 135 ( 3508) : tus of the salamander, Typhlotriton nereus 1060-1061. Bishop. Copeia, 1966( 3) :555-561. 1965a. Eurycea multiplicata. In 1967a. Food and an intestinal Catalogue of American Amphibians and parasite of the troglobitic salamander Reptiles, American Society of Ichthyolo­ Gyrinophilus palleucus necturoides. Her­ gists and Herpetologists, p. 21. petologica, 23( 1) :52-53. 1965b. Eurycea tynerensis. In ---. 1967b. Gyrinophilus palleucus. In Catalogue of American Amphibians and Catalogue of American Amphibians and Reptiles, American Society of Ichthyolo­ Reptiles, American Society of Ichthyolo­ gists and Herpetologists, p. 22. gists and Herpetologists, p. 32. Dundee, H. A. and A. Gorbman. 1960. 1968. Structure of the eye of Utilization of radioiodine by thyroid of H aideotriton wallacei, a North American neotenic salamander, Eurycea tynerensis. troglobitic salamander. Jour. Morph., Physiol. Zool., 33:58-63. 124 ( 3) : 345-351. Brown, B. C. 1967. Eurycea latitans. In Dunn, E. R. 1926. The salamanders of Catalogue of American Amphibians and the family Plethodontidae. Smith College Reptiles, American Society of Ichthyolo­ 50th Anniv. Publ., Northampton, Massa­ gists and Herpetologists, p. 34. chusetts, 441 pp. Carr, A. F., Jr. 1939. Haideotriton wal­ Eigenmann, C. H. 1900. The eyes of the lacei, a new subterranean salamander blind vertebrates of North America. II: from Georgia. Occ. Papers Boston Soc. The eyes of Typhlomolge rathbuni Stej­ Natur. Hist., 8:333-336. neger. Trans. Amer. Micros. Soc., 21:49- Conant, R. 1958. A field guide to reptiles 61. and amphibians of the United States and Eigenmann, C. H. and W. A. Denny. Canada east of the lOOth meridian. 1900. The eyes of the blind vertebrates Houghton Miffiin Co., Boston, 366 pp. of North America. Ill: The structure and Cooper, J. E. 1968. The salamander ontogenetic degeneration of the eyes of Gyrinophilus palleucus in Georgia with the Missouri cave salamander, an account notes on Alabama and Tennessee popu­ based on material collected with a grant lations. J. Alabama Acad. Sci., 39:182-185. from the Elizabeth Thompson Science Cooper, J. E. and M. R. Cooper. 1968. Fund. Bioi. Bull., 2:33-41. Cave-associated herpetozoa II: Sala­ Emerson, E. T. 1905. General anatomy manders of the genus Gyrinophilus in of Typhlomolge rathbuni. Proc. Boston Alabama caves. Bull. Natl. Speleol. Soc., Soc. Natur. History, 32:43-76. 30( 2): 19-24. Estes, R. 1965. Fossil salamanders and Deamer, D. W. 1964. Entropy and cave salamander origins. Amer. Zool., 5:319- animals. Ohio Jour. Sci., 64( 3) :221-223. 334. Dent, J. N. and J. S. Kirby-Smith. 1963. Gorbman, A. 1957. The thyroid gland of Metamorphic physiology and morphology Typhlomolge rathbuni. Copeia, 1957 ( 1): of the cave salamander Gyrinophilus pal­ 41-43. leucus. Copeia, 1936(1): 119-130. Hawes, R. S. 1946. On the eyes and Dent, }. N., J. S. Kirby-Smith, and D. L. reactions Craig. 1955. Induction of metamor­ to light of Proteus anguinus. phosis in Gyrinophilus palleucus. Anat. Quart. Jour. Micros. Sci., 86:1-53. Rec., 121:429. Herald, E. S. 1952. Texas blind sala­ Dundee, H. A. 1957. Partial metamor­ mander in the aquarium. Aquarium Jour., phosis induced in Typhlomolge rathbuni. 23( 8): 149-152. Copeia, 1957( 1) :52-53. Hubbs, C. L. 1938. Fishes from the 1962. Response of the neotenic caves of Yucatan. Carnegie lnst. Wash. salamander H aideo triton wallacei to a Publ. 491:261-295.

20 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIK Humphrey, R. H. 1922. The multiple Pylka, J. W. and R. D. Warren. 1958. A testis in urodeles. Bioi. Bull., 43( 1) :45- population of Haideotriton in Florida. 67. Copeia, 1958 ( 4 ) :334-336. Kezer, J. 1952. Thyroxin induced meta­ Rafinesque, C. S. 1832. On three new morphosis of the neotenic salamanders salamanders of Kentucky. Atlantic Jour., Eurycea tynerensis and Eurycea neotenes. 1 ( 3): 121. Copeia, 1952 ( 4) :234-237. Rosen, R. 1967. Optimality principles in Lazell, J. D., Jr. and R. A. Brandon. 1962. biology. Butterworth and Co., London, A new stygian salamander from the 198 pp. southern Cumberland Plateau. Copeia, Saylor, A. 1966. The reproductive ecol­ 1962( 2) :300-306. ogy of the red-backed salamander, ;\1ayr, E. 1963. Animal species and evo­ Plethodon cinereus, in Maryland. Copeia, lution. Harvard Univ. Press, Cambridge, 1966( 2): 183-193. Massachusetts, 797 pp. Schlampp, K. W. 1892. Das Auge des Mitchell, R. W. and J. R. Reddell. 1965. Grottenolmes (Proteus anguineus). Zeit. Eurycea tridentifera, a new species of wiss. Zool., 53:537-557. troglobitic salamander from Texas and a Schreiber, G. 1933. Prove funzionali sulla reclassification of Typhlomolge rathbuni. tiroide del Proteo. Bull. Soc. Ital. Biol. Texas Jour. Sci., 17 ( 1): 12-27. Sper., 8:1-4. Mittleman, M. B. 1942. Notes on sala­ Smith, H. M. and L. E. Potter, Jr. 1946. manders of the genus Gyrinophilus. Proc. A third neotenic salamander of the genus New England Zool. Club, 20:25-42. Eurycea from Texas. Herpetologica, 3(4): 1950. Cavern-dwelling salaman­ 105-109. ders of the Ozark Plateau. Bull. Natl. Smith, P. W. 1948. Food habits of cave Speleol. Soc., ( 12): 12-15. dwelling amphibians. Herpetologica, 4: Mohr, C. E. and T. L. Poulson. 1966. 205-208. The life of the cave. McGraw-Hill Book Stejneger, L. 1896. Description Co., New York, 232 pp. of a new genus and species of blind tailed Newcomer, R. J. 1961. Investigations into batrachians from the subterranean the status of Gyrinophilus lutescens ( Raf­ waters of Texas. Proc. U. S. inesque). ASB Bull., 8( 2) :21. Natl. Mus., 18:619- 621. Noble, G. K. 1927. Creatures of pP.r­ petual night. N atur. Hist., 27 ( 5) :405- Stone, L. S. 1964. The structure and 419. visual function of the eye of larval and Organ, J. A. 1961. Studies of the local adult cave salamanders Typhlotriton distribution, life history, and population spelaeus. Jour. Exp. Zool., 156:201-218. dynamics of the salamander genus Des­ Valentine, B. D. 1964. The external mor­ mognathus in Virginia. Ecol Monogr., phology of the plethodontid salamander, 31: 189-220. Haideotriton. Jour. Ohio Herpetol. Soc., Potter, G. E. and E. L. Rabb. 1959. Thy­ 4:99-102. roxin induced metamorphosis in a neo­ Vande!, A. 1964. Biospeologie, la biologie tenic salamander, Eurycea nana Bishop. des animaux cavernicoles. Gauthier-V il­ Zool. Ser. A. & M. College of Texas, lars, Paris, 619 pp. 1(1): 1-12. Wake, D. B. 1966. Comparative osteology Poulson, T. L. 1963. Cave adaptation in and evolution of the lungless salamanders, amblyopsid fishes. Amer. Midl. Natur., family Plethodontidae. Mem. Southern 70(2) :257-290. California Acad. Sci., 4:1-111. ---. 1964. Animals in aquatic environ­ Walters, L. H. and V. Walters. 1965. ments: animals in caves. In Handbook Laboratory observations on a cavernicolous of Physiology: Adaptation to the Environ­ poeciliid from Tabasco, Mexico. Copeia, ment, chapter 47:749-771. 1965( 2) :214-223. Manuscript received by editors June 1970

VoLUME 33, NuMBER 1, JANUARY 1971 21

The Invertebrate Cave Fauna of Georgia

0 By John R. Holsinger and Stewart B. Peck H

ABSTRACT During the early summer of 1967, extensive biological field work was carried out in the caves of northwestern Georgia. Prior to that time the biology of Georgia caves was poorly known. Speleologically, Georgia can be divided into the Appalachian region in the northwest and the Coastal Plain region in the southwest. The majority of caves occur in the Paleozoic limestones of the Appalachians, although a few are found in the early Tertiary limestones of the Coastal Plain. The data obtained from the collection of invertebrates from 29 caves in Georgia are listed systematically, and a brief discussion of the zoogeographic relationships of the troglobitic forms (obligatory cavernicoles) is given. Based on troglobite distribution, two faunal units are recognized in northwestern Georgia: the Appalachian Plateau and the Appalachian Valley. The former unit is divided into two faunal subunits. Approximately 130 species of invertebrates are recorded from Georgia caves and between 24 and 27 of them are troglobites. Major animal groups represented by troglobites in Georgia include planarians, snails, isopods, amphipods, crayfish, pseudoscorpions, spiders, millipedes, diplurans, collembolans, and beetles. The invertebrate cave fauna of Georgia also includes a large number of troglophiles (facultative cavernicoles), with about 25 genera being represented.

INTRODUCTION other purposes: (a) provide the necessary The major objective of this study is to groundwork for future studies in the ecol­ give a preliminary annotated listing of all ogy and population biology of the caverni­ known species of invertebrate cavernicoles coles of the area, (b) act as a starting point of Georgia based principally on field work for more detailed studies on the zoogeog­ in June 1967. Moreover, all of the literature raphy and evolutionary biology of the cave pertinent to Georgia cave invertebrates was fauna of the state as well as for the south­ reviewed and incorporated into the list. ern Appalachians, and (c) bring Georgia Finally, a few new records, obtained by into line with other major cave-bearing John and Ylartha Cooper, Tony Iles, and states with regard to the knowledge of its Arthur Dobson subsequent to June 1967, kinds of cave organisms. are also included in the list. In addition to Prior to June of 1967, little biological being a guide to the invertebrate fauna in work had been done in the caves of Geor­ Georgia caves, this paper should serve three gia. In contrast to other southeastern states

0 Dept. of Biology, Old Dominion University, with major cave and karst regions, the Norfolk, Virginia 23508. knowledge of Georgia cave fauna was mea­ 00 Museum of Comparative Zoology, Harvard ger. Although a few scattered collections University, Cambridge, Massachusetts 02138. Present address: Dept. of Biology, Carleton Univ., Ottawa, had been made in Georgia caves, no serious Canada. attempt had been made to systematically

VoLU:\IE .'33, NuMBER 1, }ANUARY 1971 2:3 investigate or sample the cavernicolous spe­ work of 1967. Two caves were also investi­ cies of this potentially significant area. gated in the Coastal Plain at another time Previous collecting in Georgia caves had by one of us (SBP). been sporadic, although over the years a Twenty-nine caves are covered in this few caves were visited biologically by E. study, 27 of which are located in the Appa­ Ackerly, T . Barr, K. Dearolf, L. Hubricht, lachians and two of which are located in W. Jones, C. Mohr, and J. Valentine. More the Coastal Plain. Figure 1, an outline map recent such visits were made by J. and M. of Georgia, shows the location of the two Cooper, A. Iles, and A. Dobson. A number Coastal Plain caves, and Figure 2, a map of important studies on cavernicoles have of the northwestern corner of Georgia, given cursory mention to Georgia cave shows the location of 25 of the 27 caves in forms, including papers on millipedes by the Appalachians. During June 1967, 18 of Loomis ( 1939, 1943) and Chamberlin the caves located in Figure 2 were biologi­ ( 1946), amphipods by Hubricht ( 1943) cally explored by the writers and their as­ and Holsinger ( 1969), planarians by Hy­ sistants; two other caves located in Figure man ( 1954 ) , beetles by Barr ( 1965) , and 2 were visited and similarly explored by the general checklists by Dearolf ( 1953) and writers prior to 1967. The remaining caves Nicholas ( 1960). Most of these papers, located in Figure 2 were investigated by however, covered large areas of the Appa­ Tony lies. Three other caves (Saw Mill, lachians, with only one or two caves in Cricket, and Creek Bed caves), all presum­ Georgia being included. ably located near Rising Fawn, have been This report is concerned with the inverte­ mentioned previously in the literature of brate cave fauna of Georgia and should Georgia cave faunas (see Dearolf, 1953). serve as a supplement to the biospeleologi­ These caves could not be identified during cal data already published on other impor­ tant cave states in the eastern United States. Currently, a number of state cave faunal surveys are in progress or have been published, at least partially. These include

Alabama (Peck and Peck, 1967; Cooper, ,.. ATLANTA Cooper and Peck, several papers in prepa­ ration); Florida (Warren, 1961; Peck, 1970); Texas (Reddell, 1965, 1966, 1967); and Virginia and West Virginia (Holsinger, 1963a, 1963b, and in preparation). More general treatments of state cave faunas are those of Tennessee ( Barr, 1961 ) , Virginia • GR JERS CAVE (Holsinger, 1964), Pennsylvania (Holsinger, 1971), and Maryland (Franz, in prepara­ tion). Two widely separated and geologically • CLI MAX CAVE different areas of Georgia contain limestone caves- the Appalachian Plateau and Valley in the northwestern corner, and the Coastal FIGURE 1. Map of Georgia showing the Plain in the southwest. The former area, Appalachian Valley and Plateau region situated in the "cave rich" Appalachians, is (shaded) and two caves located in the by far the most extensive cavernous region Coastal Plain. See Fig. 2 for an enlarged of the two and was, consequently, chosen map showing locations of the ca,·es in the as the target for the early summer field Appalachians.

24 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN _/ / / I { 1 ( • I ( ;(ol ( 4:{ \ ~I 11 n !/ - / I .. :; - / J .::; ' // I .LAFAYETTE I -~ I ) ,/' , , I I ' 17/ \ i v i ii /! ' / , vI :.•: I ) ; ' \i...... ____) I , I - I__ f '/ i\ 19 1 I . I !

•••

MILt I ••• 10

FIGURE 2, Map of the Appalachian Valley and Plateau region of northwestern Georgia showing caves sampled for invertebrate fauna, The names of the caves, keyed to the numbers on the map, are listed in the text,

VoLUME 33, NuMBEH 1, JANUARY 1971 25 the field work and are not found in the County ( 1 cave): 0 Roberts ( 20). Bartow standardized listing of Georgia caves by the County ( 2 caves): ° Kingston Saltpetre Georgia Speleological Survey. There is good (21) and 0 Yarborough (22). Polk County reason to believe, however, that Cricket and ( 2 caves) : Deatons ( 23) and 0 White Creek Bed caves were names given to the River ( 24). Floyd County ( 1 cave): ° Cave two entrances to what is now called Byers Springs ( 25). Cave and that Saw Mill Cave could have been another name for Sittons Cave. AREAL GEOLOGY The caverniferous limestones of north­ Each cave visited in June 1967 was western Georgia occur in two physiographic thoroughly searched for invertebrates. The provinces, the Appalachian Plateau and investigation was started just inside of the the Appalachian Valley (Fig. 2). Although entrance, and all potential habitats ( i. e., there is no clear-cut line dividing these two damp clay and silt, decaying organic debris, provinces, the eastern flanks of Lookout damp rotting wood, pools, streams, etc.) Mountain and its associated outlier, Pigeon were carefully checked throughout the Mountain, are often regarded as the front traversable part of each cave. Bait in the of the Appalachian Plateau in this area. form of liver and carrion was left in some A sequence of cavernous limestones (viz., of the caves and checked on a return visit Bangor, Gasper, St. Genevieve, and several weeks later. St. Louis) occurs along the eastern flanks of A list of the caves located on the map in Lookout and Pigeon Mountains and dips Figure 2 is given below. The numbers as­ gently to the west. These same limestones, signed to these caves are keyed to the loca­ which extend westward, are responsible for tions shown on the map. Those caves in the the numerous caves developed in extreme list denoted by asterisks were visited by northwestern Georgia (Dade County) and the writers. Some of these caves were either northern Alabama (Jackson, ~Iadison, and new to the Georgia Speleological Survey or Marshall counties, etc.). To the east of the were at best inadequately known. As a plateau front are a series of low ridges and service to the Georgia Speleological Survey, wide valleys, which compose the Appalach­ Roger Baroody, who assisted with the field ian Valley and contain exposures of sedi­ work, recorded physical data on all caves mentary rocks that vary in geologic age visited and subsequently turned over these from Cambrian to Pennsylvanian. Several of data to Richard Schreiber for deposition in these valleys are floored by Cambrian and the files of the Survey. The caves, arranged Ordovician limestones and dolomites and by county, follow. are intermittently caverniferous. The exten­ Dade County ( 12 caves) : Boxcar ( 1 ) , sive karst areas found farther north in the 0 Byers ( 2), Case ( 3), 0 Howards Water­ Appalachian Valley sections of Tennessee, fall ( 4), Hurricane ( 5), 0 Johnson Crook Virginia, and West Virginia are virtually No. 1 (6), 0 Johnson Crook No. 2 (7), non-existent in Georgia, and the limestone 0 Morrison and 0 Morrison Spring (separate terrane is usually inconspicuous. caves located ~4 mile apart) ( 8), Running The cave region of northwestern Georgia Water ( 9), Rustys (1 0), and Sittons ( 11). is drained by two major river systems. Walker County ( 7 caves): 0 Bible Springs Streams located north and northwest of La ( 12), ° Cave Spring ( 13), 0 Harrisburg Fayette flow into the Tennessee River, and ( 14), 0 Horseshoe ( 15), 0 Mtn. Cove Farm streams located east, south and southeast of ( 16), 0 Pettijohn ( 17), and Hickman La Fayette flow into the Coosa River which Gulf (located 4 miles NNW of Bible is part of the Mobile Basin drainage and Springs Cave but not shown in figure 2). ultimately joins the Alabama River. Chatooga County ( 2 caves): 0 Blowing The greatest concentration of caves in the Springs ( 18), and 0 Parker ( 19). Gordon Appalachian region of Georgia is along the

26 THE NATIONAL SPELEOLOGICAL SociETY BL-LLETIN eastern flank of Pigeon Mountain and in the lobites-obligatory cave species which are outcrops of Mississippian limestones west morphologically specialized for, and re­ of Lookout Mountain in Dade County. Most stricted to, cave habitats and are unable to of the caves in the Appalachian Valley are survive on the surface; ( 2) troglophiles­ developed in various limestone and dolomite facultative cave species which frequently members of the Knox group. In the com­ inhabit caves and complete their life cycles puter print-out list issued by the Georgia there but may occupy ecologically suitable Speleological Survey in 1967 approximately habitats on the surface; ( 3) trogloxenes­ 124 caves were recorded. Undoubtedly species which often occur in caves but fre­ many more caves exist, and significant dis­ quently return to the surface or near the coveries, such as the one mentioned below, surface ( i. e., cave entrance zone) for food; will be made there in the future. In late ( 4) accidentals-species which accidentally 1968, an exhaustive exploration of Ellisons wander, fall, or are washed into caves and Cave, located on the eastern side of Pigeon can exist there only temporarily. The eco­ Mountain in Walker County, revealed it to logical term edaphobite also is sometimes be one of the deepest and most significant applied to cavernicoles. Edaphobites are caves in the eastern United States ( Schrei­ obligatory deep-soil animals that may occa­ ber and McGuffin, 1969a, 1969b). This sionally occur in caves, but if so, their oc­ cave contains more than 9.5 miles of sur­ currence is usually sporadic (Barr, 1968, veyed passage and a vertical extent of 981 p. 43). The term endogean, as defined by feet (Schreiber and McGuffin, 1969b). Its Van del ( 1964), is also used to designate most interesting geologic feature is Fantas­ this category of occasional cavernicoles. tic Pit, a huge domepit 570 feet deep. The Many of the forms found in the follow­ breakthrough in Ellisons Cave alone is good ing list are still inadequately known as to evidence of the speleological potential of distribution and ecology and their assign­ northwestern Georgia. Other large caves, ment to one of the ecological categories is such as Byers, Pettijohn, and Cemetery Pit, tentative pending further study. The follow­ also occur in this part of the state, but un­ ing abbreviations, placed in parentheses fortunately little published information on after names, have been employed: TB = these caves is available. troglobite, TP = troglophile, TX = trog­ Only a few scattered caves occur in the loxene, ED = edaphobite. Coastal Plain of southwestern Georgia and Some of the taxa found in this list are most of these are not of great extent. How­ still poorly known taxonomically and there­ ever, these caves are developed in a physio­ fore could not be determined to species. graphic region significantly different from Other forms, such as the planarians, ter­ the Appalachians and in limestones of much restrial isopods, and diplurans, currently more recent origin. The carbonate rocks of are being studied and specific names are not this area are of early Tertiary age and are yet available. Still other material represents members of the stratigraphic series which undescribed species, and to date descrip­ contains several important caves in adjacent tions and names have not been published. northern Florida (Jackson and Washington Because of these counties). reasons and the fact that a number of major caves in Georgia remain ANNOTATED LIST to be biologically investigated, this list The Schiner system of the ecological clas­ should be regarded as preliminary. How­ sification of cavernicoles, as revised by ever, considering the number of caves sam­ Racovitza and recently redefined and clari­ pled and their random distribution in the fied by Barr ( 1963b, 1968), is used in the limestone areas, the data given in this list following list. Cavernicoles usually fall into are believed to be generally representative one of four ecological categories: ( 1 ) trog- of the invertebrate cave fauna of the state.

VoLUME 33, NUMBER 1, jANUARY 1971 27 PHYLUM PLATYHELMINTHES Colbert, Lauderdale, and Madison counties, Class Turbellaria Alabama (Peck, in prep.; Cooper, in press). Helicodiscus inermis Baker (TP) ORDER TRICLADIDA Georgia Records.-Polk.: White River Cave. Family Planariidae Walker Co.: Blowing Springs Cave. Sphalloplana georgiana Hyman ( TB) Comments.-This species ranges from New Georgia Records.-Dade Co: Howards Water­ Jersey westward to Missouri and Texas, and fall Cave (Type Locality). south to the Gulf of Mexico. It has been Comments.-This species was described by recorded from caves in Van Buren and Hyman ( 1954) on the basis of material col­ Grundy counties, Tennessee ( Hubricht, lected by Mr. Charles E. Mohr. 1964a), from caves in six counties in Ala­ Sphalloplana sp. ( TB) bama (Peck, in prep.), and from one cave Georgia Records.-Dade Co.: Byers and John­ in Bath Co., Virginia (Holsinger, unpub­ son Crook caves. lished data). Comments.-This material is presently being Family Polygyridae studied by Dr. Roman Kenk of the Smith­ Mesodon perigraptus ( Pilsbry) ( TX) sonian Institution; it may well represent new Georgia Records.-Chatooga Co.: Parker Cave. records for S. georgiana, since these two caves are not far from the type locality of Mesodon (Patera) sp. ( TX) this species (see above). Georgia Records.-Walker Co.: Harrisburg Cave. PHYLUM AscHELMINTHES Comments.-This species was previously re­ corded from three counties in Georgia Class Nematomorpha (Hubricht, 1964b). ORDER GORDIOIDEA Family Zonitidae Family Gordiidae Gastrodonta intema (Say) ( TX) Gordius sp. (Accidental) Georgia Records.-Walker Co.: Blowing Springs Georgia Records.-Polk Co.: White River Cave. Cave. Comments.-A single, undetermined, adult Comments.-This species ranges from West specimen of this genus was collected from Virginia westward to Indiana and southward a shallow, mud-bottom pool in this cave. to central Alabama ( Hubricht, in litt.). Gordian worms occasionally occur in caves It has been recorded from one cave in Ala­ (Vandel, 1964; Reddell, 1965) but are bama ( Hubricht, 1964a) and from 22 probably accidentally introduced as parasitic counties in Georgia (Hubricht, 1964b). larvae by crustacean or insect hosts. Glyphyalinia paucilirata ( More let) ( TP or TX) Georgia Records.-\Valker Co.: Harrisburg PHYLUM ANNELIDA Cave. Class 0 ligochaeta Comments.-This species has been reported ORDER OPISTHOPORA from caves in Kentucky and Tennessee Family Lumbricidae ( Hubricht, 1964a). Allolobophora trapezoides Duges (ED) Glyphyalinia sculptilis (Bland) ( TX) Georgia Records.-Dade Co.: Morrison Cave. Georgia Records.-Walker Co.: Bible Springs Coments.-This species is of European ongm Cave. and was probably accidentally introduced Comments.-This species is also recorded from into this country through commerce. It has a cave in Kentucky ( Hubricht, 1964a) and also been recorded from caves in Arkansas, from eight counties in Georgia ( Hubricht, Kentucky, Tennessee, and "Vest Virginia 1964b). ( Gates, 1959). Glyphyalinia specus Hubricht ( TB) Octolasion tyrtaeus Savigny (ED) Georgia Records.-Chatooga Co.: Parker Cave. Georgia Records.-Dade Co.: Johnson Crook Dade Co.: Morrison Cave. Walker Co.: Cave No. 2. Cave Spring and Pettijohn caves. Comments.-This species was previously known PHYLUM MoLLUSCA only from caves in Kentucky, Tennessee, Class Gastropoda and Alabama. It is white and apparently ORDER STYLOMNATOPHORA blind ( Hubricht, 1965). Family Endodontidae Zonitoides arboreus ( Say) ( TP) Helicodiscus barri Hubricht ( TP) Georgia Records.-Walker Co.: Blowing Georgia Records.-Chatooga Co.: Parker Cave. Springs and Pettijohn caves. Comments.-This small, terrestrial snail is Comments.-This species is widely distributed known only from caves to date and is re­ in the United States and is known from 45 corded from Dickson and Davidson counties, counties in Georgia ( Hubricht, 1964b). It Tennessee ( Hubricht, 1962, 1964a) and has also been recorded from caves in Ala-

28 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN bama, Kentucky, Tennessee, and Texas Family Armadillidiidae (Hubricht, 1964a; Reddell, 1965; Peck, in Armadillidium vulgare (Latreille) ( TX) prep.; Cooper, in press). Georgia Records.-Walker Co.: Horseshoe Cave. PHYLUM ARTHROPOD.-\. Comments.-Eight specimens of this common Class Crustacea and widespread species were collected from ORDER EUCOPEPODA this cave. Family Ligiidae Family Canthocamptidae Ligidium elrodii chatoogaensis Schultz ( TX) Attheyella pilosa Chappuis ( TX) Georgia Records.-Chatooga Co.: Blowing Georgia Records.-Chatooga Co.: Blowing Springs Cave. Springs Cave. Comments.-This species was fairly abundant Comments.-See Bowman, et al. ( 1968) for in the stream passage in and around damp a discussion of the systematics and ecology to wet organic debris. This subspecies was of this species. A. pilosa was found in as­ recently described by Schultz ( 1970) on sociation with Cambarus in the above cave. the basis of material collected in this cave. Family Cyclopidae Other subspecies of L. elrodii are described from caves in southwestern Virginia and Cyclops ( Acanthocyclops) vernalis Fisher ( TP) eastern Tennessee by Schultz ( 1970). Georgia Records.-Bartow Co.: Kingston Salt­ Family Oniscidae petre Cave. Cylisticus convexus (De Greer) ( TX) Comments.-Four specimens of this species Georgia Records.-Dade Co.: Morrison Cave. were collected from a shallow, mud-bottom Floyd Co.: Cave Springs Cave. Walker Co.: pool. According to H. C. Yeatman (in litt.), Bible Springs and Cave Springs caves. this is an extremely variable and wide­ Comments.-This species is common through­ spread epigean species that is frequently out the United States and has been reported found in caves. from caves in Indiana, Kentucky, Tennessee, Texas, and Virginia ORDER ISOPODA (Schultz, 1970). Family Tricboniscidae Family Asellidae Caucasonethes sp. ( TB) Asellus richardsonae (Hay) ( TB) Georgia Records.-Chatooga Co. : Blowing Georgia Records.-Chatooga Co. : Blowing Springs Cave. Dade Co.: Byers Cave. Walker Springs Cave. Dade Co. : Byers and Johnson Co.: Bible Springs, Horseshoe, Mtn. Cove Crook caves. Floyd Co.: Cave Springs Cave. Farm, and Pettijohn caves. Walker Co.: Horseshoe and Mtn. Cove Farm Comments.-This material is currently being caves. studied by Professor Albert Vandel of Comments.-Steeves ( 1963) published a re­ Toulouse, France. Several species are ap­ description of this species and added several parently represented in this collection, one or new localities. A. richardsonae is wide­ more of which may be undescribed. spread and is distributed from southwestern Miktoniscus sp. ( TP) Georgia Records.-Chatooga Virginia south-southwestward to northwest­ Co. : Blowing Springs and Parker caves. Decatur Co.: ern Georgia and northern Alabama. Al­ Climax Cave. Randolph Co.: Criers Cave. though the type locality ( Nickajack Cave, Comments.-Several species of this genus oc­ Marion Co., Tennessee) of this species is cur in caves of the eastern United States situated only a few feet from the north­ and at least one of them is probably a western comer of Georgia, A. richardsonae troglobite (i.e., M. racovitzai Vandel) was previously un-recorded from Georgia. ( Vandel, 1965 ) . The most common cav­ Asellus sp. (TB) ernicolous species, however, is M. alabamen­ Georgia Records.-Walker Co.: Pettijohn Cave. sis, a troglophile which ranges from Virginia Comments.-This undescribed species is known southwestward to Alabama and Florida and from material (ca. 15 specimens) collected may well be the species represented by the from a drip pool and the cave stream. above material. Lirceus sp. ( TP or TX) ORDER AMPHIPODA Georgia Records.-Chatooga Co.: Blowing Family Gammaridae Springs Cave. Crangonyx antennatus Packard ( TB) Comments.-Nine lightly pigmented specimens Georgia Records.-Chatooga Co.: Blowing were collected from pieces of wood sub­ Springs Cave. Dade Co.: Byers, Howards merged in a small stream. Lirceus occa­ Waterfall, and Sittons caves. Floyd Co.: sionally occurs in cave streams (viz., Ten­ Cave Springs Cave. \Valker Co.: Harrisburg, nessee and Virginia) but most populations Horseshoe, Mtn. Cove Farm, and Pettijohn do not represent troglobitic species. caves.

VoLUME 33, NuMBER 1, }ANUARY 1971 29 Comments.-This species is common in caves Comments.-Both of these records are based of the southern Appalachian region, and its on single females, which, acording to H. H. range extends from the upper Tennessee Hobbs, Jr. (in litt.) possibly represent an River basin in southwestern Virginia, south­ undescribed species, closely related to C. southwestward to Floyd Co., Georgia, and striatus. In Blowing Springs Cave, this west along the Tennessee River valley to species occurs with C. latimanus. northwestern Alabama and south-central Tennessee (Holsinger, 1969). Class Arachnida Stygobromus sp. ( TB) ORDER PSEUDOSCORPIONIDA Georgia Records.-Chatooga Co.: Blowing Family Chernetidae Springs Cave. Dade Co.: Byers, Howards ·waterfall, and Rustys caves. Walker Co.: Pseudozaona sp. ( TP) Pettijohn Cave. Georgia Records.-Chatooga Co.: Parker Cave. Comments.-This undescribed species is closely Dade Co.: Johnson Crook No. 1, Morrison, related to S. mackini (a common species in and Morrison Spring caves. Walker Co.: southern Appalachian Valley caves) and Cave Spring, Hickman Gulf, and Mtn. Cove possibly ranges westward into Jackson Co., Farn1 caves. Alabama (Holsinger, 1969). Comments.-This genus is often found in caves Stygobromus sp. ( TB) of the eastern United States in association Georgia Records.-Chatooga Co.: Parker Cave. with various kinds of mammal dung. Comments.-Only five specimens of this large, Family Chthoniidae rather unusual species (undescribed) have Apochthonius sp. ( TB or TP) been taken from temporary, mud-bottom Georgia Records.-Chatooga Co.: Parker Cave. pools in the lower level of this cave. This Dade Co.: Morrison Cave. species may be distantly related to S. Kleptochthonius magnus Muchmore ( TP ? ) mackini. Georgia Stygobromus sp. ( TB) Records.-Dade Co.: Johnson Crook Cave. Walker Co.: Mtn. Cove Farm Cave. Georgia Records.-Floyd Co.: Cave Springs Comments.-This species was originally de­ Cave. Polk Co.: White River Cave. scribed from a cave in Franklin Co., Ten­ Comments.-This undescribed species is known nessee and is only slightly, if at all, modified largely on the basis of several collections for cave life (Muchmore, 1966a). It has made from White River Cave by Ernest never been collected from an epigean Ackerly during the late 1940's. This species habitat, however. has also been recorded from a cave in Cal­ houn Co., Alabama. Its morphological affini­ Family Neobisiidae ties are with S. mackini (Holsinger, 1969). Microcreagris subatlantica Chamberlin ( TX) Georgia Records.-Chatooga Co.: Parker Cave. ORDER DECAPODA Comments.-This species was originally de­ Family Astacidae scribed from caves in Colbert Co., Alabama, Cambarus cryptodytes Hobbs ( TB) but it is also found on the surface and is Georgia Records.-Decatur Co.: Climax Cave. not modified for cave life (Chamberlin, Comments.-Although this is the only known 1962). The new record (above) was re­ Georgia record for this species, it has been cently published by Muchmore ( 1969). recorded from a well and five caves in Microcreagris Florida (Warren, 1961 ) . pumila Muchmore (TP or TX) Cambarus latimanus LeConte (TP or TX) Georgia Records.-Chatooga Co.: Parker Cave. Georgia Records.-Chatooga Co.: Blowing Comments.-This species was recently de­ Springs Cave. Dade Co.: Byers and Hurri­ scribed by Muchmore ( 1969) on the basis cane caves. Walker Co.: Mtn. Cove Farm of material collected from the above cave Cave. and a cave and an epigean locality in Comments.-According to Hobbs ( 1959 ), the Blount Co., Alabama. The genus Micro­ range of this species extends from North creagris is Holarctic in distribution and Carolina to Florida. Apparently this species contains a number of epigean and cavern­ has also been taken in nothern Alabama icolous species. Troglobitic species of the (Martha Cooper, pers. comm.), but its genus have been described from Alabama, occurrence in Georgia caves was previously Tennessee, and Virginia (Chamberlin, 1962: un-recorded. At least five, rather pale speci­ Muchmore, 1966b, 1969) but none have mens of C. latimanu.s were taken from a been found in Georgia to date. single stream pool in Blowing Springs Cave. M icrocreagris sp. ( TP or TX) Cambarus sp. (TP or TX) Georgia Records.-Dade Co.: Johnson Crook Georgia Records.-Chatooga Co.: Blowing Cave. Walker Co.: Pettijohn Cave. Springs Cave. \Valker Co.: Bible Springs Comments.-A single female of this unde­ Cave. termined species was collected.

30 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN ORDER ACARINA Cicurina pallida Keyserling ( TP or TX) Family Rhagidiidae Georgia Records.-Randolph Co.: Criers Cave. Rlwgidia sp. ( TB or TP) Cicurina sp. ( TB ? ) Georgia Records.-Bartow Co.: Kingston Salt­ Georgia Records.-Dade Co.: Byers Cave. petre Cave. Dade Co.: Byers and Morrison Comments.-According to W. J, Gertsch (in caves. Walker Co.: Bible Springs and Petti­ litt.) this is an undescribed species that is john caves. more closely related to a species group from Texas than to Comments.-Rhagidiid mites are frequently a common species in nearby Alabama caves. found in caves of the eastern United States, Apparently the genus Cicurina is represented where they usually occur in damp areas as­ by troglobites only in the more southern sociated with particles of organic debris. parts of the cave areas of North America (viz., This genus is also represented in damp leaf Georgia, Alabama, and Texas). litter habitats on the surface (especially in In the central Appalachian cave region (e.g., Virginia mountainous areas). To date only two and West Virginia) this genus cavemicolous species have been described occasionally occurs in the form of a trogloxene from North America (Holsinger, 1965), but or troglophile such as C. pallida additional (undescribed) species, including (Holsinger, unpubl. data). Barr ( 1961 the material recorded above, await further ) reported C. breviaria ( TP or TX) study. from Tennessee. Tegenaria domestica Clerk ( TP or TX) ORDER OPILIONES Georgia Records. Walker Co.: Hickman Gulf Family Phalangodidae Cave. Phalangodes (Bishopella) laciniosa Crosby and Comments.-This species has also been taken Bishop (TP) in northern Alabama and southern Tennessee Georgia Records.-Chatooga Co.: Blowing caves. Springs Cave. Dade Co.: Byers and Sittons W adotes calcaratus ( Keyserling) ( Accidental?) caves. Polk Co.: White River Cave. Walker Georgia Records.-Dade Co.: Johnson Crook Co. : Bible Springs, Cave Spring, Harrisburg, No. 1 Cave. Horseshoe, and Pettijohn caves. Family Argiopidae Comments.-This species is abundant in caves Meta menardi (Latreille) ( TP) in northern Alabama and central Tennessee Georgia Records.-Dade Co.: Morrison Cave. ( Barr, 1961; Goodnight and Goodnight, Walker Co.: Harrisburg and Mtn. Cove 1942, 1960; Peck, in prep.). It has been Farm caves. collected occasionally from epigean situations Comments.-This species is common in caves in Alabama, Georgia, North and South throughout the eastern United States and is Carolina, and Tennessee; and from both usually found near entrances. inside and outside of caves in Jackson Co., Family Clubionidae Florida (Crosby and Bishop, 1924; Good­ night and Goodnight, 1942, 1953, 1960; Liocranoides unicolor Keyser ling ( TP) Peck, in prep. ) . Georgia Records.-Chatooga Co.: Parker Cave. Dade Co.: Byers Plwlangodes (Crosbyella) spintumix Crosby and and Morrison caves. Bishop (TP) Walker Co.: Bible Springs, Hickman Gulf, Horseshoe, and Mtn. Cove Fann caves. Georgia Records.-Decatur Co.: Climax Cave. Comments.-This species is also common in Comments.-This species is known from caves caves of south-central Tennessee and north­ in Alabama, Arkansas, and Florida and em Alabama (Barr, 1961; Gertsch, in litt.). from epigean localities in Alabama, Arkan­ sas, Florida, and Mississippi (Goodnight Family Leptonetidae and Goodnight, 1942; Peck, in prep.). Leptoneta sp. ( TB or TP) Georgia Records.-Dade Co.: Byers Cave. ORDER ARANEAE Walker Co.: Harrisburg and Pettijohn caves. Family Agelenidae Comments.-According to W. J. Gertsch (in Calymmaria cavicola (Banks) ( TP or TX) litt. ) these collections represent a new Georgia Records.-Chatooga Co.: Parker Cave. species. The genus Leptoneta is also repre­ Dade Co.: Byers Cave. Walker Co.: Bible sented by a number of troglobitic species Springs and Mtn. Cove Farm caves. in the Edwards Plateau region of central Comments.-This species also inhabits caves Texas (Gertsch, pers. comm.; Reddell, in nearby Tennessee (Barr, 1961) and is 1965.) This group also occurs in caves known from 22 caves in Alabama (Peck, around the Mediterranean ( Vandel, 1964 ) . in prep.). Family Linyphiidae C icurina a rcuata Keyserling ( TP or Accidental ? ) Centromerus denticulatus (Emerton) ( TP) Georgia Records.-Walker Co.: Cave Spring Georgia Records.-Walker Co.: Horseshoe Cave. Cave.

YoLuME 33, NuMBER 1, JANUARY 1971 31 Comments.-This species is also found oc­ West Virginia (Gertsch, in I itt.). Another casionally in caves of southeastern Tennessee species of this genus (A. porteri) is a and northern Alabama. common troglophile in Texas caves (Reddell, Phanetta subterranea (Emerton) ( TB ? ) 1965) and is also reported from Tennessee Georgia Records.-Dade Co.: Byers, Howards caves (Barr, 1961). Waterfall, Johnson Crook No. 1, Morrison, and Sittons caves. Floyd Co.: Cave Springs Class Symphyla Cave. Walker Co.: Cave Spring and Harris­ Scutigerella sp. (ED) burg caves. Georgia Records.-Dade Co. : Johnson Crook Comments.-This species is common in caves Cave No. 2. Walker Co.: Harrisburg Cave. all over the eastern United States (i.e., Comments.-Although symphylans are depig­ Appalachian region and Interior Low mented and anophthalmic, they are mem­ Plateaus). bers of the soil fauna ( endogean or edaphic) Porrhomma cavernicolum (Keyserling) ( TB ? ) and are not true cave inhabitants ( Vandel, Georgia Records.-Bartow Co.: Kingston Salt­ 1964). petre Cave. Comments.-This species is also widespread Class Chilopoda in the Appalachian and Interior Low Plateau ORDER GEOPHILOMORPHA cave regions but is apparently not as com­ Family Geophilidae mon as P. subterranea. Arenophilus bipuncticeps (Wood) ( TX or Family Lycosidae Accidental) Pi rata sp. (Accidental) Georgia Records.-Chatooga Co.: Blowing Georgia Records.-\Valker Co.: Horseshoe Springs Cave. Cave. ORDER LITHOBIOMORPHA Family Nesticidae Family Lithobiidae Nesticus pallidus Emerton (TP or TX) Lithobius atkinsoni Bollman Georgia Records.-Floyd Co.: Cave Springs ( TP) Cave. Randolph Co.: Criers Cave. Georgia Records.- Randolph Co.: Criers Cave. Comments.-This species is a common trog­ Comments.-This species is found in caves loxene in caves of the United States and on the southern Coastal Plain and in epigean even Mexico. habitats farther north (Peck, 1970). Family Pholcidae N eolithobius voracior (Chamberlin) ( TP) Plwlcus sp. ( TP or TX) Georgia Records.-Decatur Co.: Climax Cave. Georgia Records.-Floyd Co.: Cave Springs Comments.-This species is also reported from Cave. three localities in Mississippi (Chamberlin, Comments.-The family Pholcidae is also rep­ 1925). resented in Texas and Mexico by caverni­ Paitobius sp. ( TX or Accidental) colous species (Nicholas, 1962; Reddell, Georgia Records.-Dade Co.: Morrison Spring 1965 ) . In addition, Barr ( 1961 ) reported Cave. the group from Tennessee caves. Comments.-This species is probably P. Family Salticidae adelus Chamberlin, which is also known M aevia vittata Koch ( Accidental) from one cave in Alabama (Peck, in prep. ). Georgia Records.-Walker Co.: Hickman Gulf Pampibius sp. ( TX or Accidental) Cave. Georgia Records.-Walker Co.: Cave Spring Family Symphytognathidac Cave. M aymena ambita (Barrows) ( TP) Comments.-This material apparently repre­ Georgia Records.-Walker Co.: Horseshoe sents a new species in a previously mono­ Cave. typic genus. Comments.-This species is known primarily from caves in Alabama, Kentucky, and ORDER SCOLOPENDROMORPHA Tennessee (Gertsch, 1960). Family Cryptopidae Family Theridiidae Scolopocryptops sexspinosa (Say) (TX or Achaearanea tepidariorum (Koch) ( TP) Accidental) Georgia Records.-Gordon Co.: Roberts Cave. Georgia Records.-Dade Co.: Johnson Crook Polk Co.: White River Cave. \Valker Co.: and Morrison caves. \Valker Co. : Pettijohn Bible Springs Cave. Cave. Comments.-This species is also recorded from Comments.-This species is common in the a few caves in nearby Alabama and Ten­ eastern United States and is known from nessee, and as far north as Monroe Co., one cave in Alabama.

32 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN Class Diplopoda S. a. austrinus and S. a. nudus Chamberlin ( 1946). ORDER CHORDEUMIDA The latter is known only from its type locality (Kingston Saltpetre Cave) but Family Cleidognoidae should now be re-examined in light of the Pseudotremia eburnea Loomis ( TB) new material as it may well represent a dis­ Georgia Records.-Dade Co.: Byers and John­ tinct species. S. a. austrinus was originally son Crook No. 1 caves. Walker Co.: Hick­ described from Manitou Cave, DeKalb Co., man Gulf, Mtn. Cove Farm, and Pettijohn Alabama by Loomis ( 1943 ), but recent caves. collecting shows it (or a very closely related Comments.-Loomis ( 1939), in his description form) to be rather common in northwestern of this species, designated Cricket Cave, Georgia. Scoterpes copei (Packard) has also Dade County as the type locality. As been reported from Georgia (i.e., Sawmill pointed out earlier, however, this cavf' Cave, Dade Co.) and Tennessee (Loomis, could not he identified during the field work. 1939, 1943; Dearolf, 1953; Chamberlin and Loomis (1939), and also Dearolf ( 1953), Hoffman, 1958), but the listed Pseudotremia sp. ( immatures) from range of this species should Creek Bed Cave, Dade Co., but the latter be restricted to south-central Ken­ eave could not be identified either. tucky (Causey, in litt.). It should be noted Pseudotremia sp. ( TB or TP) that Sawmill Cave, like Creek Bed and Georgia Records.-Dade Co.: Case, Howards Cricket caves, could not be identified dur­ Waterfall, Morrison, Morrison Spring, Run­ ing the field work and is probably known ning \Vater, and Sittons caves. \Valker Co.: to the Georgia Cave Survey by another Bible Springs and Harrisburg caves. name. Comments.-Most of the above material is Scoterpes sp. ( TB) known from immatures and females and Georgia Records.-Chatooga Co.: Parker Cave. could not be specifically determined. Dade Co.: Sittons Cave. Walker Co.: Harris­ Family Lysiopetalidae burg Cave. Abacion magnum ( Loomis) ( TX) Comments.-The specimens from Sittons and Georgia Records.-Dade Co.: Byers Cave. Harrisburg caves were immature or females Chatooga Co.: Blowing Springs Cave. Polk and could not be determined. The material Co.: \Vhite River Cave. from Parker Cave represents an undescribed Comments.-This species was originally de­ species (Causey, in litt.) . scribed from Madison Co., Alabama ( epi­ gean habitat) hy Loomis ( 1943). Chamber­ ORDER JULIDA lin ( 1946) reported A. lactarium from Family Nemasomatidae Kingston Saltpetre Cave, Bartow Co., Ameractis satis Causey ( TB or TP) Georgia, but this was probably an error Georgia Records.-Dade Co.: Morrison Cave. for A . magnum (Causey, in litt.). The Comments.-This species was first described former is known from the Coastal Plain from a cave in White Co., Tennessee by and Piedmont of the eastern United States Causey (1959). More recently, Barr (1961) (Chamberlin and Hoffman, 1958) but not reported the range as extending from Over­ from the Appalachian Valley or Plateau. ton County to Hamilton County, Tennessee, Reddell ( 1965) reported another species of and Peck (in prep. ) collected it from two this genus (A. texense) from several caves caves in Alabama. The Georgia and Ala­ in central Texas. bama records extend the range of this species Family Striariidae to the southeast. Striaria sp. ( TX) Blaniulus guttulus ( Bose) (ED) Georgia Records.-Chatooga Co.: Parker Cave. Georgia Records.-Dade Co.: Morrison Cave. Comments.-This genus is represented by two Comments.-This eyeless soil inhabitant is the troglobitic species in the United States only known species of the genus in North (Shear, 1969). America and is apparently an introduced Family Trichopetalidae form from Europe. Scoterpes austrinus Loomis ( TB) ORDER PLA TYDESMIDA Georgia Records.-Bartow Co.: Kingston Salt­ petre Cave. Chatooga Co.: Blowing Springs Family Andrognathidae Cave. Dade Co.: Johnson Crook No. 1 and Andrognathus corticarius Cope ( TX) Morrison caves. Polk Co.: De a tons and Georgia Records.-Floyd Co.: Cave Springs White River caves. Walker Co.: Bible Cave. Springs, Horseshoe, Mtn. Cove Farm, and Comments.-This epigean species has been re­ Pettijohn caves. ported from several states in the southeast­ Comments.-At least two subspecies (species ern United States (Chamberlin and Hoffman, ?) of S. austrinus occur in Georgia caves- 1958).

VoLUME 33, ~UMBER 1, }ANUARY 1971 33 ORDER POLYDESMIDA Class Insecta Family Paradoxosomatidae ORDER COLLEMBOLA Oxidus gracilis (Koch) ( TP) Family Entomobryidae Georgia Records.-Decatur Co.: Climax Cave. Pseudosinella hirsuta ( Delamare-Debouttevillc) Floyd Co.: Cave Springs Cave. (TB) Comments.-The range of this species is Georgia Records.-Bartow Co.: Kingston Salt­ tropicopolitan. It has apparently been in­ petre Cave. Chatooga Co.: Blowing Springs troduced into the United States, where it Cave. Dade Co.: Johnson Crook, Howards has become well established in the south Waterfall, Morrison, Running Water, and and west (Chamberlin and Hoffman, 1958). Rustys (?) caves. Polk Co.: Deatons Cave. Although 0. gracilis is commonly found in Walker Co.: Bible Springs, Harrisburg, greenhouses, it is reported from several Horseshoe, Mtn. Cove Farm, and Pettijohn caves in Texas (Reddell, 1965) and one caves. cave in Pennsylvania (Loomis, 1939). This Comments.-This common troglobite is also species has also been collected from caves known from caves in Alabama, Kentucky, in Mexico and one cave in Alabama (Peck, Tennessee, and the extreme tip of south­ in prep. and unpubl. data). western Virginia (Christiansen and Culver, 1968). Family Polydesmidae Polydesmus sp. (or Pseudopolydesmus sp.) Family Sminthuridae (TX) Arrhopalites sp. ( TP) Georgia Records.-Dade Co.: Creek Bed Cave Georgia Records.-Bartow Co.: Kingston Salt­ petre Cave. Walker Co.: Mtn. Cove Farm (of Loomis and De a rolf). Cave. Comments.-A single male from this cave Comments.-This genus is a widespread, was said by Loomis ( 1939) to be close to, Holarctic group with a number of well if not identical with, Polydesmus pinetorum. known troglophilic species ( Christiansen, ( =Pseudopolydesmus pinetorum--Chamber­ 1966). lin and Hoffman, 1958). Family Tomoceridae ORDER SPIROSTREPTIDA Tomocerus bidentatus Folsom ( TP) Georgia Records.-Chatooga Co.: Parker Can'. Family Cambalidae Dade Co.: Byers and Morrison caves. Cambala annulata Say (TP) Walker Co.: Bible Springs Cave. Georgia Records.-Randolph Co.: Criers Cave. Comments.-This widespread species is known Comments.-This species is also known from from numerous epigean and subterranean caves in Jackson Co., Florida and central localities in the eastern United States. It is Alabama and from many epigean localities especially common in caves in the Ap­ in the Appalachians (Hoffman, 1958; Shear, palachian region (Christiansen, 1964 ) . 1969). Tomocerus dubius Christiansen ( TP) Cambala minor Bollman (TP) Georgia Records.-Bartow Co.: Kingston Salt­ Georgia Reocrds.-Chatooga Co.: Parker Cave. petre Cave. Dade Co.: Morrison Cave. Walker Co.: Comments.-This species has been recorded Horseshoe and Pettijohn caves. from a number of caves and surface sites Comments.-This species (or species complex in the southeastern United States; it has -see Shear, 1969) is a common inhabitant also been reported from several epigean lo­ calities in California ( Christiansen, 1964 ) . of forested areas in the eastern United States. It frequently occurs in caves, how­ Tomocerus flavescens ( Tullberg) ( TP) ever, and has been reported from such Georgia Records.-Dade Co.: Johnson Crook Cave. Walker Co.: Cave Spring and Horse­ habitats in Alabama (Peck, in prep.), Ken­ shoe caves. tucky (Shear, 1969 ), Tennessee (Barr, Comments.-This species occurs across the 1961), and Virginia-West Virginia (Hol­ entire North American Continent and has singer, unpubl. data). been reported from many caves (Christian­ Cambala sp. ( TP) sen, 1964). Georgia Records.-Chatooga Co.: Blowing Springs and Parker caves. Polk Co.: Deatons ORDER THYSANURA and White River caves. Walker Co.: Family Nicoletiidae Cave Spring Cave. Nicoletia sp. ( TB or TP) Comments.-Although these records are based Georgia Records.-Walker Co.: Horseshoe on immature specimens, they probably rep­ Cave. resent Cambala minor or a closely related Comments.-This undescribed species may be form. the same as that collected from two caves

34 THE NATIONAL SPELEOLOGICAL SociETY BuLLETI~ in northeastern Alabama (Peck, in prep., species also occurs in a variety of forested Cooper, in press). Furthermore, this form habitats. may be an inhabitant of the deep soil H adenoecus puteanus Scudder ( TX) ( edaphic) rather than a true cave dweller. Georgia Records.-Dade Co.: Byers, Howards Reddell ( 1966) reported a rather common Waterfall, Johnson Crook, and Morrison troglobitic species of Nicoletia from caves Spring caves (plus Creek Bed, Cricket, and in central Texas. Saw Mill caves of Dearolf, 1953). Gordon ORDER DIPLURA Co.: Roberts Cave. Polk Co.: White River Family Japygidae Cave. Walker Co.: Bible Springs and Mtn. Cove Farm caves. Unidentified genus and species (ED) Comments.-The range of this species etxends Georgia Records.-Chatooga Co.: Blowing from northeastern Ohio and southern New Springs Cave. Walker Co.: Mtn. Cove Farm York southward along the Cumberland Pla­ Cave. teau, Appalachian Valley, and Piedmont to Comments.-Single specimens of this unidenti­ northeastern Alabama and northern Georgia. fied form were collected from damp spots It has been recorded from Charlton County under rocks (with Plusiocampa, see below) in southeastern Georgia and perhaps in in the above caves. Virtually nothing is pre­ error from Lawrence Co., Mississippi. This sently known about the distribution of this species lives in cave entrances, along cliffs rare group in eastern North American caves. and talus slopes, and in mesic forests (Hub­ Family Campodeidae bell, in litt. ) . Plusiocampa sp. (TB?) Georgia Records.-Chatooga Co.: Blowing ORDER PSOCOPTERA Springs Cave. Dade Co.: Johnson Crook No. Family Psocidae 2 and Morrison caves. Walker Co.: Bible Psillipsocus ramburii Selys-Longchamps ( TP) Springs, Cave Spring, Mtn. Cove Farm, and Georgia Records.-Bartow Co.: Yarborough Pettijohn caves. Cave. Walker Co.: Cave Spring and Harris­ Comments.-Only five cavernicolous species of burg caves. this genus have been described from North Comments.-This species is known from America ( Conde, 1949), although a wealth Europe and North America, and in the of un-studied material from southern Ap­ latter its range extends from Michigan to palachian caves indicates a number of un­ Mexico. P. ramburii also occurs in caves in described forms. Two of the five described Alabama, Tennessee, and Texas and is cave species occur in nearby Alabama (i.e., usually found on organic matter in the Madison County), but it is presently impossi­ drier parts of these caves (Gurney, 1943; ble to say whether the Georgia records repre­ Mockford, 1950; Barr, 1961; Reddell, 1966; sent new species or only mark range exten­ Peck, in prep., Cooper, in press). sions of the Alabama forms. ORDER LEPIDOPTERA ORDER ORTHOPTERA Family Tineidae ( ? ) Family Gryllacrididae Unidentified genus and species (TX) Ceuthophilus ensifer ensifer Packard ( TX) Georgia Records.-Dade Co.: Byers Cave. Georgia Records.-Dade Co.: Byers, Howards Walker Co.: Harrisburg, Mtn. Cove Farm, Waterfall, Johnson Crook, Morrison, and and Pettijohn caves. Morrison Spring caves. Comments.-Several moths were collected Comments.-This subspecies is also known from the droppings of the cave rat from localities in eastern Jackson Co., Ala­ bama, and southern Hamilton and Marion (Neotoma) in the above caves. counties, Tennessee (Hubbell, 1936 and ORDER HYMENOPTERA in litt. ) . In addition to caves, this species Family Braconidae inhabits forested talus slopes and cliffs. Aspilota sp. ( TX) Ceuthophilus gracilipes (Haldeman) ( TX) Georgia Records.-Bartow Co.: Yarborough Georgia Records.-Bartow Co.: Yarborough Cave. Dade Co.: Byers, Morrison, and Cave. Dade Co.: Johnson Crook Cave. Morrison Spring caves. Gordon Co.: Roberts Walker Co.: Horseshoe and Mtn. Cove Cave. Walker Co.: Bible Springs Cave. Farm caves. Comments.-The distribution of this common Comments.-These small wasps are commonly species runs in a wide band from Massa­ found on bait left in caves in northern chusetts and New York southward along the Alabama as well as the caves mentioned Appalachians to Alabama and northwestern above. They are probably parasitic upon Florida, and then northwestward to southern the fly larvae thaJ are intially attracted to Missouri (Hubbell, 1936 and in litt.). This the bait.

VoLUME 33, NuMBER 1, JANUARY 1971 35 ORDER DIPTERA In the above caves these flies were taken Family Heleomyzidae from carrion traps and human feces. Aecothea specus (Aldrich) ( TP or TX) Family Sciaridae Georgia Records.-Dade Co.: Johnson Crook Sciara sp. ( TP or TX) Cave. Polk Co.: White River Cave. Walker Georgia Records.-Bartow Co.: Kingston Salt­ Co.: Bible Springs, Cave Spring, Horseshoe, petre Cave. Dade Co.: Byers, and Johnson and Mtn. Cove Farm caves. Crook caves. Polk Co.: White River Cave. Comments.-This species is distributed from Walker Co.: Mtn. Cove Farm Cave. Alaska to California and eastward to On­ Comments.-These small, dark, delicate flies tario (Canada ) , Ohio, and Georgia, and is are found in moist places where their larvae common in caves (Gill, 1962). Dearolf feed on fungus (Curran, 1965; Stone et al., ( 1953) reported Aecothea fenestralis Fallan 1965). Several species may be represented from Saw Mill Cave but this may be a by the above collections, most of which were misidentification for A. specus. collected within a few hundred feet of the Amoebaleria defessa ( Osten-Sacken) ( TP or TX) cave entrances in traps baited with carrion. Georgia Records.-Dade Co.: Morrison, Run­ Family Spaeroceridae ning Water, and Rustys caves (plus Saw Leptocera sp. ( TP or TX) Mill Cave of Dearolf, 1953). Walker Co.: Georgia Record-Chatooga Co.: Blowing Harrisburg, Mtn. Cove Farm, and Pettijohn Springs Cave. Dade Co.: Johnson Crook caves. Cave. Walker Co.: Bible Springs, Mtn. Comments.-This species was previously known Cove Farm, and Pettijohn caves. to range from Michigan eastward to Penn­ Comments.-These small, dark flies are com­ sylvania and south-southwestward to Mis­ mon in caves, where they are frequent souri and Alabama. The new Georgia records scavengers on decaying organic matter mark a range extension. A. defessa is com­ (Curran, 1965; Stone et al., 1965). mon in caves of the eastern United States and has been collected as deep in caves as Family Tipulidae 150 yards from the entrance (Gill, 1962, Unidentified genus and species ( TX ? ) 1965). A population with a morphology Georgia Records.-Dade Co.: Johnson Crook similar to the one formerly assigned to Cave. Walker Co.: Bible Springs Cave. Amoebaleria sackeni Garrett was found in Comments.-The above collections were made Mtn. Cove Farm Cave, but this species was in the twilight zone. Tipulids are often recently synonymized with A. defessa ( Stey­ found in small swarms in cave entrances, skal, 1968 ) . perhaps using the moist entrances as day­ Heleomyza brachypterna ( Leow) ( TP or TX) time retreats. Georgia Records.-Walker Co.: Harrisburg Family Phoridae and Mtn. Cove Farm caves. Megaselia sp. ( TP or TX) Comments.-The above records are based on Georgia Records.-Bartow Co.: Yarborough females and should be confirmed with males Cave. Chatooga Co.: Blowing if and when available. This species has been Springs and collected from caves but was previously un­ Parker caves. Dade Co.: Byers, Johnson recorded from Georgia. (Gill, 1962). Crook, and Morrison caves. Polk Co.: White River Cave. Walker Co.: Cave Spring, Family Muscidae Harrisburg, Horseshoe, Mtn. Cove Farm, Chaetogenia sp. (Accidental) and Pettijohn caves. Georgia Records.-Bartow Co.: Yarborough Cave. Comments.-These small, dark flies are com­ Comments.-This collection was made from mon around decomposing organic matter in human feces. caves. Several species are represented in the Family Mycetophilidae above collections which were made mostly from Neotoma dung and carrion Unidentified genus and species (TP or TX) bait. Georgia Records.-Walker Co.: Bible Springs ORDER COLEOPTERA and Mtn. Cove Farm caves. Family Cantharidae Comments.-These collections were made from Unidentified genus and species ( TX) carrion traps. Georgia Records.-Chatooga Co.: Parker Cave. Family Psychodidae Walker Co.: Harrisburg, Mtn. Cove Farm, Psychoda sp. ( TP or TX) and Pettijohn cave-5. Georgia Records.-Bartow Co.: Kingston Salt­ Comments.-The above collections were all petre Cave. Walker Co.: Pettijohn Cave. larval forms and were taken from damp Comments.-These flies are frequently found areas just inside the dark zone. These beetle in moist, shady places with decaying organic larvae have also been encountered in caves debris (Curran, 1965; Stone et a!., 1965). in Alabama and Tennessee.

36 THE NATIOXAL SPELEOLOGICAL SociETY BuLLETIK Family Carabidae and Virginia (Hatch, 1933). Most of the Agonum (Rhadine) caudatum (LeConte) ( TP) southern records are from caves. Georgia Records-Dade Co.: Rustys Cave. N emadus sp. ( TP or TX) Comments.-This species is a common trog­ Georgia Records.-Dade Co.: Morrison Cave. lophile in the caves of Alabama, Tennessee, 'Walker Co.: Cave Spring, Horseshoe, and and Yirginia (Barr, 1964). It was previ­ Mtn. Cove Farm caves. ously unrecorded from Georgia caves. Comments.-All of the above collections were Anillinus sp. ( TX ? ) made from liver bait in the dark zone. Georgia Records.-Dade Co.: Morrison Cave. Members of this genus are only occasionally Comments.-This undescribed, forest-soil in­ encountered in caves, where they may exist habiting species has also been found in in large populations on bat guano. Alabama caves (Barr, 1969; Peck, in prep.). Prionochaeta opaca Say ( TP or TX) Atranus pubescens (Dejean) ( TP) Georgia Records.-Chatooga Co.: Blowing Georgia Records.-Walker Co.: Bible Springs Springs Cave. Walker Co.: Bible Springs, ca,·e. Cave Spring, Horseshoe, and Mtn. Cove Comments.-This species occurs in caves of Farm caves. the eastern United States, the Ozarks, and Comments.-All of the above collections were Texas (Barr, 1964). made from liver bait in the dark zone. This Harpalus 1 Pseudophonus) sp. (Accidental) species was previously recorded from Que­ Georgia Records.-Dade Co.: Morrison Cave. bec (Canada) and New Hampshire south Walker Co.: Bible Springs Cave. to North Carolina and west to Arkansas Pseudanophtlwlmus digitus Valentine ( TB) (Hatch, 1933). Georgia Records.-Dade Co.: Byers and Ptomaphagus whiteselli Barr ( TB) Johnson Crook caves. Georgia Records.-Dade Co.: Byers, Case, Comments.-This species was previously Morrison, and Sittons caves. known only from two females taken in Comments.-This species was originally de­ Tennessee Caverns, Hamilton Co., Ten­ scribed from Sittons Cave by Barr ( 1963a) nessee ( Barr, 1965 ) . and was also reported from Sequoyah Pseudmwpllthalmus fulleri Valentine ( TB) Caverns, DeKalb Co., Alabama by Peck Georgia Records.-Dade Co.: Byers, Howards (1966). \Vaterfall, Johnson Crook, Morrison, and Ptomaphagus sp. ( TB) Sittons caves. Walker Co.: Horseshoe Cave. Georgia Records.-Walker Co.: Bible Springs, Comments.-This species was previously Mtn. Cove Farm, and Pettijohn caves. known only from its type locality (Ten­ Comments-This undescribed species appears nessee Caverns, Hamilton Co., Tenn.) and to be more closely related to a species from Howards Waterfall Cave (see above) (Barr, Jackson Co., Alabama than to P. whiteselli 1965). The new records above not only from nearby caves in Dade County. extend the range of this species but also Ptomaphagus sp. ( TP) indicate that it is not as rare as previously Georgia Records.-Dade Co.: Johnson Crook thought (see Barr, 1965, p. 66). No. 2 and Morrison caves. Walker Co.: Pseudanophthalmus sp. ( TB) Bible Springs Cave. Georgia Records.-Chatooga Co.: Blowing Comments-This undescribed species has also Springs Cave. Walker Co.: Mtn. Cove Farm been collected from forest litter in Georgia and Pettijohn Caves. and Alabama and from caves in Alabama Comments.-These collections represent an (Peck, in prep.). undescribed species. Sciodrepoides fumatus terminans LeConte ( TX?) Tachys (Tachyura) ferrugineus (Dejean) (TP) Georgia Records.-Waker Co.: Cave Spring Georgia Records.-Decatur Co.: Climax Cave. Cave. Comments.-This species occurs rarely in Comments.-Hatch ( 1933) called this species caves in the eastern United States but is Catops (Sciodrepoides) terminans and gave known from four caves in Alabama (Peck, its range as extending from British Co­ in prep.). According to Barr ( 1964), this lumbia and New Bmnswick south to species is common in caves in Texas. Illinois and North Carolina. This species Family Leiodidae (subfamily Catopinae) was collected from liver bait in the above Catops gratiosus Blanchard (TP or TX) cave where it was probaby accidental rather Georgia Records.-Chatooga Co.: Parker Cave. than trogloxenic. Dade Co.: Morrison Cave. Walker Co.: Mtn. Family Pselaphidae Cove Farm Cave. Batrisodes globosus (LeConte) ( TP) Comments.-All of the above collections were Georgia Records.-Decatur Co.: Climax Cave. made from rotting liver bait in the dark Comments.-This is the best known and most zone. This species was previousy recorded widely distributed species in the genus and from \Vashington (state), Ontario (Canada), occurs throughout eastern North America and Xew England southward to Kentucky (Park, 1947). This species ha.s alsa been

VoLU::\IE :3:3, NuMBEH. 1. JANUARY 1971 37 taken from Sanders Cave, Conecuh County, Quedius erythrogaster Mannerheim ( TP) in southern Alabama (Peck, in prep.). Georgia Records.-Dade Co.: Morrison Cave. Batriasymmodes spelaeus Park ( TP) Walker Co.: Harrisburg, Hickman Gulf, and Georgia Records.-Chatooga Co.: Blowing Pettijohn caves. Springs Cave. Walker Co.: Bible Springs Comments.-This species ranges across the Cave. northern United States and into Canada and Comments.-This species was previously re­ is common in caves of the eastern part of ported from caves in nine Alabama coun­ the country. ties and four Tennessee counties; more re­ Tachinus fimbriatus Gravenhorst (Accidental) cently it was collected from a sandstone Georgia Records.-Walker Co.: Pettijohn Cave. rockshelter in Alabama (Park, 1951, 1958, Comments.-This single record is from a 1960, 1965; Peck, in prep.; Steeves, in male specimen collected from a carrion trap litt.). just inside the cave entrance. Macherites (Subterrochus) sp. ( TB) Georgia Records.-Walker Co.: Mtn. Cove ZooGEOGRAPHIC RELATIONSHIPS Farm Cave. Of the approximately 130 invertebrate Comments.-This undescribed species is ap­ species presently recorded from Georgia parently a troglobite (Park, 1960; Steeves, in litt.). caves, between 24 and 27 are troglobites. Family Staphylinidae The troglobites represent approximately 20 Atheta sp. ( TP) percent of the total invertebrate fauna and Georgia Records.-Bartow Co.: Yarborough about 45 percent of them are apparently Cave. Chatooga Co.: Blowing Springs and endemic to northwestern Georgia. Owing to Parker caves. Dade Co.: Byers, Johnson the fact that taxonomic studies are still in Crook, and Morrison caves. Walker Co.: progress and that further field work will Bible Springs, Cave Spring, Horseshoe, Mtn. Cove Farm, and Pettijohn caves. undoubtedly yield additional species, these Comments.-The above collections, mostly figures must be regarded as provisional. The from carrion traps and liver bait, probably troglobitic species are taxonomically dis­ represent at least three species of this genus. tributed as follows, with the number of Geodromicus brunneus Say ( TX ? ) Georgia endemics given in parentheses. Plan­ Georgia Records.-Walker Co.: Bible Springs arians, 1 ( 1 ) ; gastropods, 1; asellid isopods, and Mtn. Cove Farm caves. Comments.-This species might be an acci­ 2 ( 1 ) ; trichoniscid isopods, 1 ( 1?) ; gam­ dental rather than a trogloxene. marid amphipods, 4 (1); crayfish, 1; chthon­ Lesteva pallipes LeConte ( TP) iid pseudoscorpions, 1 or 2 ( 1 ) ; spiders, 3 Georgia Records.-Chatooga Co.: Blowing or 4 ( 2); chordeumid millipedes, 2 or 3 ( 2 Springs Cave. Walker Co.: Bible Springs and Mtn. Cove Farm caves. or 3); diplurans, 1? ( 1?); collembolans, 1; Comments.-All of the above collections were carabid beetles, 3 ( 1 ) ; leiodid beetles, 2 made from carrion traps or liver bait. ( 1 ) ; and pselaphid beetles, 1 ( 1 ) . Oxypoda sp. ( TX ? ) More than one-half of the troglobitic species Georgia Records.-Bartow Co.: Yarborough Cave. of northwestern Georgia are also found in Philonthus cyanipennis Fabricius (Accidental) caves of nearby states, notably in neighbor­ Georgia Records. -Walker Co.: Pettijohn ing Alabama and Tennessee. Cambarus cryp­ Cave todytes, a troglobitic crayfish recorded from Comments-This species was collected from liver bait. Climax Cave in southern Georgia, is also Philonthus sp. ( TP or TX) known from northern Florida. Georgia Records.-Walker Co.: Bible Springs Although taxonomic and distributional Cave. Comments.-Tbis species is presently unde­ data are still incomplete, it is nevertheless scribed. possible to analyze the distribution of the Quedius fulgidus Fabricius ( TP) troglobitic species of the Georgia Appalach­ Georgia Records.-Polk Co.: White River ians. In northwestern Georgia two zoogeo­ Cave. graphic faunal units, the Appalachian Pla­ Comments.-This species is distributed across teau unit and the Appalachian Valley unit, the United States and is known from caves in Illinois and Kentucky (Peck, unpubl. can be designated. These units correspond data). to the physiographic units of the same name

38 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN and are geologically defined on the basis the milliped Pseudotremia eburnea, and an of theoretical dispersal barriers. undetermined (undescribed?) species of the The Plateau unit lies on the eastern mar­ dipluran Plusiocampa. gin of the Appalachian (Cumberland) Pla­ ( 3) There are a few species which occur teau which, in tum, continues northwest­ in caves in one or the other Plateau sub­ ward into south-central Tennessee and west­ units but not in both: the Lookout Valley ward into northern Alabama. With some fauna includes the planarian Sphalloplana exceptions it may correspond zoogeograph­ georgiana, an undescribed species of the ically to the Cumberland Plateau faunal unit spider Cicurina, the millipede Ameractis satis, designated by Barr ( 1967). However, the the carabid beetle Pseudanophthalmus dig­ Plateau of Georgia is exceptional because: itus, and the leiodid beetle Ptomaphagus (a) its geological and physiographic rela­ whiteselli; the eastern Lookout Mountain tionship to the Cumberland Plateau is mar­ fauna includes an undescribed species (one ginal, (b) its limestones lie in close prox­ each) of Asellus, Pseudanophthalmus, Ptom­ imity to those of the Appalachian Valley, aphagus, and Macherites. ( 4) Two species and ( c) its limestones are extrinsically iso­ occur in the Plateau (Lookout Valley) and lated from those which form the broad, cav­ also in the Appalachian Valley and include erniferous western edge of the Cumberland an undetermined (or undescribed?) species Plateau in Alabama and Tennessee, the area of the pseudoscorpion Apochthonius, and the more precisely designated as the Cumber­ carabid beetle Pseudanophthalmus fulleri. land Plateau faunal unit by Barr ( 1967). ( 5) Only two species ( both undescribed ) The Plateau faunal unit can be subdivided of the amphipod Stygobromus are known into ( 1 ) the Lookout Valley subunit, situ­ exclusively from the Appalachian Valley ated entirely within Dade County and de­ caves-one from Parker Cave and one from fined as the cavernous area lying between caves in Floyd and Polk counties. Sand Mountain on the west and Lookout Although not formally covered in this Mountain on the east; and ( 2) the eastern paper, the troglobitic vertebrates of Georgia Lookout Mountain subunit (including Pig­ might also be mentioned in passing, since eon Mountain) which lies mostly within their presence in the state further empha­ Walker County. sizes the zoogeographic significance of this Five distinct patterns of troglobite distri­ area. Three species of vertebrate troglobites bution are distinguishable. ( 1 ) There are a have been reported from Georgia caves­ few species which are not only common in two salamanders and one fish. Haideotriton caves in both faunal units of northwestern wallacei (Carr), a rare troglobitic sala­ Georgia but are also found in much of the mander, was first described from a deep southern Appalachian cave region. These well at Albany, Dougherty Co., Georgia, and species show comparatively little range re­ has since been found in a number of caves striction and include the snail, Glyphyalinia in Jackson Co., Florida and in Climax Cave, specus, the isopod Asellus richardsonae, the Decatur, Co., Georgia ( Pylka and Warren, amphipod Crangonyx antennatus, the spiders 1958; Warren, 1961 ). A more widely dis­ Phanetta subterranea and Porrhomma cavern­ tributed troglobitic salamander, Gyrinophilus icolum, millipedes of the Scoterpes austrinus palleucus McCrady, previously recorded complex, and the collembolan Psendosinella from caves in southern and eastern Tennes­ hirsuta. ( 2) There are a number of forms see and Jackson Co., Alabama (Brandon, which occur in both subunits of the Plateau 1966), was recently reported from Harris­ but not in the Valley and include an undeter­ burg Cave in Walker County, Georgia by mined (undescribed?) species of the terres­ Cooper ( 1968 ) . This is the first and only trial isopod Caucasonethes, an undescribed known record to date for this species in species of the amphipod Stygobromus, the Georgia. The southern cavefish, Typhlich­ pseudoscorpion Kleptochthonius magus, an thys subterraneus Girard, a rather widely undescribed species of the spider Leptoneta, distributed troglobite whose range extends

VoLUME 33, NUMBER 1, }ANUARY 1971 39 from Missouri to southern Indiana and south ACKNOWLEDGMENTS through Kentucky and Tennessee to northern This project was partially supported by Alabama, was recently discovered in Twin an Old Dominion University Education Snakes Cave in Dade Co., Georgia, by A. Foundation Grant to J. R. Holsinger and by Iles and A. Dobson (Cooper and Iles, this National Science Foundation Evolutionary issue). This is the first Georgia record for Biology Training Grants ( GB 3167 and GB this species. 7346) through Harvard University to S. B. By and large, the troglophilic species of Peck. The field work of 1967 was substan­ tially assisted by Roger Baroody Georgia have much wider ranges than the of Char­ lottesville, Virginia and Alan Fiske of Cam­ troglohitic species and most are not restricted bridge, Massachusetts. to any one given faunal unit or subunit. We would like to thank Elizabeth Staf­ Since, by definition, these species are not ford Ferguson, Cave Files Chairman of the necessarily restricted to cave habitats, their Georgia Speleological Survey, for making powers of dispersal are usually greater than available pertinent location information and those of troglobites and, consequently, their to the numerous cave owners for allowing us ranges are less narrowly defined. Approxi­ access to their property. We also thank mately 20 genera which are represented by Anthony lies, formerly of Marietta, Georgia, troglophilic species throughout most of the and John E. and Martha R. Cooper of the southeastern cave region (Appalachians and University of Kentucky for supplying addi­ Interior Low Plateau) are also found in tional collections, and Dr. and Mrs. Walter Georgia. Four other genera which contain B. Jones of Huntsville, Alabama, for lending troglophiles in Georgia are known otherwise support and encouragement to certain aspects only from caves in the Interior Low Plateau of the project. region, primarily from northern Alabama, We are grateful to the following zoologists for their determinations central Tennessee and Kentucky. of the noted ma­ terial and other helpful comments: Dr. Of some interest are four species of Geor­ Thomas C. Barr, carabid beetles; Dr. Nell gia troglophiles which show a habitat shift B. Causey and Mr. William Shear, mil­ going from north to south along their ranges. lipedes; Dr. Kenneth Christiansen and Dr. These four species-the millipede Cambala David Culver, collembolans; Dr. Richard H. annulata, the opilionid Phalangodes spintur­ Foote, dipterans; Dr. G. E. Gates, oligoch­ nix, the carahid beetle Tachys ferrugineus, aetes; Dr. Willis J. Gertsch, spiders; Dr. and the leiodid beetle Prionochaeta opaca­ Horton H. Hobbs, Jr. and Dr. Joseph Fitz­ are more common in caves on the southern patrick, crayfish; Dr. Theodore Hubbell, ends of their ranges, especially in the Coastal orthopterans; Mr. Leslie Hubricht, molluscs; Plain, than on the northern ends. In the Mr. Robert Matthews, hymenopterans; Dr. northern parts of their ranges, especially in Edward Mockford, psocopterans; Dr. Wil­ the Appalachians, these species are more liam B. Muchmore, pseudoscorpions; Dr. common in the cool, moist habitats of forest Milton W. Sanderson, cantharid and staphy­ linid beetles; Dr. George A. Schultz, oniscoid floors. The preference for caves in the isopods; Mr. Harrison R. Steeves, II, psela­ Coastal Plain is probably influenced to a phid beetles; Dr. Harrison R. Steeves, III, large extent by the absence of ecologically asellid isopods; Dr. Andrew Weaver, centi­ suitable habitats on the surface. pedes; Dr. Pedro Wygodzinski, thysanurans; and Dr. Harry C. Yeatman, copepods.

40 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIK LITERATURE CITED Barr, Thomas C., Jr. 1961. Caves of Ten­ Zinapolys in Ameria north of Mexico. nessee. Tenn. Dept. Conserv. and Com­ Bull. Mus. Comp. Zool., 57:441-504. merce, Div. Geol., Bull. 64, 567 pp. 1946. On some millipedes of 1963a. Studies on the cavernicole Georgia. Ent. News, 57:149-152. Ptomaphagus of the United States (Cole­ Chamberlin, Ralph V. and Richard L. Hoff­ optera: Catopidae). Psyche, 70 ( 1 ):50- man. 1958. Checklist of the millipedes 58. of North America. Bull. U. S. Natl. ~Ius., 1963b. Ecological classification of 212:1-236. cavernicoles. Cave Notes, 5( 2) :9-12. Christiansen, Kenneth. 1964. A revision of 1964. Non-troglobitic Carabidae the Nearctic members of the genus Tomo­ (Coleoptera) from caves in the United cerus ( Collembola, Entomobryidae). Re­ States. Coleopterist's Bull., 18 ( 1): 1-4. vue Ecologie et Biologie du (Sol, 1 4) : 1965. The Pseudanophthalmus of 639-678. the Appalachian Valley (Coleoptera: 1966. The genus Arrhophalites Carabidae). Amer. Midi. N atur., 73 (1): ( Collembola; Sminthuridae) in the United 41-72. States and Canada. Internatl. J. Speleol. 1967. Ecological studies in the 2(1-1) :43-74. Mammoth Cave System of Kentucky. I. Christiansen, Kenneth and David Culver. The biota. Internat. J. Speleol. 3 ( 1-2): 1968. Geographical variation and evolu­ 147-204. tion in Pseudosinella hirsuta. Evolution, 1968. Cave ecology and the evo­ 22( 2) :237-255. lution of troglobites. Evolutionary Bioi., Conde, B. 1949. Campodeides cavernicoles 2:35-102. de Ia region des Appalaches. Notes Bio­ 1969. Evolution of the ( Coleop­ speologiques, 12 ( 4) : 125-137. tera) Carabidae in the southern Appalach­ Cooper, John E. 1968. The salamander ians. In P. C. Holt ( ed.), The distribu­ Gyrinophilus palleucus in Georgia, with tional history of the biota of the southern notes on Alabama and Tennessee popula­ Appalachians. Part I: Invertebrates, pp. tions. J. Alabama Acad. Sci., 39:182-185. 67-92. Virginia Polytechnic Institute Press, In Press. Biological studies in Blacksburg. Shelta Cave, Alabama. Proc. 5th Internatl. Bowman, T. E., R. Prins, and B. F. Morris. Congr. Speleol., Stuttgart, Germany. 1968. Notes on the harpacticoid copepods Crosby, C. R. and S. C. Bishop. 1924. Attheyella pilosa and A. carolinensis, as­ Notes on the Opiliones of the southeastern sociates of the crayfishes in the eastern United States with descriptions of new United States. Bioi. Soc. Washington, species. }. Elisha Mitchell Sci. Soc., 40 Proc., 81:571-586. ( 1-2) :8-26. Brandon, R. A. 1966. Systematics of the Curran, C. H. 1965. The families and salamander genus Gyrinophilus. Illinois genera of North American Diptera. 2nd Bioi. Monograph, 35:1-86. Rev. Ed. Henry Tripp Publ., Woodhaven, Causey, Nell B. 1959. Some cavernicolous New York, 515 pp. millipedes from the Cumberland Plateau. Dearolf, K. 1953. The invertebrates of 75 }. Tennessee Acad. Sci., .34( 4) :229-237. caves in the United States. Proc:. Penn­ Chamberlin, Joseph C. 1962. New and sylvania Acad. Sci., 27:225-241. little-known false scorpions, principally Gertsch, W. J. 1960. Descriptions of from caves, belonging to the families American spiders of the family Symphy­ Chthoniidae and Neobisiidae (Arachnida, tognathidae. American Mus. Novitates, Chelonethida). Bull., Amcr. Mus. Natur. no. 1981, pp. 1-40. Hist., 123 ( 6) : 299-352. Gates, G. E. 1959. Earthworms of North Chamberlin, Ralph V. 1925. The genera America caves. Bull. Natl. Speleol. Soc., Lithobius, Neolithobius, Gonibius, and 21 ( 2) :77-84.

VoLUME 33, Nul\IBEH 1, }ANUARY 1971 41 Gill, Gordon D. 1962. The heleomyzid 1965. Redescriptions of two poorly flies of America north of Mexico ( Diptera: known species of cavernicolous rhagidiid Heleomyzidae). Proc. U. S. Natl. Mus., mites (Acarina: Trombidiformes) from 113( 3465) :495-603. Virginia and Kentucky. Acarologia, 7 ( 4): 1965. Heleomyzidae. In Alan 654-662. Stone, C. W. Sabrosky, et al. A cata­ 1969. Biogeography of the fresh­ logue of the Diptera of America north of water amphipod crustaceans ( Gammari­ Mexico, pp. 808-816. U. S. Dept. Agri­ dae) of the central and southern Appa­ culture, Agr. Res. Serv., Agr. Handbook lachians. In P. C. Holt ( ed.), the no. 276. distributional history of the biota of the Goodnight, Clarence J. and Marie L. Good­ southern Appalachians. Part I: Inverte­ night. 1942. New Phalangodidae (Pha­ brates, pp. 19-50. Virginia Polytechnic langida) from the United States. Amer. Institute Press. Mus. Novitates, no. 1188, pp. 1-18. 1971. The fauna of Pennsylvania 19.53. Taxonomic recognition of caves. In the caves of Pennsylvania, variation in Opiliones. Systematic Zool., Pennsylvania Topographic and Geol. Surv. 2(4):173-179. Bull. (in press ) . 1960. Speciation among cave Hubbell, Theodore H. 1936. A mono­ Opilionids of the United States. Amer. graphic revision of the genus Ceuthophi­ Midi. Natur., 64( 1) :34-38. lus ( Orthoptera, Gryllacrididae, Raphi­ Gurney, Ashley B. 1943. A synopsis of dophorinae). Univ. Florida Publ. Bioi. the psocids of the tribe Psyllipsocini, in­ Sci. Ser., 2( 1): 1-551. cluding the description of an unusual new Hubricht, Leslie. 1943. Studies on the genus from Arizona. Annals Ent. Soc. Neartic freshwater Amphipoda, Amer., 36:195-220. III. Amer. Midi. Natur., 29(3):683-712. Hatch, Melville H. 1933. Studies on the Leptodiridae ( Catopidae) with descrip­ 1962. New species of Helicodis­ tions of new species. J. New York Ent. cus from the eastern United States. Soc., 41:187-239. Nautilus, 75:102-107. Hobbs, Horton H., Jr. 1959. Malacostraca 1964a. Land snails from the ( Astacidae). In W. T. Edmondson ( ed.), caves of Kentucky, Tennessee, and Ala­ Ward and Whipple's freshwater biology, bama. Bull. Natl. Speleol. Soc., 26( 1): pp. 883-898. 33-36. Hoffman, Richard L. 1958. Appalachian 1964b. The land snails of Geor­ Cambalidae: Taxonomy and distribution gia. Sterkiana, 16:5-10. ( Diplopoda: Spirostreptida). J. Wash. 1965. Four new land snails from Acad. Sci., 48( 3) :90-94. the southeastern United States. Nautilus, Holsinger, John R. 1963a. Annotated check­ 79(1) :4-7. list of the macroscopic troglobites of Vir­ Hyman, Libbie H. 1954. North American ginia with notes on their geographic distribution. Bull. Natl. Speleol. Soc., triclad Turbellaria Viii. Three new cave 25:2.3-36. planarians. Proc. U. S. Natl. Mus., 103: 1963b. Studies on the ecology 563-573. and geographic distribution of macro­ Loomis, H. F. 1939. The millipedes col­ scopic cavernicolous invertebrates of the lected in Appalachian caves by Mr. Ken­ central Appalachians. U npubl. masters neth Dearolf. Bull. Mus. Comp. Zool., thesis, 112 pp. 86( 4): 165-193. 1964. The biology of Virginia 1943. New cave and epigean mil­ caves. In H. H. Douglas, the Caves of lipedes of the United States, with notes Virginia, pp. 57-74. Econ-Print, Falls on some established species. Bull. Mus. Church, Virginia. Comp. Zool., 92 ( 7): 373-410.

42 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN Mockford, E. L. 1950. Psocoptera of Peck, Stewart B. and James H. Peck. 1967. Illinois. Proc. Indiana Acad. Sci., 60: Preliminary survey of the terrestrial in­ 192-204. vertebrate cave fauna of Alabama. J. Muchmore, William B. 1966a. Two new Alabama Acad. Sci., 38:216-217. species of Kleptochthonius ( Arachnida, Pylka, Joseph M., and Richard D. Warren. Chelonethida) from a cave in Tennessee. 1958. A population of Haideotriton in J, Tennessee Acad. Sci., 41:68-69. Florida. Copeia, no. 4, pp. 334-336. 1966b. A cavernicolous pseu­ Reddell, James R. 1965. A checklist of doscorpion of the genus M icrocreagris the cave fauna of Texas. I. The Inverte­ from southern Tennessee. Ent. News, brata (exclusive of Insecta). Texas J. 77( 4) :97-100. Sci., 17 ( 2): 143-187. 1969. New species and records 1966. A checklist of the cave of cavernicolous pseudoscorpions of the fauna of Texas. II. Insecta. Texas J. Sci., genus M icrocreagris ( Arachnida, Che­ 18( 1) :25-56. lonethida, N eobisiidae, ldeobisiinae). Amer. Mus. Novitates, no. 2392, pp. 1-21. 1967. A checklist of the cave Nicholas, Bro. G. 1960. Checklist of fauna of Texas. III. Vetebrata. Texas J. macroscopic troglobitic organisms of the Sci., 19 ( 2) : 184-226. United States. Amer. Midi. Natur., Schreiber, Richard and Della McGuffin. 64( 1): 123-160. 1969a. 510 feet is fantastic. Natl. 1962. Checklist of troglobitic Speleol. Soc. News, 27 ( 1): 10-16. organisms of Middle America. Amer. 1969b. Further developments in Midi. Natur., 68( 1): 165-188. Ellison's Cave. Natl. Speleol. Soc. News, Park, Orlando. 1947. Observations on 27 ( 3) :42-44. Batrisodes (Coleoptera: Pselaphidae), with Schultz, George A. 1970. Descriptions of particular reference of the American spe­ new subspecies of Ligidium elrodii cies east of the Rocky Mountains. Chicago (Packard) comb. nov. with notes on other Acad. Sci. Bull. 8( 3) :45-132. isopod crustaceans from caves in North 1951. Cavernicolous pselaphid America ( Oniscoidea). Amer. Midi. beetles of Alabama and Tennessee, with Natur., 84( 1) :36-45. observations on the taxonomy of the Shear, William A. 1969. A synopsis of family. Geol. Surv. Alabama, Mus. Pap. the cave millipedes of the United States, 31, 107 pp. with an illustrated key to genera. Psyche, 1958. New or little known spe­ 76(2): 126-143. cies of pselaphid beetles chiefly from Steeves, Harrison R., III. 1963. The southeastern United States. J. Tennessee troglobitic asellids of the United States: Acad. Sci., 33(1):39-74. the stygius group. Amer. Midi. Natur., 1960. Cavernicolous pselaphid 69(2) :470-481. beetles of the United States. Amer. Midi. Natur., 64( 1) :66-104. Steyskal, George C. 1968. The synonymy 1965. Revision of the genus of Amoebaleria sackeni Garrett. Ent. Soc. Batriasymmodes (Coleoptera: Pselaphi­ Washington, Proc., 70( 2): 113. dae). Trans. Amer. Microscopic Soc., Stone, Alan, Curtis W. Sabrosky, W. W. 84( 2): 184-201. Wirth, R. H. Foote, and J. R. Coulson. Peck, Stewart B. 1966. The systematics 1965. A catalogue of the Diptera of and ecology of the cavernicolous Ptoma­ America north of Mexico. U. S. Dept. phagus (Coleoptera: Catopidae) of the Agriculture, Agr. Res. Serv., Agr. Hand­ United States. Unpubl. masters thesis. book no. 276, 1696 pp. 1970. The terrestrial athropod Vandel, A. 1964. Biospeologie-la biologie fauna of Florida caves. Florida Ent. (in des animaux cavernicoles. Gauthier-Vil­ press). lars, Paris, 619 pp.

VoLU:ME 33, NuMBER 1, JANUARY 1971 43 196.5. Les Trichoniscidae cav­ Warren, Richard D. 1961. The obligate ernicoles ( Isopod a T errestria: Crustacea) cavernicoles of Florida. Florida Speleol. de !'Amerique du Nord. Annales de Soc. Special Paper no. 1, pp. 1-10. Speleologie, 20( 3) :347-389.

Manuscript received by editors July 1970

44 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN The Southern Cavefish, Typhlichthys subterraneus, at the Southeastern Periphery of its Range

By John E. Cooper 0 and Anthony lies n

ABSTRACT Two populations of the troglobitic amblyopsid fish, Typhlichthys subter­ raneus Girard, have been discovered in Deer Head Cove headwaters of tributaries of the Tennessee River, including the first reported in Georgia. These are the nearest known localities to Walden Gorge, which divides the Tennessee into an eastern and a western portion. Another locality for this fish in the Coosa River drainage of Alabama is now known. Additional specimens from Sell's Cave, Alabama, the only previously known Coosa locality, have been collected. The !\labile Basin populations do not differ significantly in morphology from several Tennessee River drainage populations.

INTRODUCTION MS). The range apparently does not include The troglobitic amblyopsid fish, Typh­ those parts of eastern Missouri, southern lichthys subterraneus Girard, has the most Illinois, southwestern Indiana, and western extensive range of any North American cave Kentucky which lie between the southern­ fish. This range is discontinuous, with a most limits of Wisconsin glaciation and the western component including northeastern northern limits of the Cretaceous shoreline Oklahoma, central and southern Missouri, of the Mississippi Embayment. and northern Arkansas, and an eastern com­ The southeastern periphery of the range ponent which begins in the south-central of this fish was for some time thought to lie tip of Indiana, extends south in a narrow along the main Tennessee River drainage in band through central Kentucky, widens in northern Alabama and adjacent Tennessee. central Tennessee, and covers most of north­ However, Boschung ( 1961) reported speci­ ern Alabama along both sides of the Ten­ mens from Sell's Cave (Al 876), near Col­ nessee River (although few populations linsville, DeKalb County, Alabama, which from south of the river are known). It has clearly indicated that at least one popula­ recently been discovered in southeastern tion of T. subterraneus existed in the Coosa Kentucky as well (Cooper and Beiter, in River system across the drainage divide.

0 Dept. of Zoology and Institute of Speleology, A member of this population is figured in University of Kentucky. Present address: Commun­ Smith-Vaniz ( 1968). The following new in­ ity College of Baltimore, 2910 Liberty Heights formation concerning Typhlichthys on the Avenue, Baltimore, Maryland 21215. 00 Biological Survey of Alabama Caves, National southeastern edge of its range, including the Speleological Society. initial record for the state of Georgia and

VoLu~rE 33, NuMBER 1, }ANUARY 1971 4.5 the closest records to Walden Gorge, which and crayfish move up with it, probably divides the Tennessee River into an eastern seeking food. and western portion near Chattanooga, is a Fred Byers, a local resident and cave ex­ result of the work of the Biological Survey plorer, says that he has seen white fish in of Alabama Caves, a research project of the Hurricane Cave, Dade County, a cave of National Speleological Society. This paper debouche at or near the valley floor drain­ is Contribution Number 5 from the Survey. ing the east side of Fox Mountain. A search Numbers in parentheses following some in late 1967 did not yield any indication of cave names refer to the Alabama Cave Sur­ fish here. This is far from conclusive evi­ vey system of identifying and locating caves dence that they don't exist in this area, (Tarkington, et al., 1965), which avoids though, since it took many trips to Deer the confusion sometimes resulting from the Head Cove before a specimen was ever fact that many Alabama caves have the seen. same or similar names. Alabama

CoLLECTIONS AND DiscussiON DeKalb County, Deer Head Cove Cave (Survey number not available)-The sec­ Georgia ond specimen of T. subterraneus from Deer Dade County, Twin Snakes Cave-The Head Cove was collected in this cave in the first Georgia specimen of Typhlichthys sub­ spring of 1969 by Dobson and lies. Deer terraneus was captured here in April, 1969. Head Cove Cave is the main resurgence Numerous visits had been made to the cave site for the drainage of Deer Head Cove, during the preceding year, but water there which lies between Fox Mountain to the had either been impossibly high for collect­ east and Pudding Ridge to the west, and ing or non-existent. In February, 1969, A. most of which lies in Alabama. The water Dobson and A. lies saw a large specimen in here goes to ground through a number of an inaccessible sump at the far end of the sinks scattered throughout the cove, then cave. Several subsequent trips were made coalesces and re-emerges as Allison Creek. to the cave, but they were all fruitless until This eventually joins Lookout Creek, a the April trip. In addition to the lone spe­ northeast-flowing tributary of the Tennes­ cimen collected at this time, another, in ex­ see River. The main cave here has an im­ cess of 50 mm long, was observed but not pressive cross-section but is only penetrable captured. for some 30 m before it ends in a sump. Twin Snakes Cave lies in a small bluff Attempts by The Descenders, a diving and bordering Allison Creek, about 1.7 km cave-exploring club from Atlanta, Georgia, north of Deer Head Cove Cave, discussed to penetrate to the large cave hypothesized below. The entrance is a few meters above to lie beyond the sump, have met with no stream level, and the cave itself is short, success. On the surface, about 13.5 m be­ having probably only about 60 m of acces­ hind the expansive entrance, lies a small sible passage. It is normally dry, but after hole which opens onto a steep slope de­ prolonged rain the water table rises slowly scending into a room floored with boulders. and floods the cave with clear water, some­ Along the far edge of the room at the water times completely. table is a small lake where Typhlichthys Other members of the aquatic commun­ was found. This lake probably connects ity here include the crayfish Cambarus bar­ with the resurgence stream, since it rises toni cavatus, and a small, unidentified, and falls rapidly in response to epigean gilled salamander which eluded capture be­ conditions, is sometimes highly turbid, and cause of mud stirred up in crawling through appears to flow as a mass. the water. As in many other caves, when At the time the fish was captured the the water is rising in Twin Snakes the fish main cave stream was in flood. Lakes in the

46 THE NATIONAL SPELEOLOGICAL SociETY BeLLETIN cave were several feet higher than normal The only other macroscopic aquatic fauna and very turbid. Some time had been spent seen in the cave were two larvae of the in vainly trying to locate a specimen in the salamander, Eurycea longicauda longicauda opaque waters. The capture of a single fish X guttolineata, which were observed on silt was finally accomplished by a blind sweep in shallow water at the entrance. A recently of a net through deep, swirling water. A metamorphosed specimen of this salaman­ very small specimen had been previously der was found under a rock on moist soil observed in the cave on a routine collecting 0.5 m from the water's edge. trip in December, 1968, but capture was Sell's Cave is in a deep, wooded sink at not possible. On each of two subsequent an elevation of 213 m, some 120 to 150 111 visits a single fish was observed which also northwest of Big Wills Creek, a tributary proved too elusive for capture. of the Coosa River. It is primarily a large, Other aquatic fauna found in this cave subterranean lake, some 15 111 long and 3 were a single specimen of the petromyzon­ to 5 m wide, which opens to the surface tid, Lampetra sp., and an observed speci­ through a medium-sized, horizontal en­ men of a large, gilled salamander which trance. The edge of the lake is directly ex­ may have been Gyrinophilus palleucus, re­ posed to light, although it is shaded from cently reported from Georgia caves by bright sun by forest canopy for most of the Cooper ( 1968). year. Because of the steepness of the sink DeKalb County, Sell's Cave-( AI 876 )­ there is no outflow from the lake at the en­ Boschung ( 1961) first reported Typhlich­ trance. The water is very clear and cold; thys from the Mobile Basin on the basis of Boschung ( 1961) recorded the temperature 10 specimens collected at this locality by on June 29, 1958, as 14° C. Dr. J. Ramsey him, R. E. Smith, and J. White on June 29, has provided the following physico-chemi­ 1958 ( U AI C 656). Two additional speci­ cal data collected on August 20, 1964 by mens (AU 2067) were collected here by Smith-Vaniz and Greene: water tempera­ W. Smith-Vaniz and G. N. Greene on Au­ ture, 13.3°C.; air temperature, 20.6°C. in gust 20, 1964. On June 22, 1968, Cooper, cave; dissolved 02, 7.4 ppm; C02 8 ppm; HC03, 171 ppm; CaC03, 82 ppm. The floor with the help of R. Graham, A. Dobson, of the lake slopes sharply down from the and L. Guy, collected four specimens and entrance, and water depth quickly increases observed a number of others here by div­ from just a few em to around 3 m. At its ing in the cave. Individual fish on this oc­ furthest accessible point the lake is .5 to 6 casion were found within 10 em of the m deep. There is considerable air space surface and as deep as nearly 2 m, but each above the water in the lake room and there was within 30 em or less of a limestone is rumored to be a short section of dry pas­ wall, submerged ceiling, or ledge. None was sage beyond and above this. No accessible seen on or near the bottom in deep water. underwater leads were found by Cooper All of the captured specimens had heavy in 1968. concentrations of melanophores along the DeKalb County, Browder Cave (AI 445) lateral line and myosepta} divisions, and -This cave is the second Coosa drainage two had concentrations in the dorsal mid­ locality where T. subterraneus has been line. In addition, one specimen had a found. On March 15, 1969, R. Graham, E. reticulate pattern on the posterior half of Steenburn, and A. Dobson observed three the body. This concentrated condition of specimens here and collected two of them. melanophores in cave fish is usually (but Browder Cave is in the drainage of Big not always) associated with openness of the Wills Creek, about 5 k111 north of Fort habitat to light, and that is the situation Payne and 29 air km from Sell's Cave. The at Sell's. accessible cave consists of a single, medium-

VoLUME 33, NuMBER 1, JANUARY 1971 47 sized room with a short side passage. The mous from surface drainage patterns. See, floor of the room slopes very sharply from for example, Woods and Inger ( 1957) and the entrance, and it was at the bottom of Holsinger ( 1969). Ehrlich and Raven this slope that the water containing the fish ( 1969), however, present strong evidence was found. The water was covered with that phenetic similarity in possibly allopa­ muddy scum, an invariable sign of rising tric populations, including those of the water in such caves. According to the owner troglobitic collembolan Pseudosinella hir­ of the cave, it Hoods completely and water suta, carefully analyzed by Christiansen and flows from the entrance when groundwater Culver ( 1968), may be more a matter of rises in response to heavy spring rains. similar selective regimes than any signifi­ Cooper visited the cave in July, 1968, and cant gene flow. found it completely dry. No other aquatic organisms have been seen in Browder Cave. AcKNOWLEDGl\fENTS In first reporting Typhlichthys from the Some of the work on which this paper Coosa River drainage, Boschung ( 1961) is based was supported by a National "tentatively assigned" the specimens to the Science Foundation Graduate Traineeship species T. subterraneus. Smith-Vaniz ( 1968) to Cooper, and a map-purchase grant to the apparently had no difficulty assigning his Biological Survey of Alabama Caves from specimens from Sell's Cave to this species. the Research Advisory Committee of the It was reasonable to suspect that this National Speleological Society. For assist­ peripheral population, in a drainage system ance in the field we would like to thank from which the species was previously un­ Arthur Dobson, Richard C. Graham, Lin known, might be significantly divergent Guy, George McCluskey, Eric Steenburn, when compared with others, but this is not and Randy Thorp, all members of the now indicated by the morphological evi­ National Speleological Society cooperating dence. These specimens exhibit no con­ with the BSAC. We particularly want to sistent phenotypic differences when com­ thank Graham and Dobson, who have taken pared with other populations from the a very active interest in our attempts to in­ Tennessee River drainage in northern Ala­ vestigate the southeastern limits of the bama. This observation will be elaborated range of Typhlichthys and other troglobites. in a future paper dealing with variation in Dr. and Mrs. Walter B. Jones of Huntsville, southern Typhlichthys populations. Woods Alabama, very generously provided Cooper and Inger ( 1957) indicate that T. subter­ with living and working quarters and other raneus shows considerable variation with amenities much too extensive to list here. Dr. Herbert T. Boschung, Director, ~Iuseum respect to some characters throughout its of Natural History, University of Alabama, range, but the inter-populational differences kindly loaned us the Sell's Cave specimens show no consistent geographic pattern and from the University of Alabama Ichthyo­ are of no taxonomic significance. A very logical Collections ( UAIC). Dr. J. D. Mc­ thorough study of variation in cave-depend­ Clung, Auburn U nivesrity (AU). informed ent and cave-independent characters in us of the whereabouts of specimens from Typhlichthys populations is in order. A par­ this cave collected by Smith-Vaniz and tial analysis for nine populations has been Greene. Certain data on this collection were made by Poulson ( 1961). subsequently provided us by Dr. John Ram­ It has been suggested that genetic con­ sey, Cooperative Fishery Unit. Auburn Uni­ tinuity in some aquatic troglobites may be versity. Martha R. Cooper identified the facilitated by exchange through groundwa­ crayfishes and helped in locating caves on ter systems which are more-or-less autono- topographic maps.

48 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN LITERATURE CITED Boschung, Herbert T. 1961. An anno­ Holsinger, John R. 1969. Biogeography of tated list of fishes from the Coosa River the freshwater amphipod crustaceans system of Alabama. Amer. Midi. Natur., ( Gammaridae) of the Central and South­ 66(2) :257-285. ern Appalachians. In P. C. Holt ( ed.). The Christiansen, Kenneth and D. Culver. 1968. distributional history of the biota of the Geographical variation and evolution in Southern Appalachians. Part I: Inverte­ Pseudosinella hirsuta. Evolution, 22 ( 2) : brates, pp. 19-50. Res. Div. Monograph 237-255. 1, Va. Polytech. Inst., Blacksburg. Cooper, John E. 1968. The salamander Smith-Vaniz, W. F. 1968. Freshwater Gyrinophilus palleucus in Georgia, with fishes of Alabama. Auburn Univ. Agri. notes on Alabama and Tennessee popula­ Exp. Stat., Auburn, Ala. pp. 1-211. tions. J. Alabama Acad. Sci., 39:182-185. Tarkington, Terry W., W. W. Varnedoe and Cooper, John E. and D. P. Beiter. In MS. }. D. Veitch. 1965. Alabama Caves. The southern cavefish, Typhlichthys sub­ Huntsville Grotto, Nat. Speleol. Soc., pp. terraneus, in the Eastern Mississippian 1-82. Plateau of Kentucky. Woods, Loren P. and R. F. Inger. 1957. Ehrlich, Paul R. and P. H. Raven. 1969. The cave, spring, and swamp fishes of Differentiation of populations. Science, the family Amblyopsidae of central and 165:1228-1232. eastern United States. Amer. :\hdl. Natur., 58(1) :232-256.

Manuscript received by editors December 1970

VoLUME 33, NuMBER 1, jANUARY 1971 49

Biology of Cave and Deep Sea Organisms: a Comparison

By Thomas L. Poulson ~

ABSTRACT A panel discussion held at the 1969 AAAS meeting is summarized. The cave and deep sea environments have many similarities for the organisms which have adapted to life in them. The similarities and differences in these adaptations are examined for insight into evolutionary ecology and community structure in the two environments. The characteristics of life history and age distribution, sensory adaptation, and metabolic efficiency are discussed. The effects of geographic isolation and Pleistocene invasions are contrasted. Species diversity and the factors influencing it are discussed.

The intent of this paper is to summarize 3. Evolutionary trends in cave a panel discussion held at the 1969 Boston beetles ( Peck ) Meeting of The American Association for 4. Age distributions of deep sea mol­ the Advancement of Science. I arranged lusks: inferences from growth of the program which was sponsored by the inshore and deep sea species Biology Section of The National Speleologi­ (Turner and Rhoades) cal Society. My summary, of course, repre­ B. Sensory adaptations: inference from sents my own distillation of the proceedings allometry and body proportions and and so may not adequately reflect the views behavioral evidence (Poulson) of my colleagues. I hope that anything lost C. Metabolic efficiency: is it general? in that regard is balanced by cohesion. 1. Amblyopsid cave fish: increased PROGRAM OUTLINE metabolic efficiency with in­ I. General introduction (Poulson )-Char­ creased time of isolation in caves acterization of the two environments (Poulson) and reasons for interest in caves and 2. Cave am phi pods: an exception? the deep sea. (Culver) II. Life history, sensory and metabolic 3. Deep sea animals: no difference adaptations-Evidence for the impor­ from inshore species? (Teale) tance of metabolic economy? D. Is food limiting in the deep sea? A. Life history and age distributions 1. Problems of defining food and its I. Amblyopsid cave fish; a paradigm availabili~y ( Darnell) for decreased reproductive po­ 2. Is depth zonation of animals re­ tential (Poulson) lated to food supply? (Rowe) 2. Deep sea brittlestars; an excep­ E. Open discussion tion to the rule? (Schoener) III. The fauna ,. Department of Biology, University of Notre A. History of the biota-Effects of the Dame, :\'otre Dame, Indiana. Pleistocene, persistence of relicts,

VoLUC\IE 33, NuMBER I, jANUARY 1971 51 and multiple invasion vs in situ acterized as old, climatically stable, and speciation non-rigorous (Poulson and White, 1969). 1. Terrestrial cave organisms (Peck) They are therefore predictable in time. 2. Aquatic cave organisms There are, however, major differences be­ (Holsinger) tween these two natural laboratories. Im­ 3. The deep sea epifauna and in­ proved sampling has revealed ( Sanders and fauna ( Sanders) Hessler, 1969) that the bathybenthic oozes B. Explanations of species diversity contain large numbers of species ( infauna) 1. Indices of diversity, general hy­ including many species in the same genus potheses, and examples from feeding by ingesting the sediment and ex­ caves (Culver) tracting bacteria, protozoa, or dissolved 2. The stability/time hypothesis: in­ organic matter. This raises fascinating ques­ shore vs deep sea ( Sanders) tions about the role of competitive exclusion, C. Open discussion particularly since it appears that a given species pattern may be repeated over wide pARTICIPANTS areas even on both sides of an ocean basin. David C. Culver, Dept. of Biology, U. of In caves the habitat is more dissected in Chicago, Chicago, Illinois; John R. Hol­ space and in character, both inter- and intra­ singer, Dept. of Biology, Old Dominion U., cave. There is the question whether cave Norfolk, Virginia; Rezneat Darnell, Dept. communities are stabilized in evolutionary of Biology, Texas A & M U., College Sta­ time; certainly cave faunas are more recent tion, Texas; Stewart Peck, National Museum than deep sea faunas. Finally, the low and Carleton College, Ottawa, Canada; average densities of small animals in caves, Thomas L. Poulson, Dept. of Biology, U. of .0001-1.0/ m 2, make one wonder whether Notre Dame, Notre Dame, Indiana; Gilbert competitive exclusion is even possible. These Rowe, Howard Sanders, and John Teale, densities approach those seen for compara­ Woods Hole Oceanographic lnst., Woods ble sized animals in a photographic survey Hole, Massachusetts; Amy Schoener and of the deep sea epifauna, .01-10/ m2 (Rowe Ruth Turner, Museum of Comparative Zo­ and Menzies, 1970), but are nowhere near ology, Harvard U ., Cambridge, Mass. the high densities, 25-200/ mZ, for the deep sea infauna (Sanders and Hessler, 1969). INTRODUCTION Cave animals, unlike bathybenthic ani­ The major basis for this discussion is the mals, can be maintained in the laboratory similarity in the environmental problems and studied behaviorally or physiologically faced by organisms that live permanently under natural and experimentally modified in caves and those that live permanently conditions. There are few enough cave on the ocean bottoms at depths below 2000 species so that it is possible to study the meters (the bathybenthic). It seems likely biology of each species of the community that the animals in these environments share and its interactions with other species. Thus morphological, physiological, and life his­ it will be possible to determine which spe­ tory specializations due to lack of light and cies characteristics or interactions are associ­ food limitation. These adaptations may re­ ated with communities of high or low late to similarities in community structure diversity. This level of analysis is not yet which are of interest to the evolutionary possible in the deep sea, mainly because of ecologist. Finally, there was the hope that logistic and sampling problems. Defined communication between students of the two sampling of the infauna at one location will environments might yield insights into the be necessary to define interspecific and basis for community structure in other interindividual interactions. Photography is environments. used to sample the larger and less diverse Both caves and the bathybenthos are of epifauna, but w~th this method the more a class of environments that can be char- mobile groups, like fish and shrimp, are

52 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN disturbed and rarely recorded on film. For Peck described a series of elegant studies in the infauna the problems at the initial which he raised surface, troglophilic, and taxonomic level are staggering in them­ troglobitic species of the genus Ptomaphagus selves. Often groups apparently have as through their life cycles in the laboratory. many as 10-25 species in a sample. To be The more specialized troglobites show longer sure that they are really separate species it life cycles, but the development of a non­ is necessary, as Hessler has done for one feeding larva, small clutch size, and large genus of isopods, to compare all life history eggs seen in European species is inex­ stages of each species to assess whether the plicably absent in the North American fine morphological differences between sup­ species. Perhaps the problem of a sedentary posed species are constant and therefore larva finding a rare and superdispersed food real. The next level of analysis, of habitat supply is circumvented by adult feeding or food selection and of metabolic needs, is of young in the specialized species. This underway in caves but is not yet possible seems a reasonable possibility to me since in the deep sea. For whatever reason, and the larvae of N. American species do not it does not seem to be temperature or pres­ have the degenerate sense organs or mouth­ sure, live specimens have not long survived parts seen in European forms that never the trip to the surface and so have scarcely feed. been studied alive. Hopefully perfection of The data on bathybenthic species are deep submersibles will soon allow biologists scattered and fragmentary, especially on to circumYent these problems by in situ size or age distributions, but it is clear that observation. some epibenthic groups do not conform to the paradigm set up for cave fish. Schoener LIFE HISTORY AND AGE DISTRIBUTIONS showed that the brittlestars have an inverse I started this discussion by using ambly­ relation between fecundity and egg size opsid cave fish (Poulson, 1963 and 1969) but, except for a few brooding species, all as a paradigm for the decreased compound lay hundreds to thousands of small eggs. interest rate of population growth selected Each of these species has a depth range of by food limitation over a long period of 1 to 5 thousand meters, and their fecundi­ evolutionary time. The evolutionary series ties do not have the same rank order as of swamp, spring-cave, and cave fish shows specialization inferred from narrowness of decreased fecundity associated with in­ depth distribution. Apparently the epi­ creased egg size and parental care, increased benthic crabs show a similar mix of brood­ generation time associated with increased ing species with few large eggs and non­ age at first reproduction and longevity, and brooding species with many small eggs and a tendency toward skewing of age distribu­ free-living planktonic larvae (Marshall, tions toward older age classes. This evo­ 1954). From fragmentary data it appears lutionary series culminates in a life history that a higher proportion of the infauna show or reproductive strategy which is the anti­ large eggs and direct development (San­ thesis of the "opportunism" shown in sea­ ders and Hessler, 1969). sonal and rigorous environments, and I The few data available that suggest low proposed that long life and the chance for growth rates have not yet been translated repeated reproduction allows eventual re­ into age or even size distributions. Rhoades productive success in an environment that has started to analyze scanning electron may not be conducive to reproduction every micrographs of growth rings in the shells year because of food or mate scarcity and of bivalve mollusks ( nuculoids) from the predation. intertidal to the deep sea. His preliminary Peck reviewed his work on scavenger data show about twice as many rings in beetles ( 1968 and in progress) which, as deep sea as for similar sized inshore species, with most cave organisms (Poulson, 1964), but it is not yet clear whether these rings supports the paradigm given for cave fish. are equivalent because the daily and tidal

VoLU~IE 33, NuMBER 1, JANUARY 1971 53 reabsorption of shell inshore is absent in the the open pits in the bathybenthic cetomimid deep sea. Data available on the size dis­ fish ( Gyrinomimus, Richardson and Garrick, tributions of deep-sea forms are suspect be­ 1964) may serve to expose the lateral line cause of the possible size sorting and loss organs and increase their exposure in a of small individuals during retrieval of the different way than in amblyopsid cave fish sampling gear from the ocean depths. Here where the evolutionary solution has been the core sampling being done by Sander's to place the organs in rows on the surface group will provide important information of the head. A large head may reflect in­ on a comparative basis even if there are creased sensory endowment of the less obvi­ no data on growth rates to translate size ous chemosensory systems in the macrourid into age distributions. Despite lots of un­ and brotulid fish which are also convergent ambiguous data on size distributions of in their 'vacuum cleaner' mouths (Bruun, troglobites, growth rates have been deter­ 1966; Marshall, 1954). Of course some mined only for fish. species, like the double-tailed fish Halopo­ phyrus, have obvious contact chemical SENSORY .ADAPTATION sensors on whiskerlike barbels that trail Cave and bathybenthic organisms face along the bottom as the fish swims above, similar selective pressures, so we would much as catfish do. Just as increased rela­ expect parallels in sensory adaptations. tive head size gives more area for lateral However, unlike caves, the presence of lumi­ line or taste organs in fish, so the shape of nescent organs in some species of bathy­ crustaceans does for tactile and chemo­ benthic animals (but not in as high a receptors. Comparing two species of the proportion as in the bathy- or mesopelagic) same body weight, the one with a more results in maintained selection for eyes and "slight" body build and long attenuated pigmentation. On the other hand, deep sea limbs has a higher total surface area and and cave animals both have to hunt in the so more sense organs per unit body weight. dark because the deep sea is still basically However, this modification in body build aphotic. Unfortunately it is not yet possible does not seem to be common in bathy­ in the case of deep sea forms, as it has benthic crabs and shrimp, although it is been for cavefish (Poulson, 1963; Poulson the general rule in cave crustaceans. These and White, 1969) and some invertebrates calculations, of course, prove nothing with­ (Poulson, 1964; Cooper, 1969), to relate out experimental data like Banta's (in Poul­ relative food limitation to efficiency of food son, 1964). More recently, Cooper ( 1969) finding. has given behavioral data, in terms of time The morphological adaptations of the taken to react and distance moved before epibenthic deep sea fish and crustaceans capture, for increased food finding efficiency may fuction either to avoid stirring up the in a cave crayfish vs a related surface form. soft bottom oozes where food occurs-as in the stilt-fins of the bathypteroid fish, the METABOLIC EFFICIENCY long thin legs of crabs and shrimp, and There is considerable fragmentary evi­ the high oil content of the liver that confers dence that aquatic troglobites have lower neutral buoyancy to a six-gilled shark­ metabolic rates than troglophilic or surface and/or as adaptations for finding food and species of the same group, but in most cases detecting predators. It is also possible that the controls have not been adequate. I the stilt-fins function as I have suggested argued that amblyopsid fish, which are at for the long spindly legs of specialized cave the top of the food web, are most likely salamanders (Poulson, 1964 and Brandon, to be food limited. Indeed it is clear that this issue) where the lateral line water there is a reduction of growth and metabolic movement detection system is given a wider rate in the evolutionary series of surface, "field of view" by being raised off the troglophilic, and troglobitic species (Poulson, substrate. In a more straight forward vein 1963). This metabolic efficiency is not at the

54 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN expense of activity since routine metabolic showed lower growth rates for the same rate goes down while total foraging activity species of boring clams in wood panels sub­ and percentage of time that a fish is active merged at 2000 meters than in those sub­ go up. In contrast, Culver presented data on merged at 200 meters, thus suggesting a amphipods that showed no predictable rela­ pressure effect on metabolism. tion between cave adaptation and activity The morphology of bathypelagic animals or metabolic rate either between troglophilic is correlated with energy economy ( Mar­ and troglobitic species of the genus Sty­ shall, 1960), so it is worthwhile to consider gonectes or within a polytypic troglophilic such indirect evidence for energy economy species, Gammarus minus (Culver and Poul­ among bathybenthic species. Childress ( 1969 son, 1971a). The fish and amphipod data and personal communication) shows that are rationalized when it is realized that: the metabolic rate of bathypelagic shrimp 1. Amphipods are lower down in the food decreases in parallel with depth and that web than fish; and 2. The caves of the there is a parallel decrease in size, muscula­ Valley-Ridge (where the amphi pods were ture, and gill area. This parallels the studied) have discrete organic debris in­ morphological deep sea "syndrome" so well washed at all seasons but the caves of the described by Marshall for fish where the Interior Low Plateaus (where the fish occur) metabolically expensive swim bladder is get only diffuse input of unidentifiable vestigial and even the kidney may be re­ organic matter and only in the spring (see duced to a single renal tubule! Unfortu­ Jegla and Poulson, 1970). This is just an­ nately neither the bathybenthic shrimp nor other example of how caves are a less uni­ fish have been examined in this regard. But form habitat than the deep sea; the vari­ I infer, as does Marshall, from specimens ations in food supply between caves may and in situ photographs that this syndrome be equivalent to the variation in food with is not present in any bathybenthic fish. depth in the deep sea. They have swim bladders and well de­ veloped scales and musculature, and their The matter of energy economy in deep gills are normally developed. However they sea animals rests largely on inference from do tend to be small, and so the possibility morphology or distribution of animal bio­ of subtle differences in gill area and growth mass and potential food with depth, since rate, such as that documented in similar food metabolic studies are lacking. Meta­ sized cave fish, can not be discounted until bolic studies are ambiguous. The epibenthic metabolic studies can be done. species came up in poor condition and there are rarely inshore relatives to serve Rowe argued for the importance of food as a baseline for the infauna which came limitation since, with the exception of deep up in good condition protected in the ocean trenches which act as nutrient traps mud. The necessary controls for such and would be a good control for the effects studies-related species, comparable size of food vs pressure, there is a decrease and activity, adjustment to the metabolic in abundance and biomass with depth chamber-have been recently reviewed by (Menzies and Rowe, 1969; Sanders and Culver and Poulson ( 197la). The data Hessler, 1969) which is paralleled by a at hand do not meet all these conditions decrease in potential food. The decrease in but are worth reviewing. Teale reported biomass below 2000 meters, on the order neither lower metabolic rates nor any effect of 25-50 fold per 500 meters, parallels a of pressure for a few deep sea brittlestars, decrease in downslope transport of discrete a sipunculid worm, an isopod, and a clam organic matter, such as turtle grass and as compared to inshore species but empha­ sargasso weed, but dissolved and fine par­ sized the preliminary nature of his results. ticulate organic matter that may "rain" His techniques are such that routine or down from above shows little variation and acute, and therefore near maximum, meta­ shows no clear relation to animal biomass. bolic rates are measured. However, Turner This brings up the real problem, for cave

VoLUME 33, NuMBER 1, jANUARY 1971 5.5 as well as the deep sea, as to what is or is deep sea. The major theme of the discus­ not food, for these detrital based food webs. sion was the difference in possibility for Cave-bathybenthic comparisons of food geographic isolation and subsequent speci­ limitation must be made with caution be­ ation-clearly high in caves, which are cause there are no good data on organic discontinuous in time and space, and seem­ matter in or on the substrate and, even ingly low in the deep sea, which is uniform when available, it is not clear whether that in bottom type and lacks obvious geographic measured by the usual combustion or oxi­ barriers. dation methods reflects energy extractable Peck and Holsinger showed that the by organisms. Is the low density and bio­ origin of some aquatic troglobites via in­ mass of cavernicoles related to the observa­ vasion of seacoast caves or sand and gravel tion that particulate matter washed into is quantitatively less important than invasion caves may remain for several years and that from interior karst areas during glacial ad­ its nutritive value is therefore low? If so, vances and retreats. Highland terrestrial how do we explain the apparent paradox species adapted to cool, moist forests or that deep sea biomass is higher than for aquatic species adapted to cool streams caves even though the particulate matter spread into the lowland during glacial ad­ contains less organic matter? The answer vance and either moved back into the may be in the availability to the animals. uplands or became restricted to isolated There is suggestive evidence, even for in­ pockets of cool microclimate, such as shore areas of the sea, for bacterial special­ springs and cave entrances, as the glacier ists on cellulose, chitin, and other tradi­ started to retreat again. As the glaciers tionally refractile organic compounds, for continued to retreat, the terrestrial habitats complex recycling of animal secretions, in the lowlands became dry-warm and the regurgitations, and egestions, and for direct streams became warm. As a result, the only uptake of dissolved organic molecules in­ cool-adapted populations to persist in the cluding amino acids, sugars, and vitamins lowlands were those that had access to (see references in Darnell, 1967). The caves where it remained cool all year. Those morphology of inshore vs deep sea deposit­ species that persisted in caves and evolved feeding bivalves suggests the extremes for into specialized troglobites are said to have recycling of fecal pellets. Inshore the tel­ been preadapted to caves by their physi­ liniid bivalves have a short gut and the ological tolerance to cold and their ability recycling may involve egestion of feces to sense food and mates in dark areas like low in protein and high in carbohydrates, forest litter or under rocks and ledges in growth of bacteria, and repetitions of the streams; thus they were already facultative cycle until the carbohydrate is gone. In cave forms, i.e. "troglophiles". Generally the midwater species the efficiency of di­ few of these troglophilic ancestors persisted gestion presumably increases due to a even in the highlands, and so the troglobitic longer and more coiled gut with fewer cave derivatives are said to be relicts. recyclings, and in the deep-sea it appears Like most cave species some of the deep­ that bacteria have been added to the gut sea crustaceans are Pleistocene relicts, but so that there is a symbiosis (Allen and others, like the protobranch mollusks that Sanders, 1960). date from the Cambrian, are much more ancient relicts. However, unlike the cave HISTORY OF THE BIOTA species, it is not clear how the deep sea Many of the relevant ideas about cave species became isolated. Perhaps southward biogeography have been either recently re­ spread of a boreal inshore fauna during viewed (Poulson and White, 1969) or are glacial advance was followed by a down discussed elsewhere in this issue (Culver; slope instead of northward movement along Peck and Holsinger), so my review will temperature isoclines as the glaciers re­ focus on differences between caves and the treated. A variant of this is suggested by

56 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIK the disjunction in faunas at the continental prefers to think that in situ microgeographic slope thermocline which is amplified at the speciation has been important in the deep borders of marine basins such as Mediter­ sea since there have been such massive adap­ ranean-Atlantic. During the Pleistocene tive radiations within single genera (San­ there may have been little difference be­ ders, 1968). Also in his favor is a good tween the two basins but, as they diverged correlation between species richness, nar­ postglacially toward the 37.5-39.0 ofoo vs rowness of geographic distribution, and life 36.0-36.5 °/oo salinity and 13-14 o vs 2.0- history traits that promote the chances for 2.50C temperature difference of today, some microgeographic speciation. For example a of the cold adapted Mediterranean deep-sea given area has 5 species of brittlestars, 25 species could have invaded the Atlantic bivalve mollusks, and 85 crustaceans. The and so added to its diversity. And, as a crustacean species have the narrowest depth third variant, species diversity of a basin range, suggesting the narrowest physiologi­ could increase if multiple invasions of the cal tolerance, and the narrowest geographic same inshore ancestor gave rise to different range, suggesting the most limited larval species as Barr believes happened with some dispersal. Both traits go along with the cave beetles and springtails ( Christiansen presence of few large eggs, brooding of and Culver, 1968 and 1969). In the deep eggs, and absence of planktonic larvae that sea, local diversity would be increased when would disperse over a wide range of depths those separate invading populations became and geographic areas. Thus the possibilities adapted and spread enough to overlap in for microgeographic speciation, particularly geographic range. This overlap subsequent in such sedentary infaunal crustaceans, can­ to multiple invasion is easy to envision for not be discounted. The possibilities for such caves but the more striking faunal overlap, speciation seem large in view of the demon­ on both sides of the Atlantic Ocean, along strated species differences in highly mobile a depth contour rather than between depths species like fish, between ocean basins at one spot (Sanders and Hessler, 1969) and trenches in the bathybenthic rat-tails makes it difficult to explain the deep sea ( Macrouridae) and between water masses faunal overlap. Whether planktonic larvae that differ subtly in salinity, temperature, of the epibenthic species disperse widely and nutrients in the mesopelagic lantern and survive or settle at only one depth is fish ( Melamphiidae). unknown. Whatever the case, the explana­ The influence of the Pleistocene on the tion of the narrow depth distribution re­ deep sea is debated, but it is clear that it quires a much narrower physiological toler­ has had a major influence on speciation in ance to pressure than has been suggested caves. Despite possibilities for speciation by my work on ability of cave fish to resulting from multiple invasions of one metabolically compensate for temperature ancestor during one glacial epoch (see change or even by Teales' work on the Christiansen and Culver, 1969) most work­ effects of pressure on deep sea forms ( see ers agree that the diversity in caves is above). mainly the result of multiple invasions by different \Vhatever the chances of multiple in­ troglophiles during three or four Pleistocene vasion, it is clear there has been plenty of advances and withdrawals of time for diversity to slowly increase in the the ice sheet. The several grades of mor­ deep sea since, unlike caves, the tempera­ phological specialization in cave millipedes tures and salinities have changed little over and amphipods and of physiological special­ geologic time, even during the Pleistocene, ization in cave fish are most easily explained and so new invasions probably far out­ by waves of invasion and isolation that coin­ number extinctions. Put another way, it cide with different glacial epochs. If this may not be necessary to invoke in situ is correct, then the drastic runoff, flooding, speciation in the deep sea to explain its scouring, and temperature changes sug­ high species richness. Sanders, however, gested by some for periglacial karst areas

VoLUME 33, NuMBER 1, }ANUARY 1971 57 either was not so drastic or was not so perature changes on a seasonal basis. He widespread that species already present feels that competition and predation are could not survive. Even the terrestrial and more severe inshore than in the highly larger aquatic species, that could not gain diverse and biologically controlled deep-sea access to the well-buffered non-cave ground­ communities where the environment is water available to the small isopods and constant in time with extremely predictable amphipods, have survived at least one glacial and therefore presumably less intense inter­ epoch in caves. actions. I pointed out that the importance of competitive exclusion to an explanation SPECIES DIVERSITY of the high diversity in his samples can only Culver started this last part of the pro­ be answered with more detailed sampling. gram with an elementary review of di­ Despite Sanders arguments to the contrary, versity indices. He differentiated between a repeated species pattern from dredge indices which merely reflect species number, samples at about the same depth and place i.e. indices of species "richness", and those, but from different directions does not obvi­ like the information theoretic index, that ate the problem that his samples are too reflect both species number and relative big and uncontrolled to detect small differ­ abundance. For example, species richness ences in substrate or species composition would not differ between a sample of 3 that may be important to the tiny, sedentary species with 10 individuals each and 3 with infauna. Sanders' group is now doing the 15, 10, and 5, respectively. However, an necessary repeated core sampling at one information theory index would give the location to answer this criticism. Such latter a lower diversity because the relative sampling will allow him to assess the role abundance of the three species is less even of species co-occurrence, and thus presum­ than in the fanner sample (see references ably interaction, and microhabitat patchiness in Poulson and Culver, 1969 or Culver and by correlating shapes of species abundance Poulson, 197lb for more detail). This dis­ curves in a series of cores with character tinction is critical with comparisons of of substrate, amount of "food", number of within- and between-cave diversity because related species or potential predators, etc. there are so few species, but in the deep If the cores show no differences in micro­ sea each sample contains so many species habitat patchiness, then it will become im­ and individuals that these refinements are portant to look for coevolutionary inter­ not critical. Thus Sanders uses a species actions, like those plant/pollinator and richness index and standardizes his measure­ plant/herbivore interactions that have aug­ ments to 1000 individuals using a so-called mented diversity of plant/insect communi­ rarefaction method. A typical abyssal plain ties, and feeding specificities, like those sample rarified to 1000 individuals contains shown by fish which specialize on scales, 5 species of brittlestars, 19-30 bivalve mol­ blood, fin rays, and rotting flesh of other lusks, 50-70 polychaete worms, and 70-125 fish in the tropics. crustaceans including 12 taniad shrimp, 23 The diversity of cave communities is cumacean shrimp, 30 amphipods, and 35 intermediate between the benthic communi­ isopods. ties of estuaries and the deep sea, but Sanders exp] ained that his stability/ time Culver and I ( 1969 and unpublished) do hypothesis for diversity of marine benthic not believe that this allows us to simply communities has been derived to fit field say that control of diversity in caves is observations (Sanders 1968; Slobodkin and intermediate between that in estuaries and Sanders, 1969). In his view the low species the deep sea, i. e. a mix of biotic and richness of inshore communities is due to physical control. We pointed out that ob­ physical control by extreme fluctuations, for jective categorization of "physical" and example in an estuary where salinity and "biotic" are necessary to formulate a hypo­ current change on a daily basis and tem- thesis for the control of diversity and to

58 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN devise meaningful field and laboratory ex­ ratory. The small cave amphipod selects periments to test the hypothesis. Culver small gravels whether or not its larger then summarized a scheme that includes all relative is present thus indicating no present factors that potentially affect diversity, start­ day effect of competition. But the large cave ing when a new habitat is "opened" for amphipod and the isopod only showed colonization. Initially there is a stochastic microhabitat selection when the large non-interactive equilibrium that depends on spring-cave amphipod is present thus indi­ intrinsic probabilities for colonization and cating ongoing competition. This kind of extinction of each potential colonizer. After competition which leads to competitive ex­ the numbers of individuals increases, there clusion is due to active interference of one are three parallel kinds of control. Until species with another's behavior. It is hard we have data to the contrary, we consider to envision in a uniform habitat like the that these operate simultaneously: 1. Biotic deep sea especially with so many species interaction such as competition, predation, present. In discussion I suggested the possi­ and mutualism; 2. Energy flow as deter­ bility that since there has been more evo­ mined by rates of turnover, productivity, lutionary time available in the deep sea, and life history and stabilization of energy many deep sea species show invariant micro­ flow as determined by connectedness of habitat selection like the small cave amphi­ food webs; and 3. Physical control includ­ pod studied by Culver. This suggestion is ing the subparameters of .,climate"-rigor based on the observation that competitive (extremes), variability (variance), and pre­ exclusion cannot even be demonstrated in dictability (autocorrelation )-and structural terrestrial cave communities of Mammoth heterogeneity of the habitat. After this Cave that are much more complex than the introduction, examples were given for ter­ West Virginia stream communities but show restrial and aquatic cave communities. nowhere the complexity of the deep sea communities. Culver discussed his data (1970; 197la, b) As with from 28 West Virginia stream caves where Culver's study, our work on ter­ different species combinations are present. restrial communities in the 110 kilometer Flint He illustrated the random nature of coloni­ Ridge Cave System (Poulson and Culver, zation and extinction of an isopod and three 1969 and continuing) has shown species of amphipod in a small cave and a significant negative effect of climatic rigor (flooding suggested that differences in species compo­ and evaporation rate) and a posi­ tive effect sition in larger caves where populations are of structural heterogeneity of the habitat (substrate more persistent could also be partially ex­ diversity) on between­ habitat species plained by a balance of colonization and diversity. extinction. :Most of the remaining between­ Overall the terrestrial system is a better cave variance in community composition is analogue of the deep sea than the aquatic associated with length of the cave stream cave system because the terrestrial animals and its susceptibility to spring flooding. He were sampled in 50 sites within a single showed that field data on seasonal changes large cave where all 14 regular species can in amphipod density and laboratory studies get to every sampling site, where the char­ on washout and injury in an artificial stream acteristics of a sampling site can be repli­ also confirm the importance of flooding to cated to assess any random or seasonal control of community composition. The effects, and where there is a triad of re­ length of cave is negatively associated with lated beetle species which allows assessment flooding but more important is its positive of the role of competition in control of correlation with number of tributary streams diversity. and so with possible avoidance of compe­ Correlation of species diversity with sub­ tition through microhabitat selection. He strate diversity in our original study and related field data on microhabitat/amphipod suggestions of fighting during laboratory distribution to habitat selection in the labo- interactions first led us to look for explana-

VOLUME 33, NUMBER 1, }ANUARY 1971 59 tions of species diversity in terms of differ­ results thus far are that species diversity, ential substrate selection by the three beetle evenness, and repeatability are highest in species. However, there was no competitive the least rigorous, least variable, and most exclusion along the niche dimension defined predictable passages. The differences in the by substrate since competition coefficients other cave species found in these three calculated using each species tested alone passages lend weight to my current hy­ did not differ from the coefficients found pothesis that the combination of high mo­ with species tested in pairs or triads. In bility, high reproductive potential, and addition no species showed consistent prefer­ broad physiological tolerance-what I have ence for fine gravel, sand or mud. So what, called ecological opportunism-is character­ if any, niche dimension is important in istic of animals in communities with low separating these beetles? It appears that species diversity. Thus much of my current differences in species biology are part of the emphasis is on beetle biology, particularly answer. We now have data for three stream physiological tolerance. The other emphasis passages that do not differ much in sub­ will be on perturbation studies in nature to strate diversity or "food" (average weight test my hypothesis by modifying such things loss after ignition) but do differ greatly in as food to see if, as predicted, the local rigor, variability, and predictability of flood­ species diversity drops due to colonization ing, food input, and potential drying. The by opportunists, especially crickets.

LITERATURE CITED Allen, J. A. and H. L. Sanders. 1966. See 197lb. Analysis of simple cave Sanders and Hessler, 1969. communities. III. Control of abundance. Barr, T. C., Jr. 1967. See Poulson and Amer. Midi. Natur., 85:173-187. White, 1969 and critique by Culver, this Culver, D. C. and T. L. Poulson. 197la. Bulletin. Oxygen consumption and activity in close­ Bruun, A. 1956. Animal life of the deep ly related amphipod populations from cave sea bottom. Chapter in "Notes from the and surface habitats. Amer. MicH. Natur., Galathea deep sea expedition." Macmil­ 85:74-84. lan, N.Y. 1971b. Community boundaries: Childress, J. J. 1969. The respiration of Faunal diversity around a cave entrance. deep-sea crustaceans as related to their Ann. Speleologie, 26: (in press). depth occurrence and the oxygen minimum Darnell, R. M. 1967. The organic detritus layer. Amer. Zool., 9:1100 (Abstract). problem and its relation to the estuarine Cooper, M. R. 1969. Sensory specialization ecosystem. pp. 374-382 in .. Estuaries", and allometric growth in cavernicolous Amer. Assoc. Adv. Science. crayfishes. Proc. IV Internatl. Cong. Holsinger, J. R. 1967. Systematics, speci­ Speleol. (Yugoslavia), 4-5:203-208. ation, and distribution of the subterranean Christiansen, K. and D. C. Culver. 1968. amphipod genus Stygonectes ( Gammari­ Geographical variation and evolution in dae). U. S. Natl. Mus. Bull., 259:1-176. Pseudosinella hirsuta. Evolution, 22:237- 1969. Biogeography of the fresh­ 255. water amphipod crustaceans ( Gammari­ 1969. Geographical variation and dae) of the central and southern Ap­ evolution in Pseudosinella violenta. Evo­ palachians. In P. C. Holt ( ed), The lution, 23:602-621. distributional history of the biota of the Culver, D. C. 1970. Analysis of simple southern Appalachians. Part I: Inverte­ cave communities. I. Caves as islands. brates, pp. 19-50. Virginia Polytechnic Evolution, 24:463-474. Institute Press, Blacksburg, Va. 197la. Analysis of simple cave Jegla, T. C. and T. L. Poulson. 1970. Cir­ communities. II. Niche separation and cannian rhythms-1. Reproduction in the species packing. Ecology, 52: (in press). cave crayfish, Orconectes pellucidus in-

60 THE NATIONAL SPELEOLOGICAL SociETY BcLLETIN ermis. Comp. Biochem. Physiol., 33:347- Internatl. Cong. Speleol. (Yugoslavia), 355. 4-5:189-192. Marshall, N. B. 1954. Aspects of deep sea Poulson, T. L. and D. C. Culver. 1969. biology. Hutchison, London. 380 p. Diversity in terrestrial cave communities. 1960. Swimbladder structure of Ecology, 50:153-158. deep-sea fishes in relation to their system­ Poulson, T. L. and W. B. White. 1969. The atics and biology. Discovery Repts., 31: cave environment. Science, 165:971-981. 1-122. Richardson, L. R. and J. A. F. Garrick. 1964. A Menzies, R. }. and G. T. Rowe. 1969. The new species of Gyrirwmimus distribution and significance of detrital (Pisces, Cetomimidae) from New Zealand. turtle grass, Thallassia testudinata, on the Copeia, 1964:523-525. Rowe, deep-sea floor off North Carolina. Int. G. T. and R. J. Menzies. 1969. Zon­ Revue ges. Hydrobiol., 54:217-222. ation of large benthic invertebrates in the deep-sea off the Carolinas. Deep-sea Res., Peck, S. B. 1968. A new cave catopid 16:531-537. beetle from Mexico with a discussion of Sanders, H. L. 1968. Marine benthic diver­ its evolution. Psyche, 75:91-98. sity: A comparative study. Amer. Natur., Poulson, T. L. 1963. Cave adaptation in 102:243-282. Amblyopsid fishes. Amer. Midl. Natur., Sanders, H. L. and R. R. Hessler. 1969 70:257-290. Ecology of the deep-sea benthos. Science, 1964. Animals in aquatic environ­ 163:1419-1424. ments: Animals in caves. In D. B. Dill Slobodkin, L. B. and H. L. Sanders. 1969. ( ed. ) , Adaptation to the environment, On the contribution of environmental pre­ Handbook of Physiology, 4, pp. 749-771. dicability to species diversity. In Diver­ Williams and Wilkins, Baltimore. sity and stability in ecological systems, 1969. Population size, density, 22nd Brookhaven Symposium in Biology, and regulation in cave fishes. Proc. IV pp. 82-95.

Manuscript received by the editor December 1970

VoLUME 33, NuMBER 1, }ANUARY 1971 61

The Classification of Cavernicoles

By Elery Hamilton-Smith #

ABSTRACT A seven-category system for the classification of cavernicoles is proposed, primarily on the basis of Australian experience. Parasites are separated from the Schiner-Racovitza categories to form a separate category; it is suggested in line with some other workers that the term trogloxene should not be used for accidental species; threshold species not found within the dark zone are also separated, but the complexity of the threshold fauna is emphasized. The dark zone fauna is thus separated into four categories: trogloxenes and troglobites as in the Schiner-Racovitza system, with troglophiles divided into two levels, being those species also occurring in the epigean environment (first level} and those confined to caves (second level).

The ecological classification of cavern­ these three terms are the basis of present icoles has been widely discussed, and this usage. discussion has led to the coining of elaborate Alternative patterns of classification or of or even picturesque terminology, much of terminology have been proposed by a num­ which proved to have been still-born. I ber of authors, including Hesse ( 1924 ), hesitate to add to this discussion (and cer­ Thinemann ( 1926), Dudich ( 1932), Pavan tainly refuse to coin new words) but feel and Tomaselli ( 1944-58), and Christiansen it may be useful to outline my own at­ ( 1962) . Of these, only Dudich's division of tempts to gain some greater clarity in this the trogloxenes into the two categories of matter. It may well be that my thinking regular and accidental has gained any ac­ reflects the particular characteristics of the ceptance, although even this proposal is open to considerable doubt (see Barr 1963) Australian cave fauna or even my own idi­ and will be discussed below. The most use­ osyncrasies, and I stress that this paper is ful further contribution to this question was intended as a contribution to discussion, that of Jeanne! (1926) who adopted the rather than as a presentation of "'the new basic Schiner-Racovitza system, but also system for classification of cavernicoles." drew particular attention to the fauna of the The generally accepted, but variously in­ twilight or threshold zone. J eannel also terpreted, system which is in use today is developed a useful series of categories for based upon the work of Schiner ( 1854) the description of specific biotopes within who divided the fauna of the Adelsberg the cave environment and hence for the de­ Caves into troglobites, troglophiles, and tailed categorization of fauna within a par­ occasional cavernicoles. Racovitza ( 1907) ticular cave. However, this latter classifica­ proposed the term trogloxene to replace tion complements the Schiner-Racovitza Schiner's "'occasional cavernicoles", and system and in no way replaces it.

0 South Australian Museum, North Terrace, Ade­ An examination of the literature shows a laide, South Australia 5000. variety of definitions being used for each

VoLuME 33, NuMBER 1, JANUARY 1971 63 of the Schiner-Racovitza categories and an The Australian cave fauna is particularly even wider range of interpretations of these rich in species of this kind, and the signifi­ definitions. Accepting the idea of Vandel cance of this fact would be disguised if all ( 1965) that each of these categories must troglophiles were treated as a single cate­ inevitably be heterogeneous and the obvi­ gory. I have therefore adopted the practice ous fact that any system of categories must of sub-dividing the troglophiles into two inevitably be limited according to the avail­ sub-categories: able data, it seems clear that further clari­ First-level Troglophiles - those forms fication and agreement is much to be de­ living out their total life-cycle within sired. In striving towards some clarity for caves but also known from epigean my own purposes, I have tried not only to habitats. apply general classificatory principles, but Second-level Troglophiles-those to arrive at a series of classes which express forms living out their total life-cycle something of the relationship between the within caves, species and the cave ecosystem. known only from caves, but not exhibiting any modification to the The term and concept of troglobite is cave environment. perhaps the most clearly defined and under­ stood, even though formal definitions in The usefulness of this division, even not­ common use are often unclear. Thus, Hazle­ ing the extent to which its application may ton & Glennie ( 1962) suggest "'species liv­ be biased by lack of information, may be ing permanently in the dark zone, and not seen from a recent review of the Australian in the endogean or epigean domain.'' How­ cave fauna (Hamilton-Smith, 1967) which ever, their usage of the term corresponds lists 33 species as first-level and 30 species with the generally accepted definition, such as second-level. as that of Barr ( 1968), "'obligatory species The second source of confusion arises which are unable to survive except in caves from differing interpretations of the defini­ or similar hypogean habitats." tion. Most authors appear to assume that Greater confusion surrounds the troglo­ the definition intends to convey that many philes. This is, understandably, a much individuals of the species will spend their more heterogeneous category. Although gen­ total life within a cave, while other individ­ uals erally clearly enough defined, e.g. Barr will spend their total life outside of a cave. However, by contrast ( 1968), as .,facultative species which com­ with this usage, Richards ( 1968) applies the term troglo­ monly inhabit caves and complete their life phile to the New Zealand species Pallidop­ cycles there, but also occur in sheltered, lectron turneri and Gymnoplectron waitom­ cool, moist epigean microenvironments", it oensis ( Fam. Rhaphidophoridae), but at the is well known that many troglophiles are same time describes both species as fre­ only recorded from the dark zone of caves. quently leaving the cave for periods of two However, in the absence of regressive or or three hours, feeding, and then returning adaptive modification, these are by conven­ to the cave. This behaviour pattern is very tion treated as troglophiles rather than trog­ similar to, if not identical with, that of many lobites. Barr ( 1963) cites the example of trogloxenes and it would seem more appro­ Agonum (Rhadine) longicolle (Benedict), priate if these species were considered recorded only from Carlsbad Caverns and trogloxenic or primarily so. Their relation­ nearby caves, but possessing large faceted ship to the cave ecosystem is certainly that eyes and darker pigmentation than troglo­ of a trogloxene, as their behaviour would bitic members of the genus. He points out result in a considerable energy input to the that such species might reasonably be con­ system. Thus, Barr ( 1968) refers to cave sidered as potential troglobites, but suggests crickets as among the "most important trog­ that they must be currently considered as loxenes" in terms of energy input to the troglophiles. ecosystem. The treatment of the Rhaphi-

64 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN dophoridae as troglophiles by Rit:hards ( op. and summarizes this field, but it is clear cit. ) rests upon her distinctive and atypical from his summary that although this is a interpretation of the category definition complex ecological domain there is as yet which appears to embrace forms usually con­ no satisfactory overall conceptual framework sidered as "habitual trogloxenes." It there­ for its study. In terms of considering over­ fore seems confusing and inappropriate. all cave fauna it again seems preferable Dudich ( 1932) divided the trogloxenes not to force threshold dwellers into the into the two categories of "habitual" ( Pseu­ Schiner-Racovitza categories, but to clearly dotroglobionte) and "accidental" ( Tycho­ separate them. troglobionte). This pattern has been fol­ The above line of thought leads me to lowed by a number of authors, notably use seven categories which can even be Hazleton and Glennie ( 1962). Barr ( 1963) arranged into the customary binary key as suggested that the term "trogloxene" should shown below. not be applied to accidental species. I con­ 1. Obligate parasites .. PARASITES cur strongly with his suggestion, and reserve not obligate parasites ...... 2 the term for those species which frequent 2. Does not regularly inhabit caves at specific times within their life cycle, caves . ACCIDENTAL but at other times enter the epigean domain. Regularly inhabits caves . 3 Such species are a relatively major and con­ 3. Does not live in sistent source of energy input to the ecosys­ dark zone .. THRESHOLD tem. On the other hand, acddental species Lives in dark zone...... 4 are of much less significance in this respect 4. Does not spend total life and are certainly not consistent in their in cave . . . TROGLOXENE energy input. Spends total life in cave ...... 5 Obligate parasites constitute a further 5. Also known from FIRST-LEVEL minor problem. Hazleton & Glennie (1962), epigean habitats .... TROGLOPHILE while recognizing that these relate to the Only known from cave or cave ecosystem only through their host (at similar habitat ...... 6 least in the case of bat parasites), treat them 6. Not modified to SECOND-LEVEL as trogloxenes. I find it less confusing to cave environment ...... TROGLOPHILE treat these as a separate category, rather Modified to than trying to force them into the Schiner­ cave environment...... TROGLOBITE Racovitza system. In summary, the signific.:ant points sug­ The dwellers within the threshold zone gested are the division of troglophiles into present particular problems in many faunal two sub-categories, support of Barr (and lists, and, for instance, that of Hazleton and others) in separating accidental species from Glennie does not clearly distinguish thresh­ trogloxenes, and removal of threshold species old species from dark zone species. Studies and parasites from inclusion in the Schiner­ of this fauna are relatively few, although Racovitza categories. None are completely some attention has been paid to individual novel or original but it is hoped that their species, e.g. Graham ( 1966-68). Vandel integration may prove useful. Comment and ( 1965) lists a number of European studies discussion will be welcomed.

LITERATURE CITED Barr, T. C., Jr. 1963. Eeological classifi­ Christiansen, K. 1962. Proposition pour la cation of cavernicoles. Cave Notes, 5:9- classification des animaux cavernicoles. 12. Spelunca (4e), 2:76-78. 1968. Cave ecology and the evo­ Dudich, E. 1932. Biologic der aggtekeler lution of troglobites. Evolutionary Biol­ tropfsteinhohle "Baradla" in Ungarn. ogy, 2:35-102. Speleol. Monographien, 13:1-246.

VoLuME 33, NuMBER 1, ]ANUAJW 1971 65 Graham, R. E. 1966. Crane-Hies ( Tricho­ Jeanne!, R. 1926. Faune cavernicole de ceridae, Tipulidae) in California caves. France. Paris: Lechevalier. Cave Notes, 8:41-48. Pavan, M. and R. Tomaselli. 1944-56. Not 1967. The subterranean niche of seen, listed in Vandel, A. (1965 ) . Pseudometa biologica (Arachnida:Aranei­ Racovitza, E. 1907. Essai sur les prob­ dae). Caves and Karst, 9:17-22. lemes biospeleologiques. Biospeleologica I, Archiv. Zool. Exper. Gen., 36:371-488. 1968. Spatial biometrics of sub­ Richards, A. M. 1968. The cavernicolous terranean demes of Triphosa haesitata status of some species of Macropathinae ( Lepidoptera: Geometridae). Caves and ( Orthoptera: Rhaphidophoridae). J. Aust. Karst, 10:21-29. Ent. Soc., 7:87-89. Hamilton-Smith, E. 1967. The arthropoda Schiner, J. R. 1854. Fauna der Adelsber­ of Australian caves. J. Aust. Ent. Soc., ger Lueger und Magdalener-Grotte. In 6:103-118. Schmid!, A., Die grotten und hohlen von Hazleton, M. and E. A. Glennie. 1954. Cave Adelsberg, Lueg, Planina und Loos. Wien: fauna and flora. In C. H. D. Cullingford Braunmuller. ( ed. ), British Caving (2nd. edn. ), pp. Thienemann, A. 1926. Die binnengewas­ 347-395. London:Routledge& Kegan Paul. ser Mitteleuropas. Die Binnengewasser, I. Hesse, R. 1924. Tiergeographie auf oko­ Vandel, A. 1965. (Tr. Freeman, B. E.) logischer grundlage. Jena:Fischer. Biospeleology. Oxford:Pergamon Press.

Manuscript received by editors July 1970

66 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN Resume of the 1970 Short Course In Cave Biology

A Short Course in Cave Biology, prob­ 3. Planarians by Jerry Carpenter, Univer­ ably the first of its kind held in the United sity of Kentucky. Flatworms or planarians States, was conducted in conjunction with are widespread in cave waters of the eastern the annual convention of the National Spele­ and central U.S. and are represented by a ological Society at Pennsylvania State Uni­ number of troglobitic species. Several spe­ sity on August 21, 1970. The Short Course, cies were discussed and illustrated, and a day-long event composed of 10 lectures techniques for collection, preservation, and on cave organisms and their ecology, was study were stressed. sponsored by the Biology Section of the 4. Amphipods and I so pods by John R. N.S.S. The course was attended by 50 per­ Holsinger, Old Dominion University. Am­ sons, many of whom displayed a great deal phipods and isopods, two major orders of of interest in the proceedings and expressed higher crustaceans, are common in subter­ the desire for similar courses in the future. ranean waters of much of the United States. Because of the enthusiasm of the partici­ The similarities and differences of these pants and the rapidly expanding interest two groups of cavernicoles were emphasized in cave ecology, a sequel to the Penn. State and various aspects of their ecology were session will follow at the 1971 annual con­ discussed. vention at Virginia Polytechnic Institute. 5. Crayfish by Martha R. Cooper, Univer­ The 1971 session will consist of biological sity of Kentucky. Crayfish, another major excursions into nearby Greenbrier Caverns, group of crustaceans, are represented in West Virginia. The purpose of these excur­ cave waters of the eastern and central sions will be to point out some of the vari­ United States. The troglobitic forms are ous kinds of cave organisms in their natural predominantly restricted to northern Florida, environment. the Interior Low Plateau region, and the Ozark uplift. Most of the important cave A very brief summary of the talks given species were illustrated and discussed. at the 1970 Short Course follow in their 6. Caves as Islands and Archipelagoes by order of presentation: David Culver, University of Chicago. The 1. Introductory Remarks by John R. Hol­ community structure of caves was compared singer, Old Dominion University. A brief quantitatively with those of islands and history of the development of biospeleology archipelagoes. Caves, at least when consid­ in the United States was given, followed by ered from the standpoint of aquatic com­ a definition of some of the technical terms munities, tend to be more like archipelagoes currently used in the field. The importance than islands. of cave laboratories was also pointed out. 7. Ecology of Cave Entrances by Richard 2. Protozoans by Stephen Gittleson, Uni­ Graham, Upsala College. Although cave en­ versity of Kentucky. Some of the different trances are quickly by-passed by most cave types of protozoans recently discovered in biologists in their haste to get into the dark North American cave waters were discussed recesses, this zone is both significant and and illustrated. Although troglobitic (i.e., interesting. Composition and interaction of obligatory cavernicolous) protozoans are un­ species of the transitional zone between the known, a number of species that inhabit epigean and hypogean environment were subterranean waters exhibit rather unusual discussed. trends, such as frequent encystment and 8. Bats by Charles E. Mohr, Delaware extremely small size. Nature Education Center. Although bats

VoLU:\IE :33, NuMBER 1, jANUARY 1971 67 are only trogloxenes, ecologically speaking, 10. A1nphibians and Fishes by John E. these animals make up a significant fauna Cooper, University of Kentucky. Among the of many caves, and studies on their unique cold-blooded vertebrate animals, two groups behavior are being pursued by a number of -salamanders and fishes-are represented researchers. A number of the species as­ by troglobites. Although a number of sala­ sociated with North American caves were manders are found in caves, only a few are illustrated and attention was called to two exclusively troglobitic and all but one spe­ important books recently published on bats. cies are known from the United States. Both 9. Composition of Terrestrial Ecosystems cave-associated amphibians and fishes were by Thomas L. Poulson, University of Notre discussed and illustrated. Dame. The composition and interaction of species in the terrestrial ecosystems of vari­ ous caves were discussed and illustrated. -JOHN R. HoLSINGER, Many of the examples used in this presenta­ Department of Biology, tion were drawn from recent studies con­ Old Dominion University, ducted in the Flint Ridge and Mammoth Norfolk, Virginia 23508 Cave systems of central Kentucky.

Caves as Model Systems

REVIEW: "Observations on the ecology of cult to know how much credence to give caves" by Thomas C. Barr, Jr., American many of his generalizations, since references Naturalist, 101:475-492. 1967. to the literature are few and far between. For example, he states that "decomposition In this short paper Dr. Barr has attempted of the debris (in caves) takes place slowly to link what we know about cave communi­ over the course of several months or years." ties to the rapidly developing body of eco­ This may or may not be true (no references logical theory. The importance of the paper are given), but at least cave populations of can hardly be overestimated. It has been Gammarus minus ( Amphipoda) can feed widely read by ecologists, and together with on leaves and/or their microflora after the Poulson's work on amblyopsid fishes ( Poul­ leaves have been in the water only ten days son, 1963; Poulson and White, 1969) it (Culver, 1970b). However, many of the signals the emergence of biospeleology from rather general statements a long period of taxonomy. The paper has in this section are four sections: a general introduction to the probably backed up by observations recorded cave environment, biogeography of caves, in Barr's legendary field notebooks. It is various points of interest about the ecology unfortunate that more data were not pre­ of cave animals, and a model for speciation sented here to back up his contentions. in caves. In the section on biogeography, Barr con­ The general introduction to the cave en­ firms two predictions of the geographic vironment focuses on the contrast between theory of speciation. Since cave beetles are caves of the Appalachian Valley and the limited to subterranean dispersal: ( 1) there Mississippian Plateaus. In this section Barr are more species in the Appalachian Valley presents a unified, easily readable picture of where the limestone outcrops are discontin­ the cave environment. However, it is diffi- uous, and ( 2) the geographic ranges of the

68 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN species are smaller in the Appalachian Val­ of islands. Thus we need to compare val­ ley. Barr misinterprets evidence presented leys of different sizes to make a fair test in the investigation of Pseudosinella hirsuta of the area effect. Second, there are two ( Collembola: Entomobryidae) by Christian­ general kinds of equilibria of extinction and sen and Culver (1968). While it is true immigration (Janzen, 1968): an equilibrium that P. hirsuta represents several invasions concerning the number of individuals, and by a surface ancestor, each invasion and an equilibrium concerning the number of subsequent dispersal occurred within a sin­ species. The relationship of these two equil­ gle area of continuous limestone. Therefore, ibria to caves remains unelucidated. P. hirsuta confirms Barr's contention that Another source of confusion is Barr's im­ dispersal occurs via subterranean routes, but plied contention that the causes of high not the theory of geographic speciation. diversity in the tropics will be the same as While subterranean dispersal within lime­ the causes of high diversity in Mississippian stone outcrops seems to be the rule for ter­ Plateau caves. There is no reason to believe restrial cave animals, this is not the case that this is true. for many aquatic species. Holsinger ( 1969) Since the publication of Barr's paper, found that some amphipod species are able more sophisticated theories concerning inter­ to disperse via superficial subsurface water actions in complex communities have be­ as well as through subsurface water in lime­ come available (Levins, 1968), and these stone outcrops. make his discussion of the dynamics of I found the most interesting part of the higher species diversity out of date. This paper to be that relating the biology of cave is just as well, since he seems to argue that beetles to various ecological theories, par­ more species per se make more niches ticularly those that deal with aspects of available. diversity. Unfortunately, it is also the most One of Barr's more interesting findings is confusing part. It is clear, however, that that beetle abundance in some caves fits Barr's main thesis is that geologic features, what ecologists call a broken-stick distribu­ e.g. continuity of limestone outcrops, are tion (MacArthur, 1957). Such a distribu­ the dominant factors affecting diversity. The tion, however, does not indicate broad niche confusion arises when he tries to relate di­ overlap, as Barr suggests, but rather that versity of cave beetles to general theories of the niches of the species are contiguous diversity. There are two bodies of ecolog­ and non-overlapping. ical theory that might be applicable: theories about high diversity in the tropics, and In the concluding section, Barr discusses stochastic theories of island biogeography. the speciation process in troglobites. This It is unclear which of these Barr considers theory is greatly expanded in his paper on more useful, and in fact he does not explic­ evolution of troglobites ( 1968), but my itly distinguish the two. Although he pre­ major objection to the section as presented sents an analogy between tropical and here is that it is too vague and general. temperate environments and Mississippian The status of troglophile populations with Plateau caves and Appalachian Valley caves, respect to epigean or surface populations is it seems to me that a better analogy than complex and poorly understood. (Christian­ tropical and temperate environments is one sen and Culver, 1969; Holsinger and Culver, with islands or groups of islands. Caves in in press ) , yet Barr's discussion here and in one karst valley are highly connected, and his paper on troglobite evolution does not therefore immigration rates are very high. aid in the analysis of such transitional popu­ For example, in the Greenbrier Valley of lations. What is needed is a quantitative, West Virginia, I have found (Culver, 1970a) rigorous theory of the process of cave that there was no area effect, i.e. an increase adaptation. Since Barr knows the literature in species number with increased area, which of population genetics well, he is the obvi­ is one of the most general characteristics ous person to develop such a theory.

VoLUME 33, Nu:MBER 1, }ANUARY 1971 69 LITERATURE CITED Barr, T. C. 1968. Cave ecology and the Holsinger, J. R. and D. C. Culver. In press. evolution of troglobites. Evolutionary Morphological variation in Gammarus Bioi., 2:35-102. minus Say ( Amphipoda, Gammaridae) Christiansen, K. A. and D. C. Culver. 1968. with emphasis on subterranean forms. Geographical variation and evolution in Postilla. Pseudosinella hirsuta. Evolution, 22:237- Janzen, D. H. 1968. Host plants as is­ 2.5.5. lands in evolutionary and ecological time. 1969. Geographical variation and Amer. Natur., 102:592-595. evolution in Pseudosinella violenta ( Fol­ Levine, R. 1968. Evolution in changing som). Evolution, 23:602-621. environments. Princeton Univ. Press, Culver, D. C. 1970a. Analysis of simple Princeton, N. J. 120 pp. cave communities. I. Caves as islands. MacArthur, R. H. 1957. On the relative Evolution, 24:463-474. abundance of bird species. Proc. N atl. 1970b. Analysis of simple cave Acad. Sci., 43:293-295. communities. Ph.D. Thesis. Yale Uni­ Poulson, T. L. 1963. Cave adaptation in versity. amblyopsid fishes. Amer. Midi. Natur., Holsinger, J. R. 1969. Biogeography of 70:257-290. the fresh-water amphipod crustaceans Poulson, T. L. and W. B. White. 1969. ( Gammaridae) of the central and south­ The cave environment. Science, 165:971- ern Appalachians. In P. C. Holt ( ed.), 981. The distributional history of the biota of -David C. Culver, the southern Appalachians. Part 1: the Department of Biology, invertebrates, pp. 19-50. Virginia Poly­ University of Chicago, technic Institute Press, Blacksburg, Vir­ Chicago, Illinois 60637 ginia.

REviEw: .. Cave Ecology and the Evolution major picture developed is both clear and of Troglobites" by Thomas C. Barr, Jr. accurate. Evolutionary Biology, Vol. 2:39-102, 1968. The most important section of the article deals with the evolution of troglobites. An This excellent work is required reading introduction and history of biospeleology is for every student of cave biology. It serves followed by a summary of the where and equally well as an introduction to cave ani­ what of cave animals. After an outline of mals for the student of evolution and a the nature of physical conditions in caves, source for biospeleologists. The analysis of there is a more extended treatment of food the nature and origin of the cave biomes and food webs. This is followed by a brief is the best short summary available. The discussion of the present evidence for cir­ main body of the work is well-organized, cadian rhythms in cave animals and the and the topic headings lend themselves to effect of geology on troglobite distribution easy reference. The section on the cave in the East Central U. S. An extended ecosystem is largely oriented around nearctic discussion of the question of "regressive" systems but presents a clear view of most evolution is followed by an analysis of the of the information about such systems in problems of speciation and adaptation in 1968. One can quarrel with a few minor caves in general. The interesting section points (such as the statement that the on circadian rhythms points up the incon­ energy contribution of chemosynthetic auto­ clusive nature of presently available data trophs is probably negligible); however, the and sharply poses a number of critical ex-

70 THE NATIONAL SPELEOLOGICAL SociETY BuLLETIN periments. ~ The initial historical summary the genotype associated with the new di­ of the question of "regressive evolution" in rection of evolution furnished by the cave cave animals clearly exposes much of the habitat is supported by a considerable body barren ground this idea has traversed. Read­ of evidence. The idea that this reconstruc­ ing the later discussion of speciation and tional phase can only occur after the extinc­ adaptation points out the erroneous nature tion of epigean ancestral forms is a more of the term "regressive" as applied to trog­ doubtful hypothesis. A number of recent lobitic change. A far more apt term would works have shown that a high degree of be reconstructional evolution. The basic population isolation can exist without any idea clearly presented by Barr that cavern­ apparent physical barrier to interbreeding. icoles go through a period of isolation It should then be possible for the recon­ followed by a period of reconstruction of struction phase to occur locally even in the presence of the epigean ancestor. The fact o Editor's note: There have been two experi­ remains that the ancestral forms are no mental studies on circadian rhythms published since Barr's paper was written. A study on cave longer extant in areas where most troglo­ crayfish [T. C. Jegla and T. L. Poulson, Evidence bites occur. Whether this extinction is a of Circadian Rhythms in a Cave Crayfish. J. Exp. Zool. 168:273-282 ( 1968)] shows that 3 of 7 necessary prerequisite of troglobitization, or individuals had statistically signficant circadian rhythms of metabolic rate and/or activity and 5 of a by product of the stability of the cave 7 showed some rhythmicity and other criteria ex­ environment, is a secondary matter. pected of circadian clocks, e.g. resetting after change in a light-dark cycle. A parallel study on While some students of cave biology may amblyopsid cave fish, of which some preliminary results have been published [T. L. Poulson and disagree with the emphasis on Mayrsian T. C. Jegla, Circadian Rhythms in Cave Animals. speciation, or other points in this article, it Proc. IV Internatl. Congr. Speleol. in Yugoslavia, 4-5:193-195 ( 1969)], shows that a troglophilic and is on the whole clear, concise, and useful. three troglobitic species all show clear evidence of circadian metabolic rates but that activity rhythms and effects of light-dark cycles on activity patterns -Kenneth Christiansen, are clear cut only in the troglophile; they are lost Dept. of Biology, in the troglobite isolated in caves for the longest evolutionary time and show an intermediate state, Grinnell College, like that in the cave crayfish, in the troglobite most recently isolated in caves. Grinnell, Iowa 50112

VoLUME 33, NuMBER 1, }ANUARY 1971 71