All 8X9Xe %&* VJM' number 12 September 1986 EDITORIAL STAFF Eloise F. Potter, Acting Editor Eloise F. Potter, Managing Editor John B. Funderburg, Editor-in-Chief Board James W. Hardin Department of Botany N.C State University David S. Lee Curator of Birds N. C. State Museum William M. Palmer Curator of Lower Vertebrates N. C State Museum Rowland M. Shelley Curator of Invertebrates N. C. State Museum Brimleyana, the Journal of the North Carolina State Museum of Natural His- tory, will appear at irregular intervals in consecutively numbered issues. Con- tents will emphasize zoology of the southeastern United States, especially North Carolina and adjacent areas. Geographic coverage will be limited to Alabama, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, Virginia, and West Virginia. Subject matter will focus on taxonomy and systematics, ecology, zoo- geography, evolution, and behavior. Subdiscipline areas will include general invertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and paleontology. Papers will stress the results of original empirical field studies, but synthesizing reviews and papers of significant historical interest to southeastern zoology will be included. Suitability of manuscripts will be determined by the Editor, and where neces- sary, the Editorial Board. Appropriate specialists will review each manuscript judged suitable, and final acceptability will be determined by the Editor. Address manuscripts and all correspondence (except that relating to subscrip- tions and exchange) to Editor, Brimleyana, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. In citations please use the full name — Brimleyana. North Carolina State Museum of Natural History North Carolina Department of Agriculture James A. Graham, Commissioner CODN BRIMD 7 ISSN 0193-4406 Notes on Turtle Egg Predation by Lampropeltis getulus (Linnaeus) (Reptilia: Colubridae) on the Savannah River Plant, South Carolina James L. Knight Savannah River Ecology Laboratory, Drawer E, Aiken, South Carolina 29801 AND Raymond K. Loraine l Museum of Natural History, University of Kansas, Lawrence, Kansas 66045 ABSTRACT. — Observations on turtle egg predation by the colubrid snake Lampropeltis getulus on the Savannah River Plant, South Carolina, indicate that, during the turtle nesting season, some king- snakes apparently search out and consume the contents of multiple turtle nests. This seems especially true for nests of kinosternid turtles. Future studies of predators on turtle nests within the range of L. getu- lus should take that taxon into account as a potentially prominent predator. Eggs of Sternotherus odoratus may hatch even after passing through the digestive tract of L. getulus. Kingsnakes of the colubrid genus Lampropeltis have long been known to feed on a wide variety of vertebrate prey (for a review, see Wright and Wright 1957). Of particular interest is the tendency of these snakes to consume the eggs of other reptiles, especially turtles. Brown (1979) listed two turtle eggs from two Lampropeltis getulus, and Hamil- ton and Pollack (1956) listed prey items found in L. getulus from Fort Benning, Georgia, including the eggs of lizards and snakes. Wright and Bishop (1915) reported the eggs of Pseudemys floridana and Kinoster- non spp. from stomachs of Okefenokee swamp L. getulus and observed that ". . . so addicted are they [L. getulus\ to this egg diet, that the natives consider that it is a common happening to find the snake await- ing the egg deposition." They also said that, aside from "the Florida bear, there is no form in the swamp which eats turtle's eggs in such quantity as the kingsnake. It will take a whole nest of eggs at one time, as many as 14 being found in the stomach of one snake." Ernst and Barbour (1972) cite numerous turtle species whose young are eaten by various species of snakes, but relatively few turtles whose eggs are eaten. Present address: Department of Zoology, University of South Florida, Tampa, Florida 33620. Brimleyana No. 12:1-4, September 1986 2 James L. Knight and Raymond K. Loraine Recent collections and observations on two specimens of L. getulus from the Savannah River Plant (SRP), in Aiken, Barnwell and Allen- dale counties, South Carolina, shed additional light on turtle egg-eating propensities of L. getulus and indicate that at least a small subset of the population of L. getulus on the SRP may search out nesting turtles and wait for them to lay their eggs, as suggested by Wright and Bishop (1915). On 27 May 1984, one of us (RKL) removed a L. getulus from a funnel snake trap along a drift fence near the northeast side of Ellenton Bay, a Carolina bay in the Aiken County portion of the SRP. This snake, a female with a snout-vent length (SVL) of 1118 mm, regurgi- tated 9 turtle eggs (6 ruptured, 3 intact) that, based on shape, appeared to represent several different turtle taxa. One hard-shelled egg was immediately referable to the family Kinosteridae; one was light-colored and round, apparently Chelydra; and the remaining 7 could have been assignable to any of several species of emydid turtles. On 22 June 1984, one of us (JLK) collected a female L. getulus (1257 mm SVL) along a sandy road that courses parallel to, and aver- ages about 50 m from, the edge of the Savannah River Swamp, ca. 2 km east-southeast of the mouth of Pen Branch Creek, in Barnwell County. The collector had stopped to capture a Terrapene Carolina that was in the process of excavating a nest chamber (she later laid 3 eggs in the lab). When first observed, the snake was less than a meter from the turtle, with its head and neck elevated about 10 to 12 cm off the ground and directed toward the turtle. The snake was captured, placed in a collecting bag and, upon returning to the lab, was found to have regur- gitated 4 hard-shelled eggs (2 intact, 1 damaged, 1 crushed). The snake was caged by itself and, after 3 days, defecated parts of, minimally, an additional 13 kinosternid eggs, 3 of them unbroken. Three species of kinosternid turtles have been collected at SRP: Sternotherus odoratus and Kinosternon subrubrum (Gibbons and Pat- terson 1978), and Kinosternon bauri (Lamb 1983). The eggs are most likely of S. odoratus and /or K. subrubrum, as K. bauri is comparatively rare on the SRP, the northernmost record of occurrence for the species. Unfortunately, measurements of the intact eggs yielded no information as to their identity, for all three species lay eggs of approximately the same size. Of particular interest was the number of turtle eggs present in the second snake. Gibbons (1983), discussing SRP K. subrubrum, gave a mean of 3.03 eggs/clutch, range 1-5 (N = 161). Tinkle (1961) divided a sample of adult female S. odoratus into two arbitrary size classes, the smaller exhibiting an average clutch size of 2.0 eggs and the larger aver- Kingsnake Predation on Turtle Eggs 3 aging 3.2 eggs/ clutch. If the snake located and devoured "average" clutches of K. subrubrum, then at least five or six different nests had been preyed upon, all within a fairly short period. If the same scenario is applied to "average" clutches of S. odoratus, the snake may have preyed on five to nine nests. Given the circumstances of its capture, it seems highly probable that the snake would have taken the contents of the T. Carolina nest as well. Interestingly, the three intact eggs that passed through the digestive system of the snake and were then defe- cated were incubated in the lab and hatched after approximately 50 days, yielding three S. odoratus. Imler (1945) mentioned a bullsnake, Pituophis melanoleucus sayi, with an "egg appetite to the extent that it will not eat anything else," and Legler (1960), citing a conversation with the late E. H. Taylor, men- tioned a bullsnake that "swallowed an entire clutch of newly laid eggs [of Terrapene ornata] before the female turtle could cover the nest." Per- haps some individual L. getulus behave the same way in nature. Legler (1960) stated that nest predation may have a greater effect on popula- tions than predation on hatchlings, juveniles, and adults. Our data sug- gest that L. getulus, particularly those in areas of extensive turtle nest- ing, as along the margin of the Savannah River Swamp, might contribute more than slightly to turtle egg predation totals. Any future studies of predation on turtle eggs should take this predator into account. ACKNOWLEDGMENTS.— Thanks go to R. A. Seigel, J. Iverson, and S. Novak for commenting on earlier drafts of this paper; R. A. Estes for field assistance; S. J. Morreale for incubating and hatching the S. odoratus eggs; and J. W. Gibbons for the opportunity to collect and report these observations. Manuscript preparation was supported by contract DE-AC09-76SR00819 between the U.S. Department of Energy and the University of Georgia's Savannah River Ecology Laboratory. LITERATURE CITED Brown, E. E. 1979. Some snake food records from the Carolinas. Brimleyana 1:113-124. Ernst, Carl H., and R. W. Barbour. 1972. Turtles of the United States. Univ. Kentucky Press, Lexington. Gibbons, J. Whitfield. 1983. Reproductive characteristics and ecology of the mud turtle, Kinosternon subrubrum (Lacepede). Herpetologica 39(3): 254-271. , and K. K. Patterson. 1978. The reptiles and amphibians of the Savannah River Plant. National Environmental Research Park 2:1-24. 4 James L. Knight and Raymond K. Loraine Hamilton, W. J., and J. A. Pollack. 1956. The food of some colubrid snakes from Fort Benning, Georgia. Ecology 37(3):5 19-526. Imler, R. H. 1945. Bullsnakes and their control on a Nebraska wildlife refuge. J. Wildl. Manage. 9(4):265-273. Lamb, Trip. 1983. The striped mud turtle {Kinosternon bauri) in South Caro- lina, a confirmation through multivariate character analysis. Herpetologica 39(4):383-390. Legler, John M. 1960. Natural history of the ornate box turtle, Terrapene ornata ornata Agassiz. Univ. Kans. Publ. Mus. Nat. Hist. 11(10): 527-669. Tinkle, Donald W. 1961. Geographic variation in reproduction, size, sex ratio and maturity of Sternothaerus odoratus (Testudinata: Chelydridae). Ecol- ogy 42(l):68-76. Wright, Albert H., and S. C. Bishop. 1915. II. Snakes. Pages 139-192 in A biological reconaissance of the Okefinokee swamp in Georgia: The reptiles. Proc. Acad. Nat. Sci. Phila.: 107-192. , and A. A. Wright. 1957. Handbook of Snakes of the United States and Canada. 2 vols. Comstock Publ. Assoc, Ithaca, N.Y. Accepted 30 July 1985 Observations on the Social Behavior of the Southern Cricket Frog, Acris gryllus (Anura: Hylidae) Don C. Forester Department of Biological Sciences I Institute of Animal Behavior, Towson State University, Towson, Maryland 21204 AND Richard Daniel Division of Biological Sciences, University of Missouri, Columbia, Missouri 65201 ABSTRACT. — Southern Cricket Frogs are prolonged breeders. During the reproductive season, males occupy calling territories from which they advertise for females. Mean territory size was 0.56 m2 (0.03-1.36 m2), and mean nightly movement by territorial males was 52 cm (0-205 cm). Territory size was not correlated with the number of days spent calling or with mating success. Observations on courtship behavior are presented. Anuran species are categorized as either explosive or prolonged breeders (Wells 1977a). For species composing the former group, males and females arrive synchronously at the reproductive site. In many such species, males actively search out females, and mate discrimination by the female may be limited by male assertiveness. Explosive breeders are stimulated by heavy rainfall and breed for only a fews days afterwards. Prolonged breeders often partition the reproductive site into defended calling stations. Males advertise their position by persistent vocaliza- tion, and the arrival of receptive females is typically asynchronous. In species of this type, breeding is less dependent on seasonal precipitation, and reproductive activity may continue for months. Studies on the reproductive behavior of anuran amphibians (par- ticularly prolonged breeding species) have greatly increased during the past 15 years (for a review see Wells 1977a,b; Arak 1983). The purpose of this investigation is to quantify the breeding and courtship behavior of the Southern Cricket Frog, Acris gryllus, a small, terrestrial hylid indigenous to the southeastern United States (Neill 1950). During early spring and summer, males aggregate around pools and call. Chorusing may persist throughout the summer and calling males have been reported as late as early October (Wright and Wright 1949). Females appear to arrive at breeding pools asynchronously throughout the spring and Brimleyana No. 12:5-11, September 1986 6 Don C. Forester and Richard Daniel summer, with peak oviposition from late April through June (Mecham 1964), but egg clutches have been reported during early fall (Wright and Wright 1949). Despite the fact that this species is among the most common anurans within its geographic range, little is known about its reproduc- tive biology and social structure. Our attention shall focus on social interactions between males by testing the following hypotheses: (1) call- ing males occupy a territory, (2) there is a correlation between the size of a calling territory and the number of nights a male is observed at the pond, and (3) there is a correlation between territory size and mating success. STUDY SITE This study was conducted during June and July of 1975. The study site was a complex of three small sand pits situated in a mixed pine/ de- ciduous flatwoods in Bryan County, Georgia. In most years the pits collect rain, and during the spring and summer are active reproductive sites for numerous amphibian species, including: the Southern Toad, Bufo terrestris; the Oak Toad, Bufo quercicus; the Eastern Narrow- mouthed Toad, Gastrophryne carolinensis; the Squirrel Treefrog, Hyla squirella; the Pine Woods Treefrog, Hyla femoralis; the Barking Tree- frog, Hyla gratiosa; the Southern Cricket Frog, Acris gryllus; the Little Grass Frog, Limnaoedus ocularis', the Southern Leopard Frog, Rana sphenocephala; the Crawfish Frog, Rana areolata; the Bullfrog, Rana catesbeiana; the Carpenter Frog, Rana virgatipes', the Mole Salamander, Amby stoma talpoideum; the Red-spotted Newt, Notophthalmus viri- descens; and the Striped Newt, Notophthalmus perstriatus. Our study was confined to a small (D = 3.5 m) pool with gently sloping banks and a firm bottom. The margin of the pond was covered with patches of low, dense grass. This vegetation was cover for 8 to 12 calling male A. gryllus, and sparse enough to permit observation of the males with minimal disturbance. METHODS Individual Recognition. — Male cricket frogs may be distinguished individually based on their dorsal pattern (Bayless 1969). All males observed during this study had their dorsal patterns diagramed for ref- erence. Because the breeding congress was small and never included more than eight males on any given night, individuals were easily recognized. Calling Stations. — Male Acris call from land (Wright and Wright 1949), and in the present study were always within 1 m of the shoreline. Social Behavior of Acris gryllus 1 Males were located by entering the pond at a given point each night and searching the periphery from the water. When a male was located (usu- ally by phonotaxis), a small marker was inserted into the substrate beside him. The markers were constructed from wooden dowels (D = 3 mm, L = 120 mm) to which a piece of white, waterproof tape had been attached. The identification number of the male and the observation date were printed on the tape with India ink. Calling males apparently were not disturbed by these activities. Site Fidelity by Calling Males. — Each time the position of a calling male was marked, we recorded its spatial relationship (directional angle and distance in cm) to the most recently placed marker and to the origi- nal observation point. These measurements enabled us to plot the terri- tories of individual males on graph paper. A Leitz planimeter (Model 3651-30) was used to calculate the area within each territory. Area values were based on an average of five separate measurements. Statistical Analysis. — Spearman's rank correlation procedure (Zar 1974) was used to test for correlations between territory size and the number of nights a male was observed at the pond, and between terri- tory size and mating success. The Spearman's rank procedure is a non- parametric test developed to process data obtained from a bivariate population that violates normalcy. Operational Sex Ratio. — We calculated the operational sex ratio (OSR) for the males and females observed during this study. The OSR is defined as the average ratio of fertilizable females to sexually active males at any given time (Emlen and Oring 1977). The OSR may or may not reflect the overall sex ratio of the species, particularly for prolonged breeders in which females arrive asynchronously at the reproductive site. RESULTS AND DISCUSSION Site Fidelity and Size of Territory. — Individual males moved an average of 52 cm (0 to 205 cm) between nights. Table 1 compares the mean nightly movement of each male. Nine of eleven males were observed on enough nights to facilitate calculation of their calling terri- tories. Mean territory size was 0.562 m2 (0.028-1.362 m2). We believe that this restricted movement and site fidelity warrant acceptance of our first hypothesis, that calling Acris gryllus males are territorial. We must reject our second and third hypotheses. There was no significant corre- lation between the size of a territory and the number of nights a male was observed at the pond (two-tailed Spearman's Rho, r = -5.521, P > 0.05). Neither was there a correlation between territory size and mating success (two-tailed Spearman's Rho, r = 0. 187, P > 0.05) (see Table 2). 8 Don C. Forester and Richard Daniel In our study, individuals appeared evenly spaced around the mar- gin of the pond. On only one occasion was a calling male seen invading the calling territory of a conspecific. This occurred on 16 June, when, after 1 night at the pond, Male 10 moved into the adjacent territory of Male 1. The resident male moved 1.3 m counterclockwise and continued to call for 3 nights before disappearing from the pond. The only other example of an extensive spatial shift occurred on 13 June, when Male 5 moved 2 m counterclockwise in response to rising water, which inun- dated his original calling site. This shift did not cause a change in the calling territory of the adjacent male (Male 4), and Male 5 remained at his site for an additional 13 days before leaving the pond on 26 June. Although we did not quantify intermale distance, such data are available for the species. Turner (1960) performed nearest neighbor analysis on a Louisiana population in December and April, and reported mean isolation distances of 1.94 m and 1.71 m, respectively. Behavioral Observations. — Five of the 11 males monitored during our study (observations were made on 18 nights during a 37-night period) were observed to amplex a female. Male 4 successfully amplexed two females over a 4-night span. Five of the six amplecting pairs were observed within a 5-night period during mid-June. It is probable that additional matings occurred but went undetected, for we were unable to visit the pond every night and frequently departed while some males were still advertising. The operational sex ratio at our study pond was skewed in favor of the males (5.6:0.3). However, it is likely that we underestimated the number of females present at the pond, and as a consequence we con- sider our OSR value conservative. On three occasions during the course of our study, we had the opportunity to observe male-female interactions leading to amplexus. A summary of each follows. (1) 13 June 1975. Male 4 was calling from his territory. With the exception of his pulsating vocal sac, he was hidden from direct view by dense grass. As we watched, a large female hopped into the circle of light. She appeared to be searching for the source of the sound. Her behavior included short, circling hops coupled with periodic cocking of her head from side to side. As the male continued to call, the female became increasingly active, crawling on the grass tussock and actually passing directly over the male on several occasions. Although the female circled eight times, the diameter of the circles never exceeded 8 cm. This sequence occupied just under 5 minutes and terminated when the male quickly emerged and amplexed the female. She neither resisted the male nor initiated contact with him. Social Behavior of A cris gryllus 9 Table 1. Linear movement of Acris gryllus males between consecutive nightly observations. Male Number of nights Mean Range No. 1 observed (cm) 81.00 (cm) 7 0-160 2 8 8.00 0-29 3 9 54.80 0-168 4 5 38.20 0-50 5 6 72.33 30-164 6 2 10.00 10-10 7 4 119.50 70-205 8 - - - 9 6 47.00 16-105 10 2 14.50 14-15 11 3 29.33 12-46 Table 2. Territory size, duration of calling, and mating success in a small breeding congress of Acris gryllus. No. nights observed = number of nights on which a male's calling position was marked. Days in resi- dence = the span over which the male was known to be at the pond. Male Territory size No. nights Days in No. matings No. (m2) observed 9 residence 10+ observed 1 0.502 0 2 0.127 12 13+ 1 3 1.373 16 37+ 0 4 0.180 7 8+ 2 5 1.362 11 18+ 0 6 - 3 3 0 7 0.673 6 8 1 8 0.288 12 33+ 1 9 - 2 4 0 10 0.028 6 8 0 11 0.545 7 23+ T= 14.06 1 ~x ■ = 0.562 x= 8.27 10 Don C. Forester and Richard Daniel (2) 14 June 1975. Male 7 was calling in an open spot between sev- eral clumps of grass. He lowered the pulse rate of his call and became active shortly before a female became visible. As the female approached, the male began to hop in tight circles (D = 4 cm). While moving, he continued to call. After 2.5 minutes, the male ceased calling and became stationary. Immediately the female approached to within 1 cm of the male's left side, and he quickly turned and faced her, snout to snout. After a 15-second pause, the male moved behind and amplexed the female. (3) 1 1 July 1975. Male 1 1 was calling while a female sat 3 cm away, facing the opposite direction. They remained motionless for approxi- mately 5 minutes. Suddenly the female began what we describe as a "quiver-hop" behavior, which involved quick, nervous movement of the forelimbs and elevation of the body 1 to 2 mm in a vertical position. After the female had exhibited this behavior twice in rapid succession, the male turned, moved quickly behind the female, and initiated amplexus. Calling male cricket frogs formed duets, trios, quartets, and occa- sionally quintets. The significance of this call synchrony to Acris gryllus was not tested, but similar behavior is reported to be important during mate selection by other hylids. In a study of the Pacific Treefrog, Hyla regilla, females preferred the designated bout leader during call discrim- ination trials involving a single male quarteting with itself (Whitney and Krebs 1975). The authors concluded that bout leadership must some- how imply greater fitness to a responding female. We doubt that bout leadership is indicative of male fitness in A. gryllus, for two reasons: (1) bout leadership often changed during the course of an evening, and (2) bout leadership frequently changed from one night to the next. We sug- gest, as an alternative hypothesis, that antiphonal calling may enhance the fitness of the participating males by reducing broadcast interference. This role has been documented for the Spring Peeper, Hyla crucifer, a prolonged breeder of similar size and habits (Forester and Harrison, unpubl. ms.). Among hylids, satellite behavior and sexual parasitism by noncall- ing males has been well documented (Perrill et al. 1978, 1982). To employ this behavioral strategy a noncalling male positions himself near a calling male and attempts to intercept females responding to the caller. Often, calling males respond agonistically to satellites as well as to other conspecific males that violate their calling territory. During our study, in more than 70 hours of observation, we observed neither satel- lite behavior nor agonistic encounters between males. Our failure to document social interactions between males is more likely a reflection of Social Behavior of Acris gryllus 1 1 low male density at our study pond, since both behaviors have been observed in dense populations of the closely related congener, Acris crepitans, in Indiana (S. A. Perrill, pers. comm.). ACKNOWLEDGMENTS.— S. Simon and C. Boake provided assistance and companionship in the field. This project was funded, in part, by NSF Grant BNS 73 00795 to H. C. Gerhardt. LITERATURE CITED Arak, Anthony. 1983. Male-male competition and mate choice in anuran amphibians. Pages 181-210 in P. Bateson (editor). Mate Choice. Cambridge Univ. Press, New York. Bayless, Laurence E. 1969. Ecological divergence and distribution of sympatric Acris populations (Anura: Hylidae). Herpetologica 25(3): 18 1-1 87. Emlen, Stephen T., and L. W. Oring. 1977. Ecology, sexual selection and the evolution of mating systems. Science 197:215-223. Mecham, John S. 1964. Ecological and genetic relationships of the two cricket frogs, genus Acris, in Alabama. Herpetologica 20(1):84-91. Neill, W. T. 1950. Taxonomy, nomenclature, and the distribution of southeast- ern cricket frogs, genus Acris. Am. Midi. Nat. 43(1): 152- 156. Perrill, Stephen A., H. C. Gerhardt, and R. Daniel. 1978. Sexual parasitism in the green tree frog (Hyla cinerea). Science 200: 1 179-1 180. , , and 1982. Mating strategy shifts in male green treefrogs {Hyla cinerea): an experimental study. Anim. Behav. 31:43-48. Turner, Frederick B. 1960. Size and dispersion of a Louisiana population of the cricket frog, ,4cm gryllus. Ecology 4 1(2): 258-268. Wells, Kentwood D. 1977a. The social behaviour of anuran amphibians. Anim. Behav. 25:666-693. . 1977b. The courtship of frogs. Pages 233-262 in D. H. Taylor and S. I. Guttman (editors). The Reproductive Biology of Amphibians. Plenum Press, New York. Whitney, Carl L., and J. R. Krebs. 1975. Mate selection in Pacific tree frogs. Nature 255:325-326. Wright, Albert H., and A. A. Wright. 1949. Handbook of Frogs and Toads. 3rd ed. Comstock Publ. Co., Inc., Ithaca. Zar, J. H. 1974. Biostatistical Analysis. Prentice-Hall, Englewood Cliffs, N.J. Accepted 28 June 1985 12 THE SEASIDE SPARROW, ITS BIOLOGY AND MANAGEMENT Edited by Thomas L. Quay, John B. Funderburg, Jr., David S. Lee, Eloise F. Potter, and Chandler S. Robbins The proceedings of a symposium held at Raleigh, North Carolina, in October 1981, this book presents the keynote address of F. Eugene Hester, Deputy Director of the U. S. Fish and Wildlife Service, a bibli- ography of publications on the Seaside Sparrow, and 16 major papers on the species. Authors include Arthur W. Cooper, Oliver L. Austin, Jr., Herbert W. Kale II, William Post, Harold W. Werner, Glen E. Wool- fenden, Mary Victoria McDonald, Jon S. Greenlaw, Michael F. Delany, James A. Mosher, Thomas L. Merriam, James A. Kushlan, Oron L. Bass, Jr., Dale L. Taylor, Thomas A. Webber, and George F. Gee. A full-color frontispiece by John Henry Dick illustrates the nine races of the Seaside Sparrow, and a recording prepared by J. W. Hardy supple- ments two papers on vocalizations. "The Seaside Sparrow, with its extensive but exceedingly narrow breeding range in the coastal salt marshes, is a fascinating species. All the authors emphasize that the salt marsh habitat is at peril. . . . The collection is well worth reading." — George A. Hall, Wilson Bulletin. 1983 174 pages Softbound Price: $15, postpaid. North Carolina residents add 4'/2% sales tax. Please make checks payable in U. S. currency to NCDA Museum Extension Fund. Send to SEASIDE SPARROW, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 2761 1. Core Temperatures of Non-nesting Western Atlantic Seabirds Steven P. Platania ', Gilbert S. Grant 2, and David S. Lee 3 North Carolina State Museum of Natural History, P.O. Box 27647, Raleigh, North Carolina 27611 . ABSTRACT. — Core body temperatures of 23 species of birds col- lected off the North Carolina coast did not differ with sex, weight, time of day, or season. Within the orders Procellariiformes and Cha- radriiformes, there seems to be no correlation of temperature with mass. Temperature data on injured birds are similar to those of ones recently killed. Results of this study compared favorably with those obtained by other researchers and indicate no significant differences between body temperatures of foraging and non-incubating procellarii- form birds at the nesting colonies. Temperature differences between birds taken at sea and those studied at nesting sites amount to about 1 °C and are best attributed to the activity state of the birds. Little uniform information is available on deep-body temperatures of seabirds away from nesting colonies. Comparing thermal information collected by different investigators, using dissimilar methods and sam- pling variable locations within the body, presents interpretive difficulties. The opportunity to gather temperatures from a variety of species, using uniform methods and equipment, presented itself during a long-range study into the occurrence, seasonal distribution, and food habits of sea- birds off the North Carolina coast (see Lee and Booth 1979). This paper is the first extensive report of core temperatures in actively foraging seabirds. It complements the works of others who obtained most of their information from nesting colonies, and for the most part substan- tiates their findings and speculations. MATERIALS AND METHODS Information was obtained between 1977 and 1982, primarily during spring, summer, and fall. Specimens were shot from boats traveling from 30 to 60 km off North Carolina's Outer Banks. Birds were then netted from the water and a thermistor probe (#418), feeding into a calibrated telethermometer (Yellow Springs Instruments), was inserted 1 Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, Colorado 80523. 2 Route 2, Box 431, Sneads Ferry, North Carolina 28460. 3 Direct requests for reprints to Lee. Brimleyana No. 12:13-18, September 1 986 13 14 Steven P. Platania, Gilbert S. Grant, David S. Lee through the abdominal wall near the caudal part of the sternum deep into the viscera. The maximum time between downing of the bird and the insertion of the thermistor probe was 2 minutes. Body temperature (T^) recordings stabilized within a maximum of 30 seconds of probe insertion. In order to determine if stress and shock affected core body temperature, readings were taken from any still-living birds before death and within 1 to 3 minutes of being shot. We also monitored the rate of cooling of six specimens for 20 minutes after death. Birds were later frozen in sealed plastic bags. After thawing in the laboratory, each bird was weighed to the nearest 0.1 g and sexed while being prepared for use in other studies. Level of significance is 0.05 for correlation coefficients of regressions and sample differences (using Student's /-test). Data are presented as mean + 1 standard deviation. RESULTS Table 1 presents deep body temperatures and body mass of 23 spe- cies of seabirds representing 4 orders, 14 genera, and 250 individuals. Mean T^ of male and female seabirds (Table 2) were not significantly different. In Audubon's Shearwater, Puffinus Iherminieri, and Cory's Shearwater, Calonectris diomedea, the only species with a field collect- ing base spanning 6 to 9 hours, Tu did not vary with time of day. How- ever, we made no night collections. In both of these shearwaters, as well as in the Greater Shearwater, Puffinus gravis, Tu did not correlate with time of year. These three species were collected during the longest calendar sequences (April-November). Additionally, intraspecific regres- sions of body mass versus Tu were not significant. Cooling curves were obtained on six birds ranging in size from 39.7 to 763.5 g (Fig. 1). As expected, large birds cooled more slowly than small ones. For example, in the first 20 minutes internal temperatures dropped less than 0.8 °C on Pterodroma-s'ize birds. Four of six birds showed a slight and brief increase in Tu during the first minute or two. We think this temporary increase was the result of continued cellular heat production immediately after death in the absence of convective (respiratory and circulatory) avenues of heat loss. This initial increase in temperature may mask some heat loss owing to the elapsed time between death and T^ measurements. However, temperatures of living birds and recently dead ones showed no observable differences (Table 3). DISCUSSION In collecting temperature information we attempted to eliminate as many biases as possible. Activity states of the birds immediately prior to temperature measurements undoubtedly accounted for some of the variation in the procellariiform birds whose body temperatures were summarized by Warham (1971). The difference between resting/ incubat- ing and active procellariiform birds amounted to about 2 °C (Farner Seabird Core Temperatures 15 Table 1. Deep body temperatures and body mass of seabirds. Mean + one standard devia- tion (range in parentheses). N Mass (g) Gaviiformes Gavia immer • 2 Procellariiformes Fulmarus glacialis 20 Calonectris diomedea 35 Puffinus gravis 25 Puffinus Iherminieri 35 Puffinus griseus 1 Pterodroma hasitata 9 Oceanites oceanicus 25 Pelecaniformes Phaethon aethereus 2 Sula bassanus 4 Phalacrocorax auritus 2 Charadriiformes Phalaropus lobatus 5 Phalaropus fulicaria 14 Stercorarius pomarinus 14 Stercorarius parasiticus 2 Larus marinus 4 Larus argentatus 6 Larus atricilla 11 Larus Philadelphia 2 Rissa tridactyla 10 Sterna hirundo 7 Sterna anaethetus 6 Sterna maxima 9 3588.0 ± 58.0 (3547.0-3629.0) 692.3 ± 78.5 (550.0-860.0) 591.6 ±81.8 (430.4-749.5) 615.1 ± 103.7(424.3-870.0) 206.6 ±20.5 (138.4-242.0) 774.0 441.5 ±68.8 (352.3-496.0) 33.5 ± 3.5 (25.9-39.4) 616.4 ± 12.1 (607.8-624.9) 3396.0 ± 383.0 (2898.0-3750.0) 1833.9 ±96.7 (1765.5-1902.3) 37.0 ±5.9 (31.6-46.6) 55.4 ± 9.9 (38.9-73.0) 743.8 ±58.7 (660.3-849.9) 522.8 ±9.3 (516.2-529.4) 1641.0 ±89.8 (1577.0-1774.0) 919.5 ± 120.7(778.0-1114.5) 333.7 ± 35.8 (277.9-424.6) 21 1.0 ±7.1 (206.0-216.0) 368.0 ±60.5 (294.7-448.4) 118.0 ± 14.4(95.5-142.4) 135.5 ± 15.4(117.9-154.3) 489.5 ±29.2 (456.7-543.1) Body temperature (°C) 39.7 ±0.2 (39.5-39.8) 39.9 ± 0.8 (38.0-42.0) 39.6 ±0.9 (38.2-41.0) 39.8 ±0.7 (38.6-41.2) 39.5 ± 1.0(36.5-41.2) 41.0 39.1 ±0.6 (38.0-40.0) 38.9 ± 1.3(37.0-42.2) 39.3 ± 1.1 (38.5-40.0) 40.7 ± 0.9 (40.0-42.0) 40.4 ± 0.6 (39.9-40.8) 39.9 ± 1.1 (38.8-41.5) 40.3 ± 1.0(38.2-42.6) 40.4 ± 1.3(38.4-43.3) 42.0 ±0.3 (41.8-42.2) 39.7 ± 1.0(39.2-41.2) 40.4 ±0.5 (39.5-41.0) 40.6 ± 1.3(37.8-42.0) 39.3 ±0.3 (39.1-39.5) 40.2 ±0.6 (39.4-41.2) 40.8 ± 1.1 (39.0-42.5) 40.4 ±0.8 (39.2-41.8) 40.1 ± 1.1 (38.0-41.1) 1956; Farner and Serventy 1959; Grant and Whittow 1983; Howell and Bartholomew 1961a, b; Warham 1971). Warham (1971) expressed doubt that the T^ of petrels flying at sea would be greatly increased, because of their energy-efficient methods of flight. Most of the temperatures presented here are from birds collected in flight, although some of the phalaropes were collected on the water. Nevertheless, most of the phala- ropes were actively foraging (i.e. swimming) rather than resting on the surface. We have no way of knowing how long an individual bird had been active or how long it had been resting before collection. Avian flight (especially in birds that do not soar) typically elevates body temperatures 1 to 2 °C above the level recorded for resting birds 16 Steven P. Platania, Gilbert S. Grant, David S. Lee Table 2. Body temperatures of male and female seabirds. Means not significantly different at P > 0.05. Male Female Species N 7 °C± 1SD N 13 °C+ 1SD Fulmarus glacialis 39.7+ 1.0 40.0 ± 0.8 Calonecths diomedea 15 39.610.9 15 39.7 ± 0.9 Puffinus gravis 7 39.6 ±0.7 15 39.9 ±0.8 Puffinus Iherminieri 15 39.3 ± 1.3 15 39.6 ±0.7 Oceanites oceanicus 9 38.6 ± 1.2 12 38.6 ± 1.2 Stercorarius pomarinus 4 40.1 ±2.2 9 40.3 ± 1.0 Larus atricilla 4 41.3 ± 0.8 6 39.9 ± 1.4 Rissa tridactyla 4 40.3 ± 0.9 6 40.1 ±0.4 (Berger and Hart 1974). The maximum Tu of flying birds can be seen in the upper part of the Tu range of Table 1. The T^ of seven petrel species averaged 39.7 + 0.7 °C, which is only about 0.9 °C higher than the mean compiled for 31 species by Warham (1971). This slight and insig- nificant difference may result from one or more of the following: activ- ity states of the bird, different investigator's techniques, and positioning of temperature probes (cloacal, preventricular, or visceral). We suspect, however, that it reflects the larger percentage of active birds in our sam- ples than in samples compiled by investigators working with nesting colonies. We found no body mass, sexual, seasonal, or hourly differen- ces in Tu within species. McNab (1966:54) argued that the "apparent correlation between the level of body temperature and the taxonomic group is really a corre- lation of weight and taxonomic group. (It should be noted that within both the ratites and penguins, small species have higher body tempera- tures than large species)." Warham (1971) presented evidence that the mean body temperature of petrels is significantly lower than that of non-procellariiform birds. Within the order Procellariiformes, regres- sion of T^ against body weight for our temperature (Table 1) likewise shows no correlation. Small petrels do not have higher body tempera- tures than do large ones, as Warham illustrated. This is true for our Charadriiformes as well. Our temperatures agree closely with the range of body temperatures reported by Dawson and Hudson (1970) for the orders Gaviiformes, Procellariiformes, Pelecaniformes, and Charadrii- formes. We found no evidence that stress and shock affected the body temperatures of still-living birds within 1 to 3 minutes after they were shot. The T^'s did not differ from those of recently expired birds (Table 3). Seabird Core Temperatures Table 3. Body temperatures and masses of dead and live birds. 17 Species Dead Live Temperature Mass Temperature Mass N °C+1SD g+lSD N °C+1SD g+ 1SD Calonectris diomedea 35 39.6 ± 0.9 591.6 db 81.8 12 39.5 ±0.7 577.5 ±66.6 Puffinus Iherminieri 35 39.5 ± 1.0 206.6 ± 20.5 16 39.9 ±0.9 208.9 ± 22.7 Pterodroma hasitata 9 39.1 ±0.6 441.5 ±68.8 3 39.8 ±0.7 491.1 ±34.6 Oceanites oceanicus 25 38.9 ± 1.3 33.5 ± 3.5 8 38.6 ± 1.8 32.2 ± 2.6 Phalaropus lobatus 5 39.9 ± 1.1 37.0 ± 5.9 5 40.9 ± 1.2 38.9 ± 6.7 Phalaropus fulicaria 14 40.3 ± 1.0 55.4 ± 9.9 7 40.9 ±0.9 51.8 ± 9.0 Sterna hirundo 7 40.8 ± 1.1 118.0± 14.4 4 41.4±0.8 127.3 ±20.1 Sterna anaethetus 6 40.4 ±0.8 135.5 ± 15.4 3 40.6 ± 1.1 127.6 ± 5.5 12- 10- • •• • • • • •• • • • •• ••, • ••• 5 ••. • •••• • •• • • "• -►• • •••"•••••• • • • • • • • • • 15 — I- 20 TIME Iminutesl Fig. 1. Cooling curves of six seabirds of various masses. Zero time is time of death. 1. Pterodroma, 448.9 g. 2. Pterodroma, 459.0 g. 3. Puffinus Iherminieri, 221.9 g. 4. Oceanites oceanicus, 39.7 g. 5. Calonectris diomedea, 763.5 g. 6. Puffinus Iherminieri. 232.5 g. 18 Steven P. Platania, Gilbert S. Grant, David S. Lee ACKNOWLEDGMENTS.— M. K. Clark, E. W. Irvin, and H. Baum aided in the field, and the U.S. Fish and Wildlife Service provided financial support (Contract No. 14 16 0009-80-044). H. Rahn originally suggested the study. G. C. Whittow, H. Rahn, J. E. Cooper, J. War- ham, and B. K. McNab critically reviewed the manuscript. We thank these people and agencies for their assistance. Contribution 1986-1 of the North Carolina Biological Survey. LITERATURE CITED Berger, M., and J. S. Hart. 1974. Physiology and energetics of flight. Pages 415-477 in D. S. Farner, J. R. King, and K. C. Parkes (editors). Avian Biology, Vol. 4. Academic Press, New York. Dawson, William R., and J. W. Hudson. 1970. Birds. Pages 223-310 in G. C. Whittow (editor). Comparative Physiology of Thermoregulation, Vol. 1. Academic Press, New York. Farner, Donald S. 1956. Body temperature of the fairy prion {Pachyptila turtur) in flight and at rest. J. Appl. Physiol. 8:546-548. , and D. L. Serventy. 1959. Body temperature and the ontogeny of thermoregulation in the Slender-billed Shearwater. Condor 61:426-433. Grant, Gilbert S., and G. C. Whittow. 1983. Metabolic cost of incubation by Laysan Albatross and Bonin Petrel. Comp. Biochem. Physiol. 74A:77-82. Howell, Thomas R., and G. A. Bartholomew. 1961a. Temperature regulation in Laysan and Black-footed Albatrosses. Condor 63:185-197. , and 1961b. Temperature regulation in nesting Bonin Island Petrels, Wedge-tailed Shearwaters, and Christmas Island Shearwa- ters. Auk 78:343-354. Lee, David S., and J. Booth, Jr. 1979. Seasonal distribution of offshore and pelagic birds in North Carolina waters. Am. Birds 33:715-721. McNab, Brian K. 1966. An analysis of the body temperatures of birds. Condor 68:47-55. Warham, John. 1971. Body temperatures of petrels. Condor 73:214-219. Accepted 11 April 1984 Spider Mites and False Spider Mites (Acari: Tetranychidae and Tenuipalpidae) Recorded from or Expected to Occur in North Carolina Michael K. Hennessey, David L. Stephan, and Maurice H. Farrier Department of Entomology, Box 7613, North Carolina State University, Raleigh, North Carolina 27695-7613 ABSTRACT.— Thirty-six species of spider mites have been collected from North Carolina and their host plants identified. An additional 57 species are known from the eastern United States on hosts that also occur in North Carolina. Seven species of false spider mites have also been collected in the state and their host plants identified. Twenty-nine others may occur in North Carolina, as their hosts are within the state. Spider mites (also called spinning mites, plant mites, red mites, and red spiders) and false spider mites (also called flat mites) are phytopha- gous arachnids usually with a body length of less than 1 mm in the adult stage. Some species are polyphagous and others are apparently mono- phagous. The life cycle of egg-larva-protonymph-deutonymph-adult may be spent on the host, or some stages may leave the host to estivate or hibernate in soil litter or to search for other hosts. One or more genera- tions may occur annually. Most species are known from both male and female specimens while others are apparently known only from females. Mites occur at characteristic locations on the host, such as leaf or fruit, depending on the species, and may usually be found in groups that include all life stages. They feed by puncturing plant cells with their chelicerae and eating the cell contents. The feeding, especially by large numbers of mites, may cause observable damage to plants in the form of bronzing, flecking, or curling of leaves. For this reason many species have been regarded as pests. More detailed accounts of the life histories of economically important species may be found in Jeppson et al. (1975). Several lists of spider mites and false mites from the eastern half of the United States have been published. Garman (1940) listed 15 species of spider mites and 1 species of false spider mite for Connecticut. Reeves (1963) catalogued 40 species of spider mites occurring on woody plants in New York. Mellott and Connell (1965) listed 20 species of spider mites and one species of false spider mite for Delaware. Thewke and Enns (1970) listed 38 species of spider mites representing 13 genera, and 1 1 species of false spider mites representing 3 genera, for Missouri. Brimleyana No. 12:19-27, September 1986 19 20 Michael K. Hennessey, David L. Stephan, Maurice H. Farrier Prasad (1970) recorded 20 species of spider mites and 3 species of false spider mites from Michigan. Flechtmann and Hunter (1971) catalogued 27 species of spider mites representing 10 genera for Georgia. Ten spe- cies of Tetranychidae but no species of Tenuipalpidae were recorded for North Carolina by Brimley (1938) and Wray (1967). Our list summarizes records for species of spider mites and false spider mites for North Carolina and provides information to collectors about additional species that might be found in the state when more extensive collecting is done. The list is based on published records and on approximately 1000 specimens, nearly all of which were collected in North Carolina, in the North Carolina State University (NCSU) Insect Collection, including Extension Entomology reference collections. An asterisk after the mite species name in Table 1 indicates specimens are in the NCSU collections and were associated with host plants also marked with an asterisk. Since the North Carolina climate encompasses mild coastal as well as cooler mountain elements and a very diverse flora, we included spe- cies recorded in the literature from Connecticut, Delaware, District of Columbia, Florida, Georgia, Louisiana, Maryland, Michigan, Missis- sippi, Missouri, New York, Ohio, Pennsylvania, South Carolina, Ten- nessee, Texas, and Virginia on the probability that their range includes North Carolina because suitable host plants and climate are present. The collection record given is for the state geographically nearest North Carolina, although the mite may be known from several other states. The recorded hosts for each species include one or two hosts that occur in North Carolina according to Radford et al. (1968). If more than two hosts are known, they are included under "others" and may be found by consulting the reference given for the mite species. The exception is for species in NCSU collections, where all of the hosts for our specimens are given. Common names of most hosts are listed, as these are the names that appeared with collection data; however, in some cases scien- tific names are presented, because these were given in the literature and no common names are provided in Radford et al. (1968). Nomenclature of hosts follows Radford et al. (1968) and Bailey Hortorium Staff (1976). Twenty-eight species of spider mites are represented by specimens in the NCSU collections, and eight more species are recorded in the literature as being from the state. Fifty-seven additional species may occur in the state. Five species of false spider mites are represented by specimens in the NCSU collections, and an additional two species are recorded intthe literature as having been collected in North Carolina. Twenty-nine additional species may occur in the state. Only 39% of the spider mites and 19% of the false spider mites recorded from the eastern half of the United States are known to be present in North Carolina. Thus, there is still a need for more thorough collections within the state. Future collectors are encouraged to record host information carefully, as correct plant species or cultivar identifica- tion is useful in identifying the mites and determining their host specificity. N.C. Spider Mites and False Spider Mites 21 Table 1. Spider mites and false spider mites recorded from or expected to occur in North Carolina. Collection Recorded Species record hosts Tetranychidae Aponychus spinosus (Banks) ' GA American elm, linden Beerella petiolaris Thewke 2 MO Helianthus petiolaris Bryobia praetiosa Koch* NC apple leaves and bark*, grass*, Japanese holly*, horse* [accidental], indoors*, leaf litter*, orchid*, cedar stump*, vetch*, others 3 B. rubrioculus (Scheuten)* NC apple leaves and bark*, grass*, others 3 Eotetranychus carpini borealis MO apple 2, sugar maple 2, others 4 Pritchard and Baker 2 E. carpini carpini (Oudemans) 3 NY apple, oak, others E. caryae Reeves 3 NY hickory, pecan, others E. clitus Pritchard and Baker* NC azalea*, blackberry*, others 4 E. coryli (Reck) 4 DC red maple E. crossleyi Flechtmann and Hunter ' GA chalk maple E. deflexus (McGregor) 5 SC azalea, coralberry E.frosti (McGregor) 3 OH blackberry, rose, others E. hicoriae (McGregor) 4 NC hickory, pecan, others E. lewisi (McGregor) 3 MI citrus, clover, others E. matthyssei Reeves 3 NY black locust, elm, others E. pallidus (Gar man) 3 NY alder, beech E. populi (Koch) 4 NC poplar, weeping willow, others E. pruni (Oudemans) 6 DC red maple, sugar maple, others E. querci Reeves 6 NY oaks, white birch E. sexmaculatus (Riley) 3 FL azalea, pyracantha, others E. smithi Pritchard and Baker 3 TN cotton, rose, others E. tiliarium (Hermann) 3 Atlantic coast linden, sycamore, others E. ulmicola (Reck) 2 MO American elm, elm E. uncatus Garman* NC apple*, pear*, rose*, others 3 E. sp. probably E. uncatus Garman* NC willow oak* Eurytetranychus admes MO incense cedar2, juniper 2, Pritchard and Baker 2 others 4 E. buxi (Garman)* NC boxwood* Eutetranychus banksi (McGregor) 4 FL castor bean, citrus, others Monoceronychus linki FL saw grass, others Pritchard and Baker 4 M. mcgregori Pritchard and Baker 4 FL St. Augustine grass M. scolus Pritchard and Baker 4 NC Bermuda grass, grass 22 Michael K. Hennessey, David L. Stephan, Maurice H. Farrier Table 1. Continued. Species Collection record Recorded hosts Mononychellus virginiensis (McGregor)* Oligonychus aceris (Shimer)* O. bicolor (Banks) 4 O. sp. probably O. bicolor (Banks)* O. boudreauxi Pritchard and Baker 4 O. coffeae (Neitner) 3 O. conifer arum (McGregor) 4 O. cunliffei Pritchard and Baker 4 O. endytus Pritchard and Baker ' O. sp. near O. endytus Pritchard and Baker* O. hondoensis (Ehara) 3 O. ilicis (McGregor)* O. indicus (Hirst) 3 O. letchworthi Reeves 6 O. milleri (McGregor)* O. modestus (Banks) ' O. newcomeri (McGregor) 3 O. nielseni Reeves 6 O. platani (McGregor) 3 O. pratensis (Banks) 3 O. propetes Pritchard and Baker O. stickneyi (McGregor) 3 O. ununguis (Jacobi)* O. viridis (Banks) 3 O. yothersi (McGregor)* Palmanychus steganus (Pritchard and Baker) 4 Panonychus caglei (Mellott) 3 P. citri (McGregor)* NC black locust*, locust* NC maple 4, sugar maple* NC white oak, willow oak, others NC willow oak* MS bald cypress FL camellia, grape, others FL arbor vitae, juniper, others FL pine GA American holly ', oleaster ' others 4 NC willow oak* NY Japanese cedar NC azalea*, camellia*, Cotoneaster sp.*, Japanese holly*, others 3 worldwide Johnson grass, sorghum, others NY Ostrya virginiana NC longleaf pine*, others 6 GA bamboo, corn probably apple, hawthorn, others theastern USA NY white pine NC boxwood 3, juniper 3, others 4 GA Johnson grass, wheat, others 3 NC hawthorn 2, oak 4, others 2 FL corn, rye, others NC Fraser's fir*, juniper*, others 3 GA, SC hickory, pecan NC azalea*, boxwood*, others 3 FL palmetto VA blackberry, kudzu, others NC citrus*, kumquat*, rose* P. ulmi (Koch)* Petrobia apicalis (Banks) silverberry*, others 3 NC apple leaves*, elm*, Ilex sp.*, kumquat*, peach*, others 3 GA crimson clover ', legumes ', others 3 N.C. Spider Mites and False Spider Mites 23 Table 1. Continued. Species Collection record Recorded hosts P. harti (Ewing)* NC P. latens (Miiller)* NC P. lupini (McGregor) ' GA Platytetranychus multidigituli NC (Ewing) ' P. thujae (McGregor)* NC Schizotetranychus asparagi DC (Oudemans) 4 S. camur Pritchard and Baker ' GA S. celarius (Banks) 3 GA S. garmani Pritchard and Baker 4 CT S. oryzae Rossi de Simons 3 TX S. schizopus (Zacher) 4 NY S. spireafolia Garman 4 PA Tenuipalpoides dorychaeta NC Pritchard and Baker 4 Tetranychus canadensis (McGregor)* NC T. cinnabarinus (Boisduval)* NC T. cocosinus Boudreaux 4 T. desertorum Banks* T. glover i Banks 3 T. homorus Pritchard and Baker T. lobosus Boudreaux* T. ludeni Zacher 3 T. magnoliae Boudreaux* T. marianae McGregor 3 T. mcdanieli McGregor 4 T. merganser Boudreaux* T. sp near T. merganser Boudreaux* T. mexicanus (McGregor) 3 LA NC LA NC NC LA4 NC FL NY NC NC TX clover*, wood sorrel*, others 3 cotton*, wheat*, others 3 grass, lupines honey locust arbor vitae 4, juniper*, others 4 asparagus fern, Asparagus sp. cane ', reed grass 4 bamboo, rice, others willow rice willow Spiraea alba var. latifolia, others black locust, honey locust, others cotton*, paper mulberry*, plum*, others 4 Areca sp.*, butter bean*, butterfly weed*, cotton*, Dracaena sp.*, Impatiens sp.*, Arabian jasmine*, marigold*, passion flower*, Schefflera sp.*, snap bean*, kuta squash*, tomato*, others 3 brambles, hackberry, others cotton*, grass*, others 3 cotton ash, hickory azalea*, "Nephthytis" sp. (probably Syngonium sp.)*, string bean*, wisteria*, others 7 beans 3, cotton 3, others 3 magnolia*, tulip poplar* cotton, passion flower, others apple 3, raspberry 3, others 4 cranberry*, privet*, others 4 European cranberry bush* Johnson grass, magnolia 24 Michael K. Hennessey, David L. Stephan, Maurice H. Farrier Table 1. Continued. Species Collection record Recorded hosts T. neocalidonicus Andre 4 T. schoenei McGregor* T. sinhai Baker ' T. tumidellus Pritchard and Baker* T. tumidus Banks* T. sp. probably T. tumidus Banks* T. turkestani (U gar o\ and Nikolski)* T. urticae Koch* FL butterfly bush, sweet potato, others NC apple leaves and twigs*, blackberry*, Japanese flowering cherry*, cotton*, raspberry*, weed*, others 4 GA Johnson grass, wild rye grass, others NC peanut* NC cotton*, eggplant*, Pilea sp.*, others 4 NC Schefflera sp.* NC clover*, cotton*, green bean*, lima bean*, peanut leaves*, soybean*, strawberry*, others 3 NC apple leaves and bark*, Areca sp.*, Japanese aucuba*, butterfly tree*, Japanese flowering cherry*, cotton*, cucumber*, dahlia*, Dracaena sp.*, Euonymous japonica*, day lily*, Fatshedera sp.*, gardenia*, gladiolus*, hollyhock*, Impatiens sp.*, locust*, peach*, peanut*, pear*, piggy-back plant*, primrose*, tomato*, water hyacinth*, wood sorrel*, others 3 T. yusti McGregor 3 DE soybean, grasses, others Tenuipalpidae Aegyptobia nothus NC bald cypress*, juniper*, Pritchard and Baker* others 8 A. sp. probably A. nothus NC cedar* Pritchard and Baker* Brevipalpus arcus FL camphor weed, weed Pritchard and Baker 8 B. bicolpus Pritchard and Baker 8 MD pawpaw B. butcheri Pritchard and Baker 8 FL Amaranthus sp. B. calif amicus (Banks) 8 MD apple, maple, others N.C. Spider Mites and False Spider Mites 25 Table 1. Continued. Collection Recorded Species record hosts B. colpodes Pritchard and Baker 8 FL Baccharis sp. B. columbiensis Thewke 2 MO sycamore B. docimas Pritchard and Baker 8 NC hickory, probably walnut B. ennsi Thewke 2 MO alumroot, ironweed, others B. floridanus DeLeon 8 FL red bay B. garmani Baker 8 SC alder 8, thoroughwort 8, others 2' 8 B. glomeratus Pritchard and Baker 8 VA basswood 2, oak 8, others 2 B. glymma Pritchard and Baker 8 FL lantana B. hybus Pritchard and Baker 8 FL hyacinth, weed B. lewisi McGregor 8 NC alder, grape, others B. lilium Baker 8 FL apple, azalea, others B. linki Baker 8 FL oak B. obovatus Donnadieu* NC Japanese aucuba*, azalea*, dumb cane*, Japanese holly*, Microphylla sp.*, Pittosporum sp.*, rhododendron*, others 8 B. sp. probably B. obovatus NC Dionaea sp. leaves*, maple* Donnadieu* B. ogmellus Pritchard and Baker 8 LA oak B. phoenicis (Geijskes)* NC apple*, false aralia*, goldenrod*, others 8 B. pinicola Pritchard and Baker 8 FL pine B. sayedi Baker 9 GA hickory, pecan B. xystus Pritchard and Baker 8 LA pecan 8, pignut hickory 2, others 8 Dolichotetranychus apaches FL a bromeliad Baker and Pritchard 8 D. salinas Pritchard and Baker 8 MD salt grass Pentamehsmus canadensis NY arbor vitae McGregor 8 P. erythreus (Ewing)* NC Andorra juniper*, juniper*, red cedar*, others 8 P. oregonensis McGregor* NC arbor vitae*, juniper*, others 8 P. /axz(Haller)8 DC yew Tenuipalpus argus FL yucca Baker and Pritchard 8 T. bakeri McGregor 8 FL oak, others T. carolinensis Baker 8 SC goldenrod T. celtidis Pritchard and Baker 8 FL hackberry 26 Michael K. Hennessey, David L. Stephan, Maurice H. Farrier Table 1. Continued. Species T. dasples Baker and Pritchard s T. pacificus Baker 3 T. rhysus Baker and Pritchard 8 Collection Recorded record hosts FL palmetto FL orchids FL Cyrilla racemiflora, magnolia, others * - mite specimens in NCSU collections and hosts associated with those specimens. 1 Flechtmann and Hunter 1971 2 Thewke and Enns 1970 3 Jeppson et al. 1975 4 Pritchard and Baker 1955 5 McGregor 1950 6 Reeves 1963 7 Boudreaux 1956 8 Pritchard and Baker 1958 9 Flechtmann and Davis 1971 ACKNOWLEDGMENTS.— We thank Janice Culpepper and Deanna Jones for typing the manuscript. The paper is number 9147 of the Jour- nal Series of the North Carolina Agricultural Research Service, Raleigh, North Carolina 27695-76 13. LITERATURE CITED Bailey Hortorium Staff. 1976. Hortus Third. Macmillan Publ. Co., Inc., New York. Boudreaux, H. Bruce. 1956. Revision of the two-spotted spider mite (Acarina: Tetranychidae) complex, Tetranychus telarius (Linnaeus). Ann. Entomol. Soc. Am. 49(l):43-48. Brimley, Clement S. 1938. The Insects of North Carolina. N.C. Dep. Agric. Div. Entomol., Raleigh. Flechtmann, Carlos H. W., and R. Davis. 1971. Some Acarina from Georgia pecans with notes on their biology. J. Ga. Entomol. Soc. 6(l):33-42. , and P. E. Hunter. 1971. The spider mites (Prostigmata: Tetranychi- dae) of Georgia. J. Ga. Entomol. Soc. 6(1): 16-30. Garman, Philip. 1940. Tetranychidae of Connecticut. Conn. Agric. Exp. Stn. Bull. (New Haven) 431:65-88. Jeppson, Lee R., H. H. Keifer, and E. W. Baker. 1975. Mites Injurious to Eco- nomic Plants. Univ. California Press, Berkeley. N.C. Spider Mites and False Spider Mites 27 McGregor, Earnest A. 1950. Mites of the family Tetranychidae. Am. Midi. Nat. 44(2):257-420. Mellott, John L., and W. A. Connell. 1965. A preliminary list of Delaware Acarina. Trans. Am. Entomol. Soc. (Phila.) 91:85-94. Prasad, V. 1970. Some tetranychoid mites of Michigan. Mich. Entomol. 3(1):24-31. Pritchard, A. Earl, and E. W. Baker. 1955. A revision of the spider mite family Tetranychidae. Mem. Pac. Coast Entomol. Soc. 2:1-472. , and 1958. The false spider mites (Acarina: Tenuipalpi- dae). Univ. Calif. Publ. Entomol. 14(3): 175-274. Radford, Albert E., H. E. Ahles, and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. Univ. North Carolina Press, Chapel Hill. Reeves, R. Marcel. 1963. Tetranychidae infesting woody plants in New York State, and a life history study of the elm spider mite Eotetranychus mat- thyssei n. sp. Cornell Univ. Agric. Exp. Stn. Mem. 380:1-99. Thewke, Siegfried E., and W. R. Enns. 1970. The spider-mite complex (Acarina: Tetranychoidea) in Missouri. Univ. Mo. Mus. Contrib. Monogr. 1:1-106. Wray, David L. 1967. Insects of North Carolina, Third Supplement. N.C. Dep. Agric. Div. Entomol., Raleigh. Accepted 5 March 1985 28 FISHERMAN'S GUIDE FISHES OF THE SOUTHEASTERN UNITED STATES by Charles S. Manooch, III Illustrated by Duane Raver, Jr. Remarkable for its breadth of coverage, this book details the hab- its, range, and appearance of more than 250 species offish, 150 of which are illustrated in color. Each account also includes tips on catching the fish and preparing it for the table. Manooch is experienced in the field of Fisheries Management and Technology, and Raver is nationally known for his paintings of wildlife. "An excellent general reference book for scientific or non-scientific audiences. ... It contains information not easily found in any other source." — Carter R. Gilbert, Curator of Fishes, Florida State Museum. 1984 376 pages Index Bibliography ISBN 0-917134-07-9 Price: $24.95, plus $1.25 for shipping. North Carolina residents add 414% sales tax. Please make checks payable in U. S. currency to NCDA Museum Extension Fund. Send to FISHERMAN'S GUIDE, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. Life History of the Wood Frog, Rana sylvatica LeConte (Amphibia: Ranidae), in Alabama Mark S. Davis1 and George W. Folkerts Department of Zoology- Entomology, Auburn University, Auburn, Alabama 36849 ABSTRACT. — A life history study of the wood frog, Rana sylvatica LeConte, was conducted from February 1978 to January 1980. All populations studied were in the Blue Ridge and Piedmont physiogra- phic provinces of Alabama, mostly in semideciduous forests along the flood plains of major streams. Breeding activity occured from mid- January to late February and coincided with the onset of warm winter rains. Most breeding occurred in semipermanent woodland pools. Ambystoma opacum and A. maculatum were consistent breeding associates. Usually present were Notophthalmus viridescens, Hyla cru- cifer, Pseudacris triseriata, P. brachyphona, and Rana sphenocephala. Mean clutch size in R. sylvatica was 496. Diameters of eggs and jelly envelopes are the largest reported for this species. Analysis of stomach contents indicated that adult frogs are opportunistic terrestrial feeders, but they apparently do not feed during the short, explosive breeding The wood frog, Rana sylvatica LeConte, is a small to medium-sized ranid frog with an extensive geographic range. Martof and Humphries (1959) reported its range as extending over approximately 4,044,000 square miles (more than 10,000,000 km2) from Alaska to Georgia. This range is exceeded in North America only by that of the Rana pipiens complex, which actually consists of several species. Its broad distribu- tion and the relative abundance of R. sylvatica over most of its range have prompted considerable research. Most information concerning its life history appears as scattered notes in general references, in papers presenting distributional information or ecology, in studies on amphi- bian community structure or reproductive behavior, and in accounts in various state herpetological publications. Rana sylvatica was first discovered in Alabama in 1974 (Mount 1975). Its presence was documented by three specimens from Mt. Cheaha, Cleburne County, in the east central part of the state. Prior to Mount's record, the southernmost locality for R. sylvatica was thought 1 Present address: Division of Biological Sciences, 110 Tucker Hall, University of Missouri, Columbia, Missouri 6521 1. Brimleyana No. 12:29-50, September 1986 29 30 Mark S. Davis and George W. Folkerts to be in northeastern Georgia, approximately 160 km northeast of the Alabama locality. The collection of additional specimens south of Mt. Cheaha and the paucity of information on Alabama populations pro- vided the impetus for the present study. Our attention focused on fea- tures of the frog's life history, for a cohesive study of this type (espe- cially in the southern part of the range) was lacking. Furthermore, the biology of any organism at the terminus of its range may provide insights into the adaptive significance of geographic variation in life his- tory parameters. MATERIALS AND METHODS Considerable effort was devoted to locating potential breeding ponds and breeding populations. Searching was confined chiefly to the Blue Ridge physiographic province and the upper sections of the Pied- mont Plateau. Wood frogs were collected by hand in breeding ponds and on highways during warm rains. Temperatures of water, air, or both were taken with a field thermometer at the time of collection. Most speci- mens were killed in 20% chloretone, then positioned and fixed in 10% formalin for at least 72 hours. Formalin was injected into the body cav- ity to preserve food and reproductive organs. Individuals were later transferred to 70% ethanol for permanent storage in the Auburn Uni- versity Vertebrate Museum. The stomach and intestine of each frog were removed, slit longitud- inally, and the contents washed into a culture dish. All food items were examined under a dissecting microscope and identified to the lowest possible taxon. The volume of food items was not determined. Ovaries or ovisacs were removed and their percentages of total body weight cal- culated. Ovarian or ovisacal eggs were counted, if present, and then stored in 70% ethanol. Snout-vent lengths (SVL) were determined by measuring from the tip of the snout to the posterior edge of the urostyle. Tibiofibula lengths (TFL) were taken by measuring the maximum length of the tibiofibula when the shank was completely flexed upon the thigh (Martof and Humphries 1959), and ratios of TFL to SVL were calculated. Snout length, defined as the distance from the anterior edge of the eye to the nostril, and snout height, taken as the straight-line distance from the nostril to the edge of the upper lip (Ruibal 1957), were also measured. All measurements were made with dial calipers to the nearest 0.1 mm after specimens had been kept in alcohol for at least 3 weeks. Information on reproduction was obtained primarily from field studies. Notes were made on calls of males, egg deposition, clutch size, Wood Frog Life History 31 egg development, egg mortality, and egg predators. Clutch size was determined by counting the number of eggs in six egg masses and by volumetric displacement of four additional egg masses. Estimates from volumetric displacement were obtained by placing an entire egg mass in a 1-1 graduated cylinder containing 200 ml of water. The volume of water displaced by each egg mass was recorded and then multiplied by a standard displacement volume, obtained previously for 10 eggs, to cal- culate the number of eggs present in the clutch. Larval development in the field was monitored to obtain growth and mortality data. A series of 10 or more tadpoles was collected at one pond during 1979 at varying intervals until no more tadpoles could be found. All were immediately preserved in 10% formalin, then measured and staged in the manner recommended by Gosner (1960). Rana sphenocephala tadpoles were also collected in the same ponds at the same times so that developmen- tal rates between the two species could be compared. RESULTS AND DISCUSSION Habitat and Range in Alabama Wood frogs were collected in five counties in east central Alabama (Fig. 1). Except those on or near Mt. Cheaha, Cleburne County, all collecting localities were near mesic semideciduous forests along the flood plains of large streams. Several frogs collected on Mt. Cheaha were considerable distances from running water, but were never far from mesic sites. All frogs were collected from localities in the Blue Ridge and Piedmont Plateau physiographic provinces. The floristic and geologic components of these areas have previously been described (Harper 1943, Hodgkins 1965, Johnson and Sellman 1975). The Blue Ridge, as used here, is synonymous with the Blue Ridge herpetofaunal province de- scribed by Mount (1975) and the Mountain Forest Habitat Region defined by Hodgkins (1965) and Johnson and Sellman (1975). From a geological standpoint, the terms "Blue Ridge" or "Mountain" may be inappropriate, for the general consensus among geologists is that the Blue Ridge province terminates in northern Georgia. However, the vegetative distribution patterns and faunal components differ suffi- ciently from the Ridge and Valley province and Piedmont to warrant recognition of the Blue Ridge as a separate entity in Alabama (Johnson and Sellman 1975, Mount 1975). All Piedmont localities for breeding ponds and adult frogs were in the northern subdivision known as the Ashland Plateau. To the north, this part of the Piedmont makes contact with the Blue Ridge, but the transition is gradual with a continuous gradation of the Piedmont into the uplands. The southern subdivision 32 Mark S. Davis and George W. Folkerts of the Piedmont, known as the Opelika Plateau, is geologically less complex. The different surface configurations and geological structures in the two plateaus have resulted in some differences in vegetation dis- tribution (Johnson and Sellman 1975, Golden 1968). Mount (pers. comm.) stated that his more recent studies of the herpetofauna of this area indicate that a more distinct transition exists between the Opelika and Ashland Plateaus of the Piedmont than between the latter and the Blue Ridge, if a distinction is to be made. The southernmost locality known for R. sylvatica in North Amer- ica lies just south of the Tallapoosa River in Horseshoe Bend National Military Park, Tallapoosa County, Alabama. The site is near the boun- dary of the Ashland and Opelika plateaus in the central Piedmont. The southern boundary of the range of the wood frog in Alabama approxi- mates the southern edge of the Ashland Plateau, although it is likely that many populations are isolates. One wood frog has been collected in Calhoun County, Alabama (L. G. Sanford, pers. comm.), at the northern edge of the Blue Ridge, and represents the northernmost record for this species in Alabama. The known range in Alabama thus extends from the northern edge of the Blue Ridge along its contact with the Ridge and Valley province to the southern edge of the Ashland Plateau in the central Piedmont. Scat- tered populations probably occur in suitable habitat in that part of the Ridge and Valley province south of the Coosa Valley. Mount (1975) mentioned that wood frogs might occur in the higher elevations of the Appalachian Plateau in extreme northeastern Alabama (Jackson Coun- ty). If so, they are probably derived from populations that moved southward on the Cumberland Plateau from Tennessee and not from populations in the Blue Ridge. Known localities for R. sylvatica in Georgia are limited to five counties in the Blue Ridge of the northeastern part of the state (Wil- liamson and Moulis 1979; C. W. Seyle, pers. comm.) (Fig. 2). No spec- imens have been collected in the 160-km-long area between the Georgia and Alabama wood frog populations, apparently because this area of Georgia has been inadequately surveyed (R. E. Daniel, C. W. Seyle, pers. comms.). Since suitable habitat does occur in this area, we feel that the Alabama wood frog populations are continuous with those in northeastern Georgia (Fig. 2). Based on our knowledge of the habitat requirements of this species in Alabama (and in the southern Appala- chians), the presumed range in the intervening area is thought to be limited to the Blue Ridge (Blue Ridge, Cohuttas, Talladega Upland subdivisions), the southern part of the Great Valley, and the Upland and Gainesville Ridges subdivisions of the northern Piedmont (see Wharton 1978). Wood Frog Life History 33 Fig. 1. Known Alabama localities for Rana sylvatica determined during this study. Open circles represent breeding localities; inset shows position in the state of counties from which R. sylvatica is recorded. Adult Characteristics Snout-vent lengths of adult male wood frogs in Alabama averaged 50.0 mm (SD = 5.5, N = 20), and adult females averaged 60.0 mm (SD = 3.16, N = 18). These values are smaller than those given by Martof and Humphries (1959) for wood frogs in northern Georgia and western North Carolina (males: x = 54.8 mm; females: x = 66.8 mm). Berven (1982a) discovered size differences along an altitudinal gradient from Maryland (lowland populations) to western Virginia (montane popula- tions). Mountain males and females were larger (males: x = 55.3 mm; females: x = 64.4 mm) than individuals from lowland populations (males: x = 41.7 mm; females: "x = 47.7 mm). Because Berven hypothe- sized that selection acted primarily on egg size and that selection for 34 Mark S. Davis and George W. Folkerts increased fecundity would secondarily favor large body size, comments on the size of Alabama R. sylvatica are reserved for the later section on egg size. Martof and Humphries (1959) established evidence for a latitudinal gradient in relative leg length in wood frogs and found that the frogs with the longest legs occur in the southern Appalachians. We calculated TFL/SVL ratios for 19 adult males and 19 adult females to determine if this trend was evident in Alabama R. sylvatica. Tibiofibulas of males averaged .602 of the SVL, those of females averaged .625 — a value iden- tical to that obtained by Martof and Humphries (1959) for both sexes. Ruibal (1957) reported a latitudinal and altitudinal clinal gradient in snout length for R. pipiens and pointed out evidence for a similar latitudinal gradient in R. sylvatica. Blunt snouts were defined as those with high height /length (H/L) values (> 1.15), pointed snouts as those with low H/L values (~ 1.00). Wood frogs from northern Canada (locality not given) possessed blunter snouts (x = 1.30 mm, R = 1.07 to 1.50, N = 14) than those from New York (x = 1.11 mm, R = 1.00 to 1.22, N= 15). Snout lengths were measured on Alabama wood frogs to determine if this apparent cline continued. The mean H/L value was found to be 0.83 (R = 0.74 to 0.95, N = 37). These results further substantiate the evidence for a clinal increase in snout length southward. Martof and Humphries (1959) and Martof (1970) described Appalachian wood frogs as having blunt snouts, apparently a subjective description for no quan- titative H/L analysis was performed. The coloration of adult R. sylvatica in Alabama is typical of the Appalachian phenotype described by Martof and Humphries (1959). A color photograph resembling the Appalachian phenotype may be found in Behler and King (1979, Fig. 216). Breeding Ponds, Breeding Associates, and Breeding Season All 14 breeding congregations of R. sylvatica found were in shallow (usually < 45 cm), temporary pools in or adjacent to forests. These pools fill with winter rains from December through February. Most were located in semideciduous woods along the flood plains of large streams. Three sites were found in pastures; however, these were bor- dered by semideciduous woods and probably had been wooded in the past. Breeding ponds differed in the amount and type of vegetation. Ponds in open (pasture) situations received more sunlight and were gen- erally characterized by vigorous growths of Eleocharis sp., Juncus sp., and Carex spp. Woodland pools generally had fewer rushes and sedges, probably because of reduced sunlight. Peltandra virginica, Sagittaria Wood Frog Life History 35 Fig. 2. Distribution of Rana sylvatica at the southern terminus of its range. Solid circles indicate counties (not localities) in Georgia and Alabama where specimens have been taken. Presumed range is indicated by hatching (see expla- nation in text). latifolia, Saururus cernuus, Sparganium americanum, and Ranunculus sp. were the most common plants in these situations. Alnus serrulata, Quercus spp., and Cornus spp. were usually present along the edges of the pond or in shallow water. A fairly consistent assemblage of breeding associates was present with R. sylvatica during the breeding season. Amby stoma maculatum and A. opacum were present at every site. Notophthalmus viridescens, Hyla crucifer, Pseudacris triseriata, and P. brachyphona were common associates. Bufo americanus and Rana sphenocephala were always pres- ent in pasture breeding ponds. Collins and Wilbur (1979) reported that, in Michigan, R. sylvatica, H. crucifer, and P. triseriata were breeding associates, particularly in temporary aquatic habitats. A number of previous accounts described wood frogs as explosive breeders that generally spend only a few days in the breeding ponds (see Seale 1982), and we found this to be true of Alabama populations. Males begin calling with the onset of the first heavy, warm winter rain 36 Mark S. Davis and George W. Folkerts from mid- January to late February. Calling began on 21 February dur- ing the 1979 breeding season (air and water temperatures 11 °C). Although egg deposition was completed in all pounds by 1 March, males continued to call intermittently until 5 March. Similar postbreed- ing calling was noted by Waldman (1982). Breeding occurred sporadi- cally between 21 February and 1 March whenever air temperatures were above 5 °C. Vigorous calling occurred on 22 February (air 16 °C, water 14 °C) and 24 February (air 16 °C, water 12 °C). No calls were heard after 5 March even though air and water temperatures were above 10 °C. In 1980, males began calling on 17 January after unseasonably warm weather (air 15.5 °C, water 9 °C, at 2130 CST), but calling ceased early the next morning when the temperature dropped considerably (air 4 °C, water 9 °C, at 0200). Males were in full chorus in all ponds visited on 22 January (air 9 °C, water 12 °C, at 1915). Egg deposition was completed in all ponds by 22 January. Development of Ovarian Eggs Females collected throughout the year yielded information on egg development. Body weights of preserved specimens before ovary re- moval, ovary weights from each specimen, and ovary weights as a per- centage of body weights are given in Table 1. Eggs were stored in ovisacs in one female that had completed ovulation. For this specimen, ovisacs (instead of ovaries) with ripe oocytes are expressed as a percentage of total body weight, as indicated. Examination of females indicated that ovarian weight (expressed as a percentage of body weight) remains fairly constant from early Sep- tember to late November. No preovulating females were collected immediately prior to the breeding season, but we assume that the great- est increase in size of oocytes occurs during this time (later stages of vitellogenesis). All gravid females collected in the breeding ponds had completed ovulation and mature ova were present in the ovisacs. Ovi- sacs in a gravid female collected in the breeding pond on 18 January composed 41.7% of the total body weight. Ovarian weight in spent females drops to 3.9 to 5.0% of total weight. The ovaries of a female collected on 14 May were macroscopically similar in appearance to those of spent females. In this female, ovaries equaled 5.3% of total body weight. Because no females were collected between May and September, we could not determine when oocyte enlargement begins; however, based on the size and appearance of oocytes in females collected during Sep- tember, we estimate that enlargement begins in July or August. Because ripe oocytes in gravid females compose such a large percentage of total Wood Frog Life History 37 Table 1. Body and ovarian weights (g) and ovarian weight as a percentage of body weight for adult female Rana sylvatica collected in Alabama at different times of the year. Date Body weight Ovary weight Ovary weight as pecentage of body weight 3 September 22.4 3.3 14.7 29 September 38.4 3.7 9.6 12 November 24.9 3.1 12.4 12 November 20.4 2.9 14.2 23 November 31.3 4.3 13.7 23 November 27.4 4.0 14.6 23 November 29.1 4.8 16.5 17 January 52.0 21.7* 41.7 (female in breeding pond) 18 January 10.5 0.6 5.0 (spent female) 18 January 17.6 0.7 3.9 (spent female) 14 May 13.2 0.7 5.3 * ovisac weight body weight, an extended period is probably necessary for a female to reach reproductive condition. Redshaw (1972) reported that amphibian oocyte enlargement from 450/i to 1400/i required a period of 9 months. Sexual Dimorphism, Calling, and Amplexus Sexual dimorphism is more pronounced during the breeding sea- son. Males are generally much darker than females, the ground color ranging from deep brown to almost black. Howard (1980) noted that this darker color matched the dark color of the water in breeding ponds. Females are usually tan to reddish brown during the breeding season. Darker females are observed occasionally, but they are never as dark as males. The margin of the toe webbing between the digits of the hindlimb is markedly convex in males. The male thumb (first digit on the fore- limb) and the musculature in the forelimb are also enlarged during this time, as in other ranid species. Noble and Farris (1929) thought that the additional surface area provided by convexity of the toe webbing allowed males more mobility in the water, a suggestion consistent with the behavior of males during the breeding season. The thumbs of male 38 Mark S. Davis and George W. Folkerts wood frogs in Alabama are slightly enlarged throughout the year, becoming more conspicuously so during the breeding season. Toe web- bing in females remains concave during the breeding season, a condition found in both sexes throughout the rest of the year. The call of male wood frogs in Alabama is similar to that described for males elsewhere (Thoreau 1881, Hinckley 1882, Dickerson 1906, Smith 1961, Martof 1970, Minton 1972). It usually consists of two high- pitched croaks or snappy clacks, and may be described as a nasal "back- up," repeated rapidly several times in succession. Solitary males call less frequently, repeating the call only once or twice at varying intervals. Calling males float or swim at the water surface with forelimbs hanging down and hindlimbs projecting posteriorly. The digits on the hindlimbs are expanded, exposing maximum webbing surface. In large aggregations, males move and interact frequently (see Wright 1914, Noble and Farris 1929, Wright and Wright 1949, Howard 1980, Berven 1981). Calling males are extremely wary and dive below the water sur- face at slight disturbances, concealing themslves under leaf litter and decaying vegetation on the bottom, or hiding among roots or emergent vegetation. When calling from fairly open water, males are nearly unap- proachable. If the pond is small, with emergent vegetation, one can usu- ally approach close enough to observe floating males. Calling males can be heard continuously (though often sporadically) from dusk until dawn, but the chorusing is usually strongest immediately after sunset. Males in almost all Alabama populations call only at night; diurnal choruses were heard at only one breeding pond (W. Baker, pers. comm.). Two lethargic males were collected from the bottom vegetation and leaf litter at one pond during midday. The apparent diel restriction of calling activity is not as conspicuous in more northerly populations (Wright and Wright 1949, Howard 1980, Berven 1981, Waldman 1982). This might be a function of the extremely small population sizes in Ala- bama and the resultant lack of stimulation by large numbers of conspecifics. Females in Alabama populations are less conspicuous than chorus- ing males and usually remain below the surface of the water. This behavior is similar to that noted by Banta (1914) and Noble and Farris (1929). Only one female was seen floating on a pond surface; all others collected in breeding ponds were taken under water while in amplexus. Amplexus is axillary (pectoral), with males clasping females just poste- rior to the forelimbs. One interspecific amplexing pair was observed during this study — a male wood frog clasping a female R. sphenocephala. When approached, the male released his hold and swam away. The female, partly covered Wood Frog Life History 39 with vegetation, remained on the bottom of the pond. The freshly de- posited R. sphenocephala egg mass (eggs had completely cleared the cloacal opening) was resting on the posterior surface of the female and ob- scured her hind limbs. Whether or not the male R. sylvatica had extruded sperm over the eggs is not known. The cause of the apparent breakdown in isolating mechanisms in this case is also unknown. Per- haps a breakdown in habitat isolation is a partial explanation. Rana sylvatica and R. sphenocephala were found as breeding associates only in sites where the forest had been removed. In Alabama, R. sylvatica usually breeds in woodland pools, whereas R. sphenocephala breeds in a variety of open aquatic habitats, as well as in woodland pools. Eggs of both species were found in only one woodland pool during this study. Rana sphenocephala has often been seen breeding in woodland pools in other areas of Alabama (R. H. Mount, pers. comm.). Some ecological separation may occur in the part of the state where the two species are sympatric. Nelson (1971) mentioned a female R. sylvatica that was clasped by a male R. pipiens. None of the R. sylvatica eggs fertilized by the R. pipiens developed beyond gastrulation. Moore (1955) found that devel- opment did not proceed beyond gastrulation in experimental laboratory reciprocal crosses of R. sylvatica and R. pipiens. Interspecific pairing of male R. sylvatica with other amphibians in the laboratory was reported by Wright (1914). OVIPOSITION The eggs of R. sylvatica in Alabama are laid as submerged globular masses, usually attached to vegetation. Often, upper portions of the egg mass become emergent. Moore (1949) pointed out that the deposition of submerged egg masses by northern ranid species (those adapted to cool climates) is an adaptation that helps protect the developing embryos from freezing. The rapid drop in water level that often occurs in tem- porary pools in Alabama may cause exposure of the egg masses in cer- tain situations. Desiccation then becomes an added mortality factor. The depth of water in which oviposition occurs is fairly consistent, averaging 15 to 20 cm. Wood frogs characteristically have communal oviposition sites. The advantages of this behavior have been discussed (Wells 1977, Howard 1980, Seale 1982, Waldman 1982, Waldman and Ryan 1983). Commu- nal oviposition sites (COS) were encountered in this study only in breed- ing ponds with larger populations. The largest such site was in a pond south of Mt. Cheaha where 147 egg masses, arranged in two layers, were found in an area 1.5 x 1 m square. Another COS (65 egg masses) 40 Mark S. Davis and George W. Folkerts was in a small woodland pond in northern Tallapoosa County. In both ponds all egg masses were restricted to the COS. The third largest popu- lation (60 egg masses) was in a pasture breeding pond, where communal oviposition occurred, but to a lesser extent. In this pond 31 egg masses were in a communal site, and the rest were deposited in small clumps separate from the COS. The 1 1 other wood frog breeding ponds discov- ered during this study were characterized by extremely small popula- tions (compare Howard 1980, Berven 1981, Seale 1982, Waldman 1982). The number of egg masses found in each of these ponds varied from 4 to 28, and the tendency toward communal oviposition was less pronounced. Clutch size varied from 350 to 709 eggs per mass (x = 496). Ovarian and /or ovisac counts indicated that oviposition may occur once or twice during the breeding season. The number of ovarian eggs per female ranged from 618 to 966. When all eggs are deposited at one time, the resultant egg mass appears as two fused masses, indicating that females empty each ovisac separately. If a female moves to another site after emptying one ovisac, the resultant egg mass represents approximately one-half the ovarian complement. This probably accounts for much of the apparent variability in clutch sizes observed in the field. Even so, there is some variability in reproductive potential, as evidenced by the range in egg complements seen in gravid females. This is probably attributable to a combination of individual and ontogenetic variation. Seale (1982) found no significant difference between clutch size and ovarian egg counts in Pennsylvania wood frogs (clutch: x~ = 895; ovarian eggs: x = 840). Although there are few data available concerning ovarian egg counts, several authors have presented information on clutch size (Table 2). There is some evidence for smaller clutch size in the southern parts of the range, although this trend may be obscured by altitudinal differences (Berven 1982a). Clutch size probably varies in response to different selection pressures throughout the geographic range, creating a chaotic pattern of variation. Furthermore, variation in clutch size should be viewed with respect to differences in adult body size and egg size. At the southern terminus of the frog's range, the probability of egg mortality resulting from freezing is reduced and may be a factor in decreased clutch size. Moore (1949) pointed out that the submerged egg masses of northern species of Rana were poorly adapted for higher pond temperatures because diffusion of oxygen would not be rapid enough to supply the metabolic needs of embryos in the center of the egg mass. Thus, the smaller egg masses characteristic of southern popu- lations of R. sylvatica would allow a more rapid diffusion of oxygen to these inner embryos. However, Savage (1961) claimed that egg masses possess intercapsular channels and that gaseous diffusion need not take Wood Frog Life History 41 place through the entire egg mass. If so, smaller clutch size resulting from selective pressures for small egg mass size would be an inapprop- riate hypothesis. The eggs of Alabama wood frogs are the largest reported for any population of R. sylvatica (x = 2.9 mm diam., SD = .08, N = 50). Com- parison of these values with previously published information indicates a general trend for egg diameter to increase southward (Table 2.). Berven (1982a) hypothesized that selection has acted primarily on egg size, and that other reproductive traits such as clutch size, body size, and age at first reproduction have evolved secondarily. Different selec- tive pressures in different environments would confer differential selec- tive advantages on particular sizes of eggs, clutches, and adults (see Berven 1982a,b, for discussion). The large size of eggs in Alabama populations of R. sylvatica is probably a consequence of increased fitness (larger size) of larvae hatch- ing from these eggs (see Berven 1982a,b), a phenomenon that would result in faster growth rates and shorter larval periods. In Alabama populations, selection for rapid metamorphosis would probably result from breeding exclusively in temporary ponds. An additional selective pressure for more rapid metamorphosis in R. sylvatica may be the con- current breeding of R. sphenocephala in the same sites. This does not seem to be the case farther north. Berven's hypothesis concerning the relationship of large egg size to large body size is difficult to support with data from Alabama wood frogs. Although egg size is largest in Alabama populations, adults are somewhat smaller than those reported in other parts of the southern Appalachians (see earlier mention). Because determination of different age classes was not possible during this study, size comparisons and determinations of age and size at first (and subsequent) reproduction await further study. Larger sample sizes obtained by future workers will probably help to clarify this situation. The diameters of egg jelly envelopes for R. sylvatica in Alabama are larger than values reported in other parts of the range. Diameters of inner envelopes averaged 6.6 mm (R = 5.4 to 7.2 mm, N = 50); outer envelopes averaged 14.0 mm (R = 12.4 to 17.3 mm, N = 50). Few data are available on more northerly populations (Table 2). The reasons that jelly envelopes of Alabama wood frogs are so much larger than those in northern populations are not obvious. Perhaps jelly deposition is con- trolled by egg size, with larger eggs receiving more jelly. Egg Fertility, Development, and Predation Fertility, although variable, was quite high, and several egg masses exhibited 100% fertility. Three clutches were entirely infertile, perhaps a result of oviposition in the absence of a clasping male. Early mortality 42 Mark S. Davis and George W. Folkerts of developing eggs was occasionally observed. These eggs usually were infested with fungi, which probably invaded after egg death rather than having been the cause of mortality. Wood frog egg masses were easily recognized in breeding ponds by their characteristic shape and the large size of their jelly envelopes. Another distinguishing feature was a greenish color imparted to the jelly envelopes by a unicellular green alga. Dickerson (1906) first noted the presence of this alga and assumed that the relationship was mutualistic. Gilbert (1942) also observed this alga in jelly envelopes of wood frog eggs and identified it as Oophila amblystomatis, a species characteristi- cally found in the egg jelly of Ambystoma maculatum. Surprisingly, there has been little inquiry into the relationship between wood frog eggs and algae by subsequent workers (see mentions by Pope 1964, Gatz 1973). Although the relationship between A. maculatum and Oophila has generally been viewed as mutualistic (Gilbert 1942, 1944; Hutchin- son and Hammen 1958; Hammen and Hutchinson 1962), a higher rate of mortality has been related to the presence of the alga in some cases (Anderson et al. 1971, Gatz 1973). Further investigation concerning the relationship between the alga and R. sylvatica eggs is warranted. All egg predators observed during this study were invertebrates. Mayfly naiads (Siphlonuridae, Ephemerellidae) and isopods (Asellidae) were often present between adjacent egg envelopes within egg masses. Caddisfly larvae (Phryganeidae) fed on the external surfaces of egg masses, and one leech, Macrobdella decora, was found feeding on an egg mass. Cory and Manion (1953) found this same leech destroying the majority of wood frog eggs in some situations in Indiana, and thought that its presence in certain populations of R. sylvatica might constitute a check on population size. Since only one M. decora was observed dur- ing our study, the effect of this species on Alabama wood frog popula- tions is probably minimal. Hudson (1954) reported newts, Notophthalmus viridescens, feeding on wood frog eggs in Pennsylvania. This salamander was a potential egg predator in Alabama wood frog breeding ponds, but predation was never observed during our study. The large diameters of egg jelly enve- lopes of R. sylvatica in Alabama populations might reduce newt predation. Hatching, Larval Development, and Larval Mortality The length of the period between egg deposition and hatching var- ies directly with water temperature. Under field conditions, wood frog tadpoles generally hatch in 7 to 9 days after eggs are deposited (water temperatures variable, 5 to 17 °C). Larvae hatch at a fairly advanced developmental stage, usually stage 20 (gill circulation, Gosner 1960) or Wood Frog Life History 43 T3 u « E g .2 w <3 "S ►n > w c e > 00 O «* U i » r- sO On >» oo c 2 c ~ S3 r o £ 2 ^ 2 a "2 £ r- -• On » s o ? 5 S ffl U PQ ^ 1 > oo £ 3 Cd On 13 -a cd xi O 2 Q £ § 8 & *** — * C/3 *£ Cfl O ed Xi CM CM 00 oo On Os — ' — ' C5 c c s u u 2 > > c h m £ a> a> CD CQ 9 £2 oo ^t «n On OO Tt *> so 3 II O Pi CM II IX o so «-< II II |X IX i-l i-' .5 o Tt oo CM oo CM CM CM ON CM «-3 CM CM* cm ii CM J1 Ix ift II II IX |X gC Ift o o §§ §§ «-» CN II II cd £ 00 00 ON o o o § §8 6 6 6 8£o§ — g < etj a: o 0.10). Distance moved between captures ranged from 40 to 760 m (x= 390 m). The recapture data were not appropriate to derive an estimate of actual population density (White et al. 1982). However, there are approximately 13.6 ha of habitat suitable for hognose snakes (excluding salt marsh and aquatic sites) on the spit. Using only the 66 individuals captured, the absolute minimum popula- tion density of H. platy rhinos was 4.8 snakes/ ha. Two hatchlings were found dead on the road on 15 and 19 August, and were 18.5 and 19.0 cm SVL. No hatchlings were captured during the late summer, so their growth rate could not be determined. Two juvenile males (SVL < 36.0 cm) and one adult male (62.0 cm SVL) were recaptured more than a month after initial capture. These two size classes exhibited average summer growth rates of 2.2 and 1.0 cm/ month, respectively. Piatt (1969) observed higher growth rates in H. platyrhin- os juveniles (3.4 cm/ month) and lower rates in large males (0.8 cm/ month). Growth rates for female size classes could not be determined. Notes on Eastern Hognose Snake 53 CO 0 03 O CO "co "D D 03 O _q CO 0 -Q E 60 55 50 45 40 35 • male • a a a oDo a aaa a a a a a a an a a a female 120 125 130 135 140 145 Number of ventral scales 150 Fig. 1. Scale counts used to sex 66 Heterodon platy rhinos individuals found on Assateague Island, Virginia, June-September 1981. Females were significantly longer (t = 3.8, df = 64, p < 0.02) and also exhibited greater body mass (t = -2.74, df = 64, p < 0.01) than males (Table 1). Covariance analysis was used to scale body mass for differen- ces in snout-vent length, using SVL as the covariate. Female H. platy- rhinos were no heavier than males of the same SVL (F i,63 ) = 2.61, p > 0.10). Sexual dimorphism in body length has been described in several populations of Heterodon (Edgren 1961, Piatt 1969) and for other spe- cies (Fitch 1981, Gibbons 1972, Shine 1978). Piatt (1969) attributed the sexual size dimorphism in H. platyrhinos to faster growth rates in females. Larger females of some species produce larger clutches, thereby possibly promoting selection for increased body size in females (Shine 1978, Semlitsch and Gibbons 1982). Of the 66 individuals captured, 55 were judged to be normal in coloration, 3 were melanistic, and 8 were intermediate (very dark with some light markings). All melanistic snakes were adults. Observations on the number of dorsal blotches revealed that females have more blotches than males (t = 4.22, df = 64, p < 0.001; Table 1). Moreover, these means were also different (more than 2 SE) from values reported 54 David Scott by Edgren (1961) for the Delmarva Peninsula population of H. platy- rhinos. This difference suggests a change in gene frequency of the "blotch" allele(s), possibly owing to founder effect and the genetic isola- tion of the island hognose population, or perhaps a change of selective pressures in an island environment. Most of the 72 captures were made from early to middle morning. Only five snakes were captured in late afternoon (1700 to 1930), and none were found on the five night searches. Substrate temperature ranged from 24 °C to 39 °C for 52 captures. Snakes appeared to be most active when substrate temperature was 32 °C to 35 °C (N = 22). The data were grouped into early and late summer captures of adults (males > 36.0 cm, females > 40.0 cm SVL) to test whether sexes differed in their summer activity patterns. Seventeen adult males and 8 adult females were captured in early summer (prior to 8 July), and 1 1 males and 14 females in late summer. These proportions were tested using a binomial test of proportions (Lewis 1966), which tended to indi- cate differences in activity (U = 1.76, p < 0.08). In addition, for the period 1 1 June to 12 September, adult males had a median capture date of 23 June. The median for females was a month later (23 July). Nine of the 10 largest females were captured after 22 July. Females in this popu- lation probably laid their eggs in late June or early July, assuming an incubation period of 45 to 55 days (Piatt 1969). In contrast to Piatt's study, in which few adult females were captured after laying eggs, females on Assateague Island appeared to be most active after oviposi- tion. Females were less active early in the summer when they were gravid, as has been reported for other species (Jackson and Franz 1981, Shine 1979). ACKNOWLEDGMENTS.— I thank J. W. Gibbons, J. Congdon, S. Morreale, R. Semlitsch, T. Lamb, and C. Vincent for comments on the manuscript. J. Hoover and R. Schneider assisted with field work. This research was supported by a grant from the U.S. Fish and Wildlife Ser- vice to R. D. Dueser and W. E. Odum of the University of Virginia. Manuscript preparation was aided by Contract EY-76-C-09-0819 between the U.S. Department of Energy and the University of Georgia (Institute of Ecology). LITERATURE CITED Conant, Roger. 1975. A Field Guide to the Reptiles and Amphibians of Eastern and Central North America. Houghton Mifflin, Boston. Corrington, John D. 1929. Herpetology of the Columbia, South Carolina region. Copeia 1929(72):59-83. Duellman, William E., and A. Schwartz. 1958. Amphibians and reptiles of southern Florida. Bull. Fla. State Mus. 3(5): 181-324. Notes on Eastern Hognose Snake 55 Table 1. Body mass, snout-vent length (SVL), and dorsal blotch number in male and female hognose snakes, Heterodon plat y rhinos (x ± 1 SE). Sex N_ Body mass (g) SVL (cm) Dorsal blotches Male 38 94.9 ± 9.6 44.2 ± 1.8 21.8 ± 0.27 Female 28 140.6 ± 14.5 54.4 ± 2.7 23.7 ± 0.37 Edgren, Richard A. 1961. A simplified method for the analysis of clines: geo- graphic variation in the hognose snake (Heterodon plat y rhinos Latreille). Copeia 196 1(2): 125- 132. Fitch, Henry S. 1981. Sexual size differences in reptiles. Univ. Kans. Mus. Nat. Hist. Misc. Publ. 70. Gawne, Constance E. 1966. Shoreline changes on Fenwick and Assateague Islands, Maryland and Virginia. BS thesis, Univ. Illinois, Urbana. Gibbons, J. Whitfield. 1972. Reproduction, growth, and sexual dimorphism in the canebrake rattlesnake (Crotalus horridus atricaudatus). Copeia 1972(2):222-226. , and J. W. Coker. 1978. Herpetofaunal colonization patterns of Atlantic Coast barrier islands. Am. Midi. Nat. 99:219-223. Jackson, Dale R., and R. Franz. 1981. Ecology of the eastern coral snake (Micrurus fulvius) in northern peninsular Florida. Herpetologica 37(4):21 3-228. Lee, David S. 1972. List of the amphibians and reptiles of Assateague Island. Bull. Md. Herp. Soc. 8(4):90-95. Lewis, Alvin E. 1966. Biostatistics. Reinhold Publ. Corp., New York. Lynn, William G. 1936. Reptile records from Stafford County, Virginia. Copeia 1936(3): 169-171. Martof, Bernard S., W. M. Palmer, J. R. Bailey, and J. R. Harrison. 1980. Amphibians and Reptiles of the Carolinas and Virginia. Univ. North Carolina Press, Chapel Hill. Piatt, Dwight R. 1969. Natural history of the hognose snakes Heterodon pla- tyrhinos and Heterodon nasicus. Univ. Kans. Publ. Mus. Nat. Hist. 18:253-420. Semlitsch, Raymond D., and J. W. Gibbons. 1982. Body size dimorphism and sexual selection in two species of water snakes. Copeia 1982(4):974-976. Shine, Richard. 1978. Sexual size dimorphism and male combat in snakes. Oecologia 33:269-277. 1979. Activity patterns in Australian elapid snakes (Squamata: Ser- pentes: Elapidae). Herpetologica 35(1): 1-11. White, Gary C, D. R. Anderson, K. P. Burnham, and D. L. Otis. 1982. Capture-recapture and removal methods for sampling closed populations. LA-8787-NERP. Los Alamos National Laboratory, Los Alamos, New Mexico. Accepted 5 August 1985 56 A DISTRIBUTIONAL SURVEY OF NORTH CAROLINA MAMMALS by David S. Lee, John B. Funderburg, Jr., and Mary K. Clark This book lists all the mammals of North Carolina and offers spe- cies accounts and range maps for all of the non-marine species. Intro- ductory chapters describe the plant communities of the state as they relate to mammal distribution and discuss local zoogeographic patterns. 1982 72 pages Softbound Price: $5, postpaid. North Carolina residents add 4'/2% sales tax. Please make checks payable in U. S. currency to NCDA Museum Extension Fund. Send to MAMMAL BOOK, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. A Study of Variation in Eastern Timber Rattlesnakes. Crotalus horridus Linnae (Serpentes: Viperidae) Christopher W. Brown and Carl H. Ernst Department of Biology, George Mason University, Fairfax, Virginia 22030 ABSTRACT. — Variation was examined in specimens of Crotalus horridus from the eastern United States in an attempt to investigate the status of its two described subspecies, C. h. horridus and C. h. atricaudatus, as defined by Gloyd. A particular effort was made to duplicate the results of a study by Pisani, Collins, and Edwards, who concluded that the subspecies were invalid. Maximum likelihood fac- tor analysis and step-wise discriminant analysis on the same morpho- logical characters, plus several others relating to adult size and pattern, produced evidence that the two subspecies of C. horridus are valid in the eastern portion of its range. However, standard morphological characters alone are not sufficient to discriminate between the two forms. Rather, adult size and pattern differences, in conjunction with the number of dorsal scale rows and ventral scales, best differentiate C. h. horridus from C. h. atricaudatus. Two subspecies of the rattlesnake Crotalus horridus are thought to occur in the eastern United States (Conant 1975): C. h. horridus, the timber rattlesnake, and C. h. atricaudatus, the canebrake rattlesnake. Gloyd (1940) defined the former as having 23 dorsal scale rows, a lower number of ventral and caudal scales, an absent or faint postocular stripe, and less brilliant contrast between the ground color and pattern. He defined the latter as having 25 dorsal scale rows, a higher number of ventral and caudal scales, larger size, and more brilliant markings. The geographic range of the two races is shown in Figure 1. Crotalus horridus shows considerable variation in the western por- tion of its range; C. h. atricaudatus is not known to occur in Oklahoma, yet specimens of C. h. horridus from southeastern Oklahoma resemble C. h. atricaudatus in color and pattern (Webb 1970). Anderson (1965) found that populations of C. h. horridus from western Missouri pos- sessed a reddish-brown middorsal stripe like that of C. h. atricaudatus from southeastern Missouri. Gloyd (1940:186) also reported that "the middorsal stripe of reddish brown, although very conspicuous in typical (C. h. atricaudatus) specimens, is not a good definitive character because of its common occurrence in specimens of C. h. horridus from western localities." Smith (1961) regarded Illinois specimens from Jackson Brimleyana No. 12:57-74, September 1986 57 58 Christopher W. Brown and Carl H. Ernst 7" Fig. 1. Range of Crotalus horridus (from Klauber 1972). County and southward as intergrades, because the specimens of atri- caudatus from counties bordering the Mississippi River more closely resembled horridus in some characters. A study by Pisani et al. (1973) concluded that, on the basis of 13 morphological characters, the recognition of subspecies in C. horridus could not be justified. They examined specimens from localities through- out the range, including western populations where intergradation is thought to occur. The purpose of this study was to examine variation in pattern and adult size differences in addition to those morphological characters used by Pisani et al. (1973) in eastern C. horridus to determine if a more comprehensive study of the species is needed. MATERIALS AND METHODS Data were obtained on 337 museum specimens from New Hamp- shire, Vermont, Massachusetts, Connecticut, New York, New Jersey, Pennsylvania, Maryland, Virginia, North Carolina, South Carolina, Georgia, and Florida. However, only 101 specimens were suitable for the analyses used here in that they were complete in all characters exam- ined. Twenty-one specimens were from localities of probable intergrada- tion, and so were treated separately. Of the remaining 80, 10 were juve- niles and were eliminated from some analyses. Localities of the 101 specimens used are shown in Figure 2. The characters used in this study Variation in Crotalus horridus 59 Fig. 2. Localities of specimens used in this study. Each circle represents at least one Crotalus horridus. Solid circles represent specimens from localities of probable intergradation. A question mark indicates an unknown locality for the state. Dashed lines are approximate range limits for each subspecies (see Fig. 1). 60 Christopher W. Brown and Carl H. Ernst are listed in Table 1; the first 13 are those of Gloyd (1940) and were also used by Pisani et al. (1973). However, the method of counting cross- bands was probably different in this study; the band was not counted if it was interrupted by at least one scale of ground color (Fig. 3). Specimens not from localities of probable intergradation were clas- sified a priori into one of the two forms (subspecies) based on that race's distribution as defined by Gloyd (1940). The 21 probable intergrades (Fig. 2) not used in the analyses were classified into groups based on their localities: those from within the range of C. h. horridus, those from within the range of C. h. atricaudatus, and those from localities lying between the two ranges. Fig. 3. Method of counting dorsal scale rows (numbered at top) and crossbands. Of the three apparent bands, only one complete crossband would be counted here, for only one is uninterrupted by any scales of ground color. Variation in Crotalus horridus 61 Table 1. Characters examined in this study of variation in Crotalus horridus. The first 13 were used by Pisani et al. (1973). No. Character Description 12 13 14 ADS DSM PDS VS CS 6 DCS 7 LSL 8 RSL 9 LIL 0 RIL 1 BCB TCB SVL 5 TL 6 HL 7 MS 8 GC 19 POS Anterior dorsal scale rows, counted at one head-length posterior to the occipit (see Fig. 3). Dorsal scale rows at midbody. Posterior dorsal scale rows at one head-length anterior to the anal plate. Number of ventral scales, not including the anal plate (Dowling 1951). This method was not used by Pisani et al. (1973T Number of caudal scales, starting with the first complete scale posterior to the anal plate. Number of divided caudal scales. Left supralabials. Right supralabials. Left infralabials. Right infralabials. Number of complete body crossbands, counted between the head and anal plate (Fig. 3). This is not the method used by Pisani et al. (1973), who were not sufficiently clear on how crossbands were distinguished from blotches. Number of complete tail crossbands. Although many specimens possessed tail markings that suggested banding, relatively few had tail crossbands that were clearly entire. Most specimens had a dark-colored tail with no markings, the dark color extending well anterior to the vent. Ratio of tail length to snout-vent length. Tail length was mea- sured from the posterior margin of the anal plate to the base of the first rattle segment. Adult snout-vent length. Individuals longer than 750 mm were considered adults, but this may have failed to exclude a few sub-adults. Adult tail length. Adult head length, measured from tip of rostrum to line joining posterior tips of mandible (Peters 1964). Middorsal stripe, coded as zero for either faint or completely absent and as one for clearly present. Ground color, an attempt to measure pattern contrast. Because many museum specimens had lost some of their original color, this was coded as either zero to denote light colors, such as pale brown, tan, pinkish, yellowish, and pale gray, or as one to denote dark colors, such as plain brown, dark gray, and dark olive-greenish. Some melanistic specimens were examined, but none were used in the analyses because they lacked other essen- tial characters. Postocular stripe, coded as zero for absent or faint and as one for clearly present on one or both sides of the head. 62 Christopher W. Brown and Carl H. Ernst "o s: o. o e en &> "O c c/a o >> o 03 V c oo o3 -o jm c 'as o >> u a > C/3 _>. 03 C 03 03 C 03 <*i T3 ;■— I C O cu o JC tD H c/3 CU Bfl JZ X) 03 H "O C/3 c '55 o j>^ CJ *03 u c en 03 2 >> •a Efl c '55 o >> o "o3 cu c 00 03 b LV3 s_^ «/-> sO 00 "* v-> On «r> o d d £^ Q oo < Q ^t Tf On m rs sO — Os 00 cs B "♦J o d d d o CU o c .5 ft 'a? a C to j3 CU "l_l •€ O > 03 > 03 O 03 Ih 03 CJ Id _^ „_ ~— > c 3 5 i- O £ =3 U O o 03 C 03 Variation in Crotalus horridus 63 Table 3. Results of discriminant analyses. The first analysis was performed on the 13 characters used by Pisani et al. (1973); all 19 characters were used in the second analysis. First Second analysis analysis Number of variables in discriminant function 5 4 Eigenvalue 1.088 2.989 Wilks' lambda 0.479 0.251 Approximate F-value (P = 0.01) 16.103 48.577 Canonical correlation 0.722 0.866 Coefficients for canonical variable -0.156 (BCB) -0.109 (HL) -0.216 (VS) -0.119(VS) -0.346 (TCB) -0.376 (ADS) -0.419 (DSM) -2.217 (MS) -0.514 (PDS) Constant 57.593 35.626 A maximum likelihood factor analysis (Dixon and Brown 1979), in which all variables are evaluated simultaneously, was employed prima- rily to determine the existence of groups that correspond to subspecies. Two factor analyses were conducted, first on the 13 morphological characters used by Pisani et al. (1973), and then on all 19 characters. To analyze group integrity, we used stepwise discriminant analysis, which, like the factor analysis, evaluates all variables simultaneously (Dixon and Brown 1979). Again, two discriminant analyses were conducted, one on the characters used by Pisani et al. (1973) and one on all 19. The maximum number of discriminant functions to be derived in a one discriminant analysis is either less than the number of groups or the same as the number of discriminating variables, whichever is smaller (Nie et al. 1975). Because there are only two groups in this study, there is only one discriminant function. Three criteria for evaluating this func- tion are the eigenvalue, canonical correlation, and Wilks' lambda. The eigenvalue is a measure of the total variance explained by the discrimi- nating characters. The canonical correlation is a second measure of the function's ability to discriminate among the groups. Wilks' lambda is an inverse measure of the discriminating power in the characters that have not been removed by the discriminant function. A smaller lambda, then, means more information is accounted for in the discriminant function. In Biomedical Computer Programs (BMDP), the Wilks' lambda is transformed into an approximate F-value. Since there is one discriminant function, there can only be one canonical variable, which is the linear combination of variables entered that best discriminates among the groups (the largest one-way ANOVA F-value) (Dixon and Brown 1979). The canonical variable is adjusted so that the pooled within-group variance is one, and its overall mean is 64 Christopher W. Brown and Carl H. Ernst zero. The canonical variable is then evaluated at the group mean for each specimen, and all cases are plotted in a histogram to demonstrate separation of distinct groups. Table 3 lists the constant and canonical coefficients of the discriminating characters for each analysis. Figure 6 shows a comparison of the histogram from each discriminant analysis. Analysis was performed at the George Mason University Comput- ing Services on the Cyber 170-720 computer system. The P-series of the BMDP (Dixon and Brown 1979) was used, as were all default proce- dures, except the second factor analysis, in which four factors were requested. Specimens Examined: Carnegie Museum (CM): S 9130; 36497, 40186, 40187, 40192, 54721, 91446, 91447, 91482-91484, 91582, 91583, 91677, 92053, 92056, 92057, 92063, 92065 North Carolina State Museum (NCSM): 2347, 5744, 8035, 8041, 8121, 8520, 8725, 9638, 9655, 9772, 9879-9885, 9888, 10229, 10779, 10920, 11017, 11259, 11874, 11875, 12011, 12061, 12108, 12112, 12113, 12263, 12266, 12795, 12857, 12894, 12911, 13899, 14011, 14111, 14141, 15678, 15793, 15926, 16657, 16711, 17056, 17059, 17105, 17150, 19241, 19359, 19595, 19641,21808 National Museum of Natural History (USNM): 8372, 9973, 10519, 14755 (2 specimens), 17959, 19970, 20651, 29362, 44313, 49958, 101858, 102714, 107879, 108687, 110487, 127601, 129094, 129759, 130167, 130168, 139618-139620, 145377, 156804,210092,218911. RESULTS Four factors accounting for 52% of the variance were produced in the factor analysis of the first 13 characters. The variation in characters CS and R correlated most closely with factor 1; ADS and DSM with factor 2; RIL with factor 3; and LSL, RSL with factor 4. Character variation that correlated less than 0.500 with any factor was not consi- dered significant. The factor loadings and eigenvalues are summarized in Table 2, and estimated factor scores for the 80 specimens used are plotted in Figure 4. In the factor analysis of all 19 characters, 4 factors were requested to limit the number produced. These accounted for a cumulation of 52% of the variance, but the characters SVL, TL, HL, and POS correlated most closely with factor 1; CS, R, and TL with factor 2; ADS and DSM with factor 3; and MS with factor 4. These results are summarized in Table 2. Factor scores for the 70 specimens used are plotted in Figure 5. Fig. 4. Scatterplots of estimated factor scores for specimens from the factor analysis of the first 13 characters. Solid circles represent one or more specimens of Crotalus h. horridus; open circles represent specimens of C. h. atricaudatus. Tail measurements (characters CS and R) correlated most closely with factor 1; dorsal scale rows (ADS and DSM) correlated with factor 2; infralabials (RIL) correlated with factor 3; and supralabials (LSL and RSL) correlated with factor 4. 2.4 J 0 1.8; 1.2: 0 0 *° °# °o F A c o° 0 0 0: .60; 0.0* -60: • 0 % s OOI 1 0 o A -.750; 4 - 1.50: •2.25 o • • •o« ^•o,.^ , *Q /r 2.10 -1.40 -.700 0.00 .700 1.40 2.10 FACTOR 1 3.00 0 2.25 1.50 00 °0° ; F A .750 ••'•-I • • ° 0 •o'V C • • • mmrm M £ ° T 0.00 0 R-.750 3 -1.50 ^ ° 0 0 00: 0 • •° o# 0 0 • -2.25 •• 0 0 2.10 -1.40 -.700 0.00 .700 1.40 2.10 FACTOR 2 3.75 3.00 2.25 f 1.50 A I .750 T 0.00 -.750 -1.50 -2.25 • • • 0 0 • ••o. 000 0 ° ° i %M& .*<*oo* 8 •nrfti', >cf° 8* °o • • : 3.75 3.00 2.25 I 1.50 C T .750 O R 0.00 4 -.750 -1.50 -2.25 •- "a!>. JU OQ»««L o o -2.10 -1.40 -.700 0.00 .700 1.40 2.10 FACTOR 2 -2.4 -1.6 -.80 0.0 .80 1.6 2.4 FACTOR 3 66 Christopher W. Brown and Carl H. Ernst Initial stepwise discriminant analysis of the first 13 characters pro- duced 5 discriminating characters. They were, in order of their increas- ing ability to discriminate, characters BCB, VS, TCB, DSM, and PDS. None were strongly correlated; the highest was 0.365 between VS and DSM. The single discriminant function had a significant F-value (P = 0.01) of 16.103 and a canonical correlation of 0.722. These results, including the value of the Wilks' lambda, are summarized in Table 3. Thirteen (16%) of the 80 specimens used in this analysis were incorrectly classified into the two groups: C. h. horridus, 6 (13.6%), and C. h. atri- caudatus, 7 (19.4%). Discriminant analysis of all 19 characters produced 4 discriminat- ing characters: MS, ADS, HL, and VS. Characters MS and HL had a weak correlation of 0.633, the next highest correlation being 0.366 between HL and VS. The discriminant function had a significant F- value (P = 0.01) of 48.577 and a canonical correlation or 0.866. These results, including the value of the Wilks' lambda, are summarized in Table 3. Five (7%) of the 70 specimens used in this analysis were incor- rectly classified into the groups: C. h. horridus, 4 (11.1%), and C. h. atricaudatus, 1 (2.9%). Group means and standard deviations of all 19 characters are pre- sented for both nonintergrades and intergrades in Tables 4 and 5, respectively. DISCUSSION One purpose of the factor analysis performed was to determine, by inspection of the plotted factor scores, whether clusters of individuals occur that correspond to subspecies. Analysis of the first 13 characters reveals little or no clustering in any of the scatterplots. Factor 1 (ab- scissa) versus factor 2 (ordinate) appears to have the best clustering of the six graphs (Fig. 4A). Separation seems to occur along the vertical axis. Crotalus h. atricaudatus tends to cluster in the first two quadrants, while C. h. horridus tends to cluster in quadrants three and four, indi- cating separation on the basis of dorsal scale rows (factor 2). However, overlap is wide. More than 25% of the specimens of C. h. atricaudatus lie below the first two quadrants. No other plot (Fig. 4B-F) demon- strates any distinct clustering. On the basis of the first 13 characters, therefore, no subspeciation can be recognized. The plots from the analysis of all 19 characters, however, show contrary results. Factor 1 (abscissa) versus factor 2 (ordinate) demon- strates clustering along the horizontal axis: C. h. atricaudatus tends to Fig. 5. Scatterplots of estimated factor scores for specimens from the factor analysis of all 19 characters. Solid circles represent one or more specimens of Crotalus h. horridus; open circles represent specimens of C. h. atricaudatus. The X's represent specimens of both. Adult size measurements and postocular stripe (characters SVL, TL, HL, and POS) correlated most closely with factor 1; tail measurements (CS, R, and TL) correlated with factor 2; dorsal scale rows (ADS and DSM) correlated with factor 3; and middorsal stripe (MS) correlated with factor 4. 1.8 1.2 I .60 C T 0.0 O R -.60 2 -1.2 -1.8 -2.4 • o o» o i "v • o o • o : • • • • • o • • •< • o o •o Boo"o j o ° ° 6> ° : O ; O 3.0 o 2.4 o F 1.8 o A C 1.2 o °#o o T • o o o 0 .60 R 3 °-° - .60 - 1.2 • o o " f M • • • \ - • • • • 3f o o» o o • o • -1.8 • -1.8 -1.2 -.60 0£) .60 1.2 1.8 FACT O R 1 1.8 -1.2 -.60 0.0 .60 \2 1.8 FACTOR 1 3.0; o 2.4 j o 1.8: o 1-2: o • ° o o : .60; QO- o° o o a* o° ° : .60 • ■\2\ "Pi o • t : • -1,8; • • • o us o 1.0 # ° 9,° • o o • o .50 % ( ° • o ) ° oo 0.0 o ° -.50 •. •• • • • • • •.• o • • - 1.0 •••«!» • • • -1.5 o • • -2JD 8 • -2.10 -1.40 -.700 000 .700 1/*0 2.10 FACTOR 2 -2.10 -1.40 -.700 0.00 .700 FACTOR 2 1.40 2.10 1.5 1.0 F .50 A C 0J3 T O -.50 R 4 -1.0 -1.5 -2.0 V ° ° o°°oo< • o o o : o oo o s • • r. • • • ° ° : o | •• • •• • o ° o 1.5 1.0 F .50 A C 0.0 T O-.50 R 4-1.0 -1.5 -2.0 o "o 'i o o*° o o o o °o ^0 o oo o o • o ° ."Jr%. . o «•*••! i • • ° • • 8 ■1/»0 -.700 OJOO .700 1*0 2.10 2.80 FACTOR 3 ■1.8 -1.2 -.60 OJO .60 1.2 1.8 F ACT O-R 1 68 Christopher W. Brown and Carl H. Ernst occur in quadrants one and four, and C. h. horridus in quadrants two and three (Fig. 5A). Factor 1 (adult size and postocular stripe), then, appears to differentiate C. horridus into two forms. Of those specimens of C. h. atricaudatus occurring in the second and third quadrants, only one lacks a postocular stripe, and all are under 1000 mm snout-vent length, the smallest individuals of their group. For example, the speci- men of C. h. atricaudatus having the largest negative factor 1 score is only 765 mm snout-vent length (probably a subadult). Similarly, those specimens of C. h. horridus lying in the first and fourth quadrants are the physically largest individuals of their group. Such large or small individuals, though not typical of their group, can be expected. Factor 2, which includes character R, demonstrates that the ratio of tail length to snout-vent length as an indicator of size is not as reliable as the lengths themselves (factor 1) in distinguishing the two groups. For instance, a large snake having a correspondingly large tail could have the same ratio as a smaller snake, or even one of a different species. Factor 1 (abscissa) versus factor 3 (ordinate, Fig. 5B) produces a scatterplot much like plot 5A, indicating that factor 3 (dorsal scale rows), like factor 2 (adult tail measurements), is relatively unimportant in differentiating the two subspecies. Factor 1 (adult size) again pro- duces good separation of the two groups in plot 5B with the same individ- uals lying far to the left or right of their respective groups as seen in plot 5A. Since factor 2 (adult tail measurements) and factor 3 (dorsal scale rows) have been shown to be unimportant in distinguishing the two groups, the plot of factor 2 versus factor 3 would be expected to demon- strate no clustering, and this is observed in Figure 5C. In the plot of factor 2 (abscissa) versus factor 4 (ordinate), cluster- ing occurs along the vertical axis, with most specimens of C. h. atricau- datus in the first two quadrants and those of C. h. horridus in the last two (Fig. 5D). A similar plot occurs for factor 3 versus factor 4, as would be expected (Fig. 5E). Factor 4 (middorsal stripe) therefore appears to differentiate the specimens into two groups: those possessing a distinct middorsal stripe (factor 4 greater than zero, which corre- sponds to C. h. atricaudatus) and those possessing an indistinct middor- sal stripe or none at all (factor 4 less than zero, which corresponds to C. h. horridus). Specimens with factor scores outside the normal range of variation for their group were examined more closely in plots 5D and 5E to determine why they clustered with the "wrong" group. Those few C. h. horridus that possessed a distinct middorsal stripe (factor 4 greater than zero) were all from localities in North Carolina and Georgia where intergradation might occur. None, in other words, came from localities well to the north of the C. h. atricaudatus range. Those few specimens of C. h. atricaudatus having a large negative factor 4 score all possessed a faint middorsal stripe, rather than lacked one entirely, which classified Variation in Crotalus horridus 69 Table 4. Mean character values (x) and standard deviations (s) of all 19 charac- ters for the two subspecies of Crotalus horridus. Numbers in paren- theses are sample size. One specimen from each subspecies was not included because of unknown sex. M = male, F = female. C. h. horridus C. h. atricaudatm M (14) F(21) M(ll) F (22) x, s x, s x, s x, s ADS 25.29, 1.54 25.38, 0.97 27.54, 1.44 26.14, 1.64 DSM 23.43, 1.22 23.38, 0.80 24.46, 0.93 24.18, 1.05 PDS 18.57, 0.76 18.67, 0.73 19.27, 0.65 19.00, 0.62 VS 163.00, 2.94 166.67, 3.42 167.09, 2.34 169.54, 3.04 CS 23.64, 1.01 19.81, 1.12 25.54, 2.66 20.23, 1.90 DCS 1.50, 1.65 1.48, 1.03 3.18, 2.99 1.04, 1.13 LSL 13.86, 0.86 13.71, 1.10 13.36, 0.92 14.09, 0.92 RSL 13.86, 1.01 13.33, 0.86 13.27, 0.90 14.00, 0.98 LIL 14.50, 1.02 14.76, 0.89 15.54, 0.69 15.04, 0.95 RIL 15.21, 0.70 14.86, 1.01 15.54, 1.29 15.18, 1.14 BCB 11.14, 3.50 9.14, 3.90 12.54, 3.67 11.82, 2.58 TCB 0.57, 1.02 0.00, 0.00 1.54, 2.12 0.36, 0.95 R 0.09, 0.01 0.07, 0.00 0.09, 0.01 0.07, 0.01 SVL 939.71, 133.96 849.52, 59.77 1049.91, 128.87 1097.91, 116.14 TL 81.71, 10.62 56.95, 5.52 91.00, 13.03 72.86, 8.17 HL 42.42, 4.36 39.25, 2.98 47.52, 4.57 48.24, 4.34 MS 0.29, 0.47 0.10, 0.30 1.00, 0.00 0.86, 0.35 GC 0.36, 0.50 0.67, 0.48 0.09, 0.30 0.41, 0.50 POS 0.29, 0.47 0.14, 0.36 0.82, 0.40 0.96, 0.21 them into the C. h. horridus group. Of the properly classified C. h. horridus specimens, about 20% possessed an indistinct stripe and 90% lacked one altogether. Since adult size and middorsal stripe appear to be the most impor- tant factors, one plotted against the other (factor 1 versus factor 4) should yield good separation of the groups along both axes, which is the case in Figure 5F. Crotalus h. atricaudatus clusters in the first quadrant, and C. h. horridus in the third quadrant. The individuals lying outside their respective clusters are a combination of aberrant individuals in the previous plots and have already been discussed. Characters DSM, PDS, VS, BCB, and TCB were determined to be the combination of variables that best discriminated in the discriminant analysis of the first 13 characters. Pisani et al. (1973) reported characters CS, VS, DSM, and R (in decreasing order of discriminating ability) as the most discriminating in their analysis. Some differences would be expected in light of the different geographical areas sampled. In addi- tion, the method of counting crossbands was different in this study (see 70 Christopher W. Brown and Carl H. Ernst 1 * < g u U » 2 -c £ "& ll 60 £ O XI O ; O en c/5 «| CO s a cd O cd cd XI u C S cd s- 0) o «n , CA c TD Bl cd cn i^T Q S m r- o O O O — co NO o — 'OOCOOCNOOO — CNO O NO 00 rt o o o IX 0000 O ^t CN CN O O O no o o »T) 00 o o o o m Tf Tt NO >n o o CO wS CN co — ^ r- no ^r ON fx ON OO O O OO CN 'BH «g NO O ON O © © «n © on in NO NO ON ON CO © © © © © © CO m © ir— o —^ —' © © © © in — no ON — — © © © © © © © © © © © © Tf — O © © IX © in© - vo - ^h^^-^^^m inNOTt — ON © © © © © © 00 S cn M jj Q^Q^c^nJ^w UQ jo! _, co pa ti s u u J oi cq H J J cn H EC 2 O a- Variation in Qrotalus horridus 71 above). For example, in their study mean body bands ranged from 23.14 to 25.05 among all groups. In our study, body crossbands aver- aged from only 9.14 to 12.54 (Table 4). The differences in mean tail bands is similar. In our first analysis these characteristics discriminated between the two subspecies, whereas they did not in Pisani et al. (1973); however, their method may have measured an entirely different charac- ter variable than the one they intended. In the discriminant analysis of all 19 characters, 2 of the 6 added characters discriminated. These were HL and MS. Because head length is probably indicative of the total length of the adult snake (Klauber 1938, 1972), it appears that adult size and pattern are important in dis- criminating between the two subspecies. Dorsal scale rows and number of ventrals also discriminated here, as in Pisani et al. (1973), except that the discrimination was by ADS instead of DSM. Characters BCB and TCB did not discriminate in the second analysis. In comparing the two analyses, we find that all 19 characters allowed better discrimination. This is evident in the larger eigenvalue (2.989 vs. 1.088), the larger canonical correlation (0.866 vs. 0.722), the smaller Wilks' lambda (0.251 vs. 0.479), and better classification of indi- viduals into the two groups (7% incorrectly classified vs. 16%). In the canonical variable histograms (Fig. 6), separation of the two groups is much better in the analysis of all 19 characters, again showing the importance of size and pattern. Comparison of the discriminating characters ADS, VS, and HL between nonintergrades and intergrades (Tables 4 and 5) shows that, as expected, the mean character values of the intergrade specimens lie between the mean character values of C. h. horridus and C. h. atricau- datus, regardless of sex. Since the remaining characters other than MS did not discriminate, their mean values for the intergrade specimens are not expected to be intermediate or even different from the mean values of either horridus or atricaudatus. Interestingly, all intergrades pos- sessed a distinct middorsal stripe, much like intergrades reported from western localities (Gloyd 1940, Smith 1961, Webb 1970). Of the three new pattern characters tested in this study (MS, GC, POS; Table 1), POS and, especially, MS were important. None, how- ever, was completely free from subjectivity in measurement. In some cases, the distinction between light- and dark-colored or indistinct and clearly visible was a fine line. Use of old museum specimens, many faded by preservatives, may have induced too much subjectivity, how- ever unintentional. Some dark-colored specimens had a middorsal stripe that had apparently faded to an almost white color, making the stripe unusually conspicuous. Had the specimens been living, the stripe may have been inconspicuous. In other specimens the ground color was faded and difficult to determine. Another problem is the inadequacy of the coding scheme for GC. Gloyd (1940) and others (Wright and Wright 1957, Conant 1975) mentioned that C. h. horridus has two color 72 Christopher W. Brown and Carl H. Ernst phases — the typical dark one, as tested for in this study, and a yellow one. There was no possible way, in certain cases, for this study to determine whether a light-colored C. h. horridus specimen was truly the yellow phase, as opposed to a badly faded normal dark phase, or the color of a typical C. h. atricaudatus specimen. Use of ground color as a discriminating character obviously requires fresh or living specimens and an improved coding scheme. The evidence presented here suggests that, on the basis of differen- ces in adult size and pattern, two subspecies of Crotalus horridus (as described by Gloyd 1940) occur east of the Appalachians. There the races are clearly more distinct than in the western populations. Stand- ard morphological characters alone are not sufficient to separate the two taxa; rather, adult size and pattern differences, in conjunction with the number of dorsal scale rows and ventral scutes, best discriminate C. h. horridus from C. h. atricaudatus. This combination of size, pattern, and morphological differences needs to be examined in western popula- tions, preferably on living or freshly collected specimens for accurate determination of color and pattern. We feel that the results of our study are preliminary and that a comprehensive study of variation throughout the entire range of C. horridus is needed. ACKNOWLEDGMENTS.— We thank C. J. McCoy, Carnegie Museum of Natural History (CM); William M. Palmer, North Carolina State Museum of Natural History (NCSM); and George R. Zug and W. Ronald Heyer, National Museum of Natural History, Smithsonian Institution (USNM), for allowing us to examine their specimens of Cro- talus horridus. We are also grateful to Madeleine Kennedy and Charles Crumly for their unselfish help with the analyses and interpretation. LITERATURE CITED Anderson, Paul. 1965. The Reptiles of Missouri. Univ. Missouri Press, Columbia. Conant, Roger. 1975. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Houghton Mifflin Co., Boston. Dixon, W. J., and M. B. Brown (editors). 1979. BMDP-79 Biomedical Compu- ter Programs P-Series. Univ. California Press, Berkeley. Dowling, Herndon G. 1951. A proposed standard system of counting ventrals in snakes. Br. J. Herpetol. 1:97-99. Gloyd, Howard K. 1940. The Rattlesnakes, Genera Sistrurus and Crotalus. Chi- cago Acad. Sci., Chicago. Klauber, Lawrence M. 1938. A statistical study of the rattlesnakes, V. Head dimensions. Occ. Pap. San Diego Soc. Nat. Hist. 4:1-53. . 1972. Rattlesnakes, Their Habits, Life Histories, and Influence on Mankind. 2nd ed. 2 vol. Univ. California Press, Berkeley. Variation in Crotalus horridus 73 -• o e v Z Z 2 Z Z I o o wvtvtZ >> 03 c c3 £ o u TD C 03 , — , cu o * — ' Ui 60 O a 03 a •— 03 ^3 2 m c ^_, a on U -sf IS OJ U -C <*H *-" o <*-H o c , "o 3 <+* 3 u X) .5 03 U > <+H o a c o l/» ♦ 1 s t/> <*> O ^d 03 1 GO 03 •.— • -C 74 Christopher W. Brown and Carl H. Ernst Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent. 1975. SPSS Statistical Package for the Social Sciences. 2nd ed. McGraw-Hill, Inc., New York. Peters, James A. 1964. Dictionary of Herpetology. Hafner Publ. Co., New York. Pisani, George R., Joseph T. Collins, and Stephen R. Edwards. 1973. A re-eval- uation of the subspecies of Crotalus horridus. Trans. Kans. Acad. Sci. 75:255-263. Smith, Philip W. 1961. The Amphibians and Reptiles of Illinois. 111. Nat. Hist. Surv. Bull. 28:1-298. Webb, Robert G. 1970. Reptiles of Oklahoma. Stovall Mus. Publ. 2, Univ. Oklahoma Press, Norman. Wright, Albert H., and A. A. Wright. 1957. Handbook of Snakes of the United States and Canada. Vol. II. Comstock Publ. Assoc, Ithaca, N.Y. Accepted 25 February 1985 Seasonal, Thermal, and Zonal Distribution of Ocean Sunfish, Mola mola (Linnaeus), off the North Carolina Coast David S. Lee North Carolina State Museum of Natural History, P.O. Box 27647, Raleigh, North Carolina 27611 ABSTRACT. — Most previous information on the ocean sunfish, Mola mola, has been derived from beached specimens and contributed little to our understanding of typical distributional patterns of the spe- cies. More than 60 encounters with Mola mola in North Carolina's offshore waters reveal that this fish is an epipelagic migrant, occurring in shallow water (10 to 40 fathoms in depth) commonly in the spring between mid-March and mid-June. In the fall it has been seen less frequently (mid-October through November), and the species is essen- tially absent in the winter. In spite of its cosmopolitan distribution, little information is avail- able concerning the natural history of the ocean sunfish, Mola mola (Linnaeus). This is particularly true in the southeastern United States, where nearly all records are of animals found awash in the surf. Because such records may reflect atypical patterns of movement and distribu- tion, observations on the .seasonal, thermal, and zonal distribution of Mola at sea are of interest. Between 1977 and 1986, I conducted 126 offshore trips for trye primary purpose of monitoring seasonal occur- rence and abundance of marine birds and mammals. During this period, however, I also incidentally observed other pelagic organisms (see Lee and Booth 1979, Lee and Palmer 1981). All but seven of the offshore survey trips departed from either Oregon Inlet or Hatteras Inlet, Dare County. Of the seven trips that did not, five were from Beaufort Inlet, Carteret County; one was from Wil- mington, New Hanover County; and one was from Virginia Beach, Vir- ginia. Each daylong outing lasted 10 to 1 1 hours and typically followed predesignated transects of 20 to 55 miles (32 to 88 km) from the point of departure and into the Gulf Stream. All of the Oregon Inlet and Hatter- as Inlet surveys extended to at least the 100-fathom contour, and many went several miles beyond the 1,000-fathom contour. Trips were made at all seasons, but monthly coverage was uneven (see Table 1). Ideally, water surface temperature, directional movement, and time and location of sightings were recorded for each sunfish observed. Data are not uni- form, however, because some charter boats lacked LORAN and other recording equipment, sea conditions necessitated abbreviated record Brimleyana No. 12:75-83, September 1986 75 76 David S. Lee -o o «- -S c « c -3 c ° I § a J2 O o o o > U > s- 3 C/5 o o OO O X sO «§ £ o. C/3 .1 £ 2 £ Cu 3 < o Mola Distribution off North Carolina Coast 77 keeping, and field effort was focused on seabirds. Furthermore, surveys of ocean sunfish from boats are difficult, because surface conditions and angle of view normally limit subsurface visibility. Variations in surface conditions from one trip to another make comparisons of trip-by-trip tallies meaningless. Nevertheless, cumulative records show patterns of zones of occurrence, as well as seasonal movement and abundance. In the North Atlantic M. mola ranges north to the Gulf of St. Lawrence, Newfoundland, southern Iceland, northern Norway, and the Kola Peninsula (Martin and Drewry 1978). It is not common in the tropics (Parin 1968). Information on seasonal movements is mostly con- jectural, suggesting passive transport by ocean currents or^ foraging while following passively drifting coelenterates and ctenophores (see Martin and Drewry 1978). The species is generally regarded as pelagic and solitary, but there are reports of M. mola moving in pairs or small groups (Whitley 1931, Smith 1965), and there are several records of summer occurrences, both of free-swimming and surf-washed individu- als, in bays such as Sandy Hook, New Jersey (Breder 1932), Isle of Wight Bay, Maryland (Schwartz 1964), and Monterey Bay, California (Myers and Wales 1930). Records of Mola along the southeast coast of North America are scarce, although farther north (e.g., New Jersey; Townsend 1918) it is fairly well established that these headfish occur regularly. Most north- ern records are of summer encounters. Brimley (1939) documented the occurrence of M. mola in North Carolina, providing information on one specimen and three other records; Anderson and Cupka (1973) compiled eight records for South Carolina. The species is known from waters off other southeastern states, including the Gulf of Mexico (Dawson 1965), but generally it appears on state faunal lists with no details of occurrence (e.g., Briggs 1958). Both Mola mola and Mola (formerly Masturus) lanceolata Lie- nard, the sharp-tailed mola, are found off the North Carolina coast (Brimley 1939, Funderburg and Eaton 1952). Although Dawson (1965) commented on the difficulty of identifying ocean sunfish at sea, several distinctive field characters separate these two fish. I was able to identify M. mola by its dull, nearly uniform color, the rounded dorsal or ventral fins, and the short blunt shape of its tail (which could be confirmed in 70% of sightings). Because nearly all fish seen were considerably greater than 1 meter total length, I assumed most were adults. Most sunfish were sighted while they were swimming about 0.5 to 1.5 m below the surface. In their "sunning" behavior the fish's sides were always below the surface. Usually the dorsal fin, and occasionally the ventral fin, projected above the surface. Projecting fins were normally held at angles of 45 to 70 degrees and were constantly undulating. This allowed sunfish to be sighted from distances of more than 100 m under calm conditions. Observed fish whose fins did not project above the surface could not be detected for more than 20 to 25 meters from the boat. Normally the fish did not dive at the approach of the boat, but 78 David S. Lee simply maneuvered out of its way. They sounded only if the boat was on a collision course. Boat captains say the fish are rarely if ever hit by their boats. Sunfish were seen on calm days, days with considerable swells, and days when small white caps were prevalent, although reduced visibility made comparative counts useless. When seas were quite rough (20+ mph winds, high swells, and extensive white caps), no fish were found; but under these conditions we occasionally sighted marine turtles, sharks, and porpoises. I suspect the sunfish were then swimming deeper, and our failure to see them was not simply a result of the poor subsur- face visibility. Information pooled from 60 sightings of Mola mola personally obtained and other available records from the North Carolina coast suggest that the species is not randomly distributed by season or loca- tion. Although field effort was not uniform, the records obtained are informative, in that the majority are from areas and seasons having min- imal opportunities for observation (see Table 1). Season: Mola mola is essentially absent off the North Carolina coast during the winter (see Table 1). Although I have made few winter trips (N = 20), I have no reason to assume ocean sunfish occur regularly at this season, for boat captains and others also have not encountered them in the winter. The earliest spring record is for 16 March, and the earliest fall record is for 17 October. The species is most commonly seen in the spring. Surprisingly, the fish do not occur regularly in our waters during summer. Boat captains say they occasionally see ocean sunfish in the summer, but some of these could be the more tropical M. lanceola- ta. Interestingly, a large part of our survey time during summer was spent in the Gulf Stream, where M. lanceolata could be expected, but none was verified. In the summer of 1985 I personally encountered M. mola eight times on only 4 of 15 offshore trips, all between 17 and 29 August, a period when relatively calm water usually provides optimum subsurface visibility. No other summer records are available in spite of rather extensive offshore surveys in this season. The fact that only three M. mola were encountered in the fall (17 October through 20 November) suggests a different fall migration route, or perhaps a seasonal absence of surface "sunning" behavior. The three dated North Carolina records provided by Brimley (1939) are all for May. Anderson and Cupka (1973) also reported Mola from April (2) and May (1); but their other records were from December (2), January (1), and February (2), sug- gesting winter occurrence in South Carolina (see below). Location: This fish was seldom seen in areas of deep water (> 100 fathoms); most occurred in an offshore zone between 10 and 40 fathoms deep (x - 28.19 fathoms). Most were seen more than 10 miles from shore, although one fish was seen while the survey boat was still in sight of land (19 April 1980). Only six records were in water 40 to 100 Mola Distribution off North Carolina Coast 79 fathoms deep, and one December record is from 500 fathoms. Except for the December fish, individuals were not encountered beyond the inner edge of the continental shelf (100 fathoms), although nearly half of our survey time was spent in these deeper waters. Additionally, Charles Manooch, National Marine Fisheries Laboratory, Beaufort, informed me that all of the 15 Mola seen by him were between 20 and 30 miles from shore and in water 17 to 25 fathoms deep. Off South Carolina, ocean sunfish (species not determined) have been reported over water about 42 m (23 fathoms) deep (Anderson and Cupka 1973). Interestingly, Lee and Palmer (1980) documented the regular ocurrence of leatherback turtles, Dermochelys coriacea, another reputed coelen- terate feeder, to be restricted, or nearly so, to shallow waters inshore of the 100-fathom contour. Manooch reported an adult M. mola in Core Sound (Harkers Island, fall date not recorded), and the site of Brimley's (1939) Swans- boro record is Bogue Sound. Although Myers and Wales (1930) noted that young individuals were of regular occurrence during the summer in Monterey Bay, California, I am not aware of any records from estuarine bays. There are no reports of Mola, for example, in the Chesapeake Bay. However, Steve Ross (pers. comm.) captured a single adult from near the mouth (< 20 ppt) of the Neuse River near Long Creek on 16 May 1980 in a gill net. This is the only truly estuarine occurrence of which I am aware. Water Temperature'. Ocean surface temperatures were recorded for 20 of my 60 North Carolina sightings at sea, and temperature approxi- mations (±2 °C) are possible for 13 others based on temperatures recorded at other locations near the sighting. The coldest water in which I encountered M. mola was 6.8 °C on 16 March 1984, which was also the date of the earliest spring record. The warmest water was 29.4 °C on 13 June 1979, the date of the latest spring sighting. Most encounters were at temperatures between 10 and 18 °C. On all dates a surface temperature gradient was recorded, with coolest waters generally closest to land and warmest waters within the Gulf Stream. Seasonal and ther- mal distributions (Fig. 1) suggest that, although maximum and min- imum temperatures may be critical, these fish are not simply moving into deeper, warmer waters during cool periods, or into cooler inshore waters during warm seasons. Similar findings were reported for several species of marine turtles off the North Carolina coast (Lee and Palmer 1980). Time of Day for "Sunning": Surface "sunning" behavior was noted for most periods of the day, the earliest at 0732 EST and the latest at 1432. Additionally, several sunfish were seen in "mid- to late after- noon," but exact times were not recorded. Miscellaneous: All sunfish observed were solitary, although on sev- eral occasions individuals were found within half a mile of each other. 80 David S. Lee Except for the one December record, none of the Mola I saw were known to be associated with jellyfish or other fishes, nor were any asso- ciated with sargassum beds, floating boards, or other objects. Manooch (pers. comm.), however, reported diving in water 20 to 30 miles off Beaufort and seeing one M. mola associated with a large number of "sea nettle type" jellyfish on 12 March 1976. Probably coelentrates are not easily seen from above the surface. At any rate, none of the coelenter- ates or ctenophores that could offer a prey base were seen regularly. The only jellyfish typically seen on any of the surveys was Physalia, and it invariably was in the Gulf Stream, offshore of the areas inhabited by Mola. Likewise ocean sunfish were not found along "tide lines," current edges, sites of local upwellings, or other areas where many marine organ- isms tend to congegrate.. Migration and Movement: In that Mola mola is well known north of North Carolina in summer and south of the state in winter (Anderson and Cupka 1973) and is rare or absent from North Carolina waters during these periods, most individuals seen off our coast are probably migrants. All spring individuals whose orientation was recorded (about one-half of the total) were swimming north. Their lack of apparent for- ward movement may be deceptive; when the boat was in motion (10 to 18 knots), the fish appeared to remain in one area. On several occasions, however, sunfish were watched moving past and out of sight of our idling boat (in one case the boat was broken down) in a short time period. As previously implied, movement was within a wide band gen- erally over the 10- to 45-fathom contour. The records from mid to late August 1985 are interesting in that this was the only summer in 10 years of offshore study that I have seen ocean sunfish. Although late August at first appears early for "fall" migration, I should point out that many southbound sea birds appear in North Carolina offshore waters at this time. Furthermore, various migratory sport fish locally appear or reappear in this same time period. Nevertheless, southward fall migration of M. mola would appear to occur primarily in October and November, with movements perhaps starting as early as late August in some years. Most ocean sunfish were noted between mid-April and mid-May when about 80% of the total sightings were compiled. It may be that south of the Hatteras area migration occurs farther offshore. This is suggested by the few sightings made off Beaufort (5 in 175 trips made by Manooch, pers. comm.; none in 25 trips made by Wayne Irvin, pers. comm., or me). In this area, comparable water zones and the inner edge of the Gulf Stream are much farther from land than off the northern Outer Banks where most of my surveys were conducted. DISCUSSION The ocean sunfish, Mola mola, is best regarded as an epipelagic migrant in North Carolina's offshore waters. In the spring it can be Mola Distribution off North Carolina Coast 81 °c 30-i 28- 26- 24- 22- 20- 18- 16- 14 - 12- * • + ■ Vv Gulf Stream v / Shelf Break 10- (ca 100 fathoms) • 8 - -___ Near-Shore 6- 1 1 1 1 i i i i i i i i JAN FEB MAR APR MAY jUN JUL AUG SEP OCT, NOV DEC N (4) (8) (3) (3) (1) (1) Fig. 1. Thermal distribution of Mola mola compared to surface temperature gradients. Ranges and means of temperatures taken in immediate vicinity of Mola (N = 20). Average monthly sea-surface temperatures for three areas of the North Carolina continental shelf north of Cape Hatteras (from Newton et al. 1971). quite common. On 18 and 19 April 1980, 15 were seen each day despite sea surface conditions that offered less than maximum visibility. Five were counted on 14 May 1981, but on all other days only one or two verifiable M. mola were seen per trip. Sunfish actually were more com- mon than Table 1 indicates. I often observed two to three times as many individuals as reported, but these sightings were not recorded, either because specific identity could not be confirmed or because other survey priorities were more urgent at the moment. The dearth of M. mola sightings during fall is difficult to explain, especially since Anderson and Cupka (1973) stated that a boat captain reported at least 30 molas (species undetermined) in late autumn of 1970 and 1971 off South Carolina. Local seasonality of occurrence of Mola based on beach stranded specimens may be misleading. Along the Atlantic coast injured, sick, or dead fish could be displaced long distances by the Labrador Current, long shore current, or Gulf Stream. The fact that six of the seven M. lanceolata from North Carolina (Brimley 1939, Funderburg and Eaton 82 David S. Lee 1952, NCSM records) are winter records seems contradictory to the known habits of this tropical species. Such occurrences should not be interpreted to mean that they are a regular part of the offshore fauna in winter. The same point could be argued for five of the eight Mola reported from South Carolina beaches in December, January, and Feb- ruary (Anderson and Cupka 1973). In both cases individuals may have been numbed by cool sea conditions and transported northward from, to date, undetermined "wintering areas." ACKNOWLEDGMENTS.— Steven P. Platania and Mary Kay Clark, both of the North Carolina State Museum, assisted with many of the offshore surveys. Charles S. Manooch III, National Marine Fisher- ies Service, Beaufort Laboratory, and E. Wayne Irvin, NCSM, supplied supplemental data from their trips off Beaufort. George Burgess, Flor- ida State Museum, and Steve Ross, NCSM, both assisted in locating several pertinent literature sources including local records, and reviewed the contents of this note. John E. Cooper provided useful comments on the manuscript. The study was financed in part by contract # 92375- 1130-621-16, U.S. Fish and Wildlife Service Laboratory, Slidell, Louisiana. LITERATURE CITED Anderson, William D. Jr., and D. M. Cupka. 1973. Records of the ocean sunfish, Mola mola, from the beaches of South Carolina and adjacent waters. Chesapeake Sci. 14(4):295-298. Breder, Charles M., Jr. 1932. Fish notes for 1931 and 1932 from Sandy Hook Bay. Copeia 1932(4): 180. Briggs, J. C. 1958. A list of Florida fishes and their distribution. Bull. Fla. State Mus. 2:223-318. Brimley, H. H. 1939. The ocean sun-fishes on the North Carolina coast. The pointed-tailed Masturus lanceolatus and the round-tailed Mola mola. J. Elisha Mitchell Sci. Soc. 15(2):295-303. Dawson, C. E. 1965. Records of two headfishes (Family Molidae) from the north-central Gulf of Mexico. Proc. La. Acad. Sci. 28:86-89. Funderburg, J. B., Jr., and T. H. Eaton. 1952. A new record of the pointed- tailed ocean sunfish, Masturus lanceolatus, from North Carolina. Copeia 1952(3):200. Lee, D. S., and J. Booth. 1979. Seasonal distribution of offshore and pelagic birds in North Carolina waters. Am. Birds 33(5):7 15-721. , and W. M. Palmer. 1981. Records of leatherback turtles, Der- mochelys coriacea (Linnaeus) and other marine turtles in North Carolina waters. Brimleyana 5:95-106. Martin, F. D., and G. E. Drewry. 1978. Development of fishes of the mid- Atlantic bight: an atlas of eggs, larval and juvenile stages. Volume VI. U.S. Fish and Wildlife Service FWS/OBS-78/ 12. Mola Distribution off North Carolina Coast 83 Myers, G. S., and J. H. Wales. 1930. On the occurrence and habits of ocean sunfish (Mola mola) in Monteray Bay, California. Copeia 1930(1): 1 1. Newton, J. G., D. H. Pilkey, and J. O. Blawton. 1971. An Oceanographic Atlas of the Carolina Continental Margin. N.C. Dept. of Conservation and Development. Parin, N. V. 1968. Ikhtiofauna Okeanskoi Epipelagiali [Ichthyofauna of the Epipelagic Zone]. Akademiia Nauk SSSR. Institut Okeanologii, Moscow. (Translated by U.S. Department of Interior and National Science Founda- tion, Washington, D.C., 1970.) Schwartz, F. J. 1964. Fishes of the Isle of Wight and Assawoman bays near Ocean City, Maryland. Chesapeake Sci. 5(4): 172-193. Smith, J. L. B. 1965. The Sea Fishes of Southern Africa. 5th ed. Central News Agency, Ltd., South Africa. Townsend, C. H. 1918. The great ocean sunfish. Bull. N.Y. Zool. Soc. 21:1677-1679. Whitley, G. P. 1931. Studies in Ichthyology. No. 4. Rec. Aust. Mus. Syd. 18(3):96-133. Accepted 15 November 1985 84 ATLAS OF NORTH AMERICAN FRESHWATER FISHES by D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, J. R. Stauffer, Jr., and many collaborators This timely book provides accounts for all 777 species of fish known to occur in fresh waters in the United States and Canada. Each account gives a distribution map and illustration of the species, along with information on systematics, distribution, habitat, abundance, size, and general biology. ". . . represents the most important contribution to freshwater fishes of this continent since Jordan and Evermann's 'Fishes of North and Middle America' over 80 years ago." — Southeastern Fishes Coun- cil Proceedings. 1980 825 pages Indexed Softbound ISBN 0-917134-03-6 Price: $25, postpaid. North Carolina residents add 4l/2% sales tax. Please make checks payable in U. S. currency to NCDA Museum Extension Fund. Send to FISH ATLAS, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. ATLAS OF NORTH AMERICAN FRESHWATER FISHES 1983 SUPPLEMENT by D. S. Lee, S. P. Platania, and G. H. Burgess The 1983 supplement to the 1980 Atlas of North American Fresh- water Fishes treats the freshwater ichthyofauna of the Greater Antilles. In addition to this bound supplement, there are 19 accounts, mostly species not described in 1980, in looseleaf form to be added to the 1980 volume. Illustrated by Renaldo Kuhler. 1983 67 pages Indexed Softbound Price: $5, postpaid. North Carolina residents add 4!/2% sales tax. Please make checks payable in U. S. currency to NCDA Museum Extension Fund. Send to FISH ATLAS, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. A Late Quaternary Herpetofauna from Saltville, Virginia J. Alan Holman The Museum, Michigan State University, East Lansing, Michigan 48824 AND Jerry N. McDonald 715 Saratoga Avenue, Newark, Ohio 43055 ABSTRACT. — The late Quaternary herpetofauna from Saltville, Vir- ginia, consists of at least two salamanders, two anurans, two turtles, and four snakes; all are forms that can be found living in the area today. The fossil herpetofauna originated from three 14C dated strati- graphic units. Based on the presence of all 10 taxa of the herpetofauna in Units W2 (lower) and W3, it is reasonable to conclude that this fauna has been in place for the last 13,500 to 15,000 years. Because the most northern area where all members of the Saltville herpetofauna may be found living together today is in extreme northeastern Penn- sylvania, the herpetofauna is clearly not a "Boreal" one. Moreover, Boreal temperatures, as we know them today, would not provide enough warm days for the eggs of Chelydra serpentina, Chrysemys picta, or Elaphe cf. E. obsoleta to hatch. The late Quaternary fluvial and lentic sediments of the Saltville Valley in Virginia have yielded the remains of large mammals for more than two centuries (Jefferson 1787, Peterson 1917, Boyd 1952, Ray et al. 1967, McDonald and Bartlett 1983). Most of these remains were found during construction activities related to agriculture or the production of salt. The first purely scientific excavation in search of late Quaternary vertebrates at this locality was conducted jointly by Virginia Polytechnic Institute (VPI) and the Smithsonian Institution (SI) in 1966 and 1967. In 1978 and 1981 Charles Bartlett, Jr., performed salvage excavations at several locations in the valley for the Town of Saltville, and in October 1980 Bartlett and J. McDonald began controlled excavations in the valley. In 1982 McDonald initiated the Saltville Project, a multidisciplinary investi- gation of the late Quaternary history of Saltville Valley that included the collaboration of several specialists from different institutions in eastern North America. Late Quaternary deposits in the Saltville Valley have been shown to span some 27,000 years, including a continuous record for approximately the last 15,000 years (McDonald 1984, 1985a), making this locality unusually useful for the documentation of Brimleyana No. 12:85-100, September 1986 85 86 J. Alan Holman and Jerry N. McDonald environmental change in the middle Appalachian region through the late Wisconsin and Holocene. The first known herptile specimen to be collected at Saltville was a partial limb bone of an anuran (fam., gen. et sp. indet.) collected on 1 1 August 1966, by the VPI-SI field crew (Catalog and field notes, 1966, VPI-Smithsonian Saltville Expedition). Bartlett found the second specimen — a costal bone of the Painted Turtle, Chrysemys picta (USNM 404721)— on 30 October 1978 (C. S. Bartlett, Jr., field notes, 30 October 1978). The 1980-1984 Radford University excavations recovered numer- ous herptile specimens by wet screening the finer fluvial sediments and closely examining thinly sliced lentic deposits of clay and silt. Vertical and horizontal provenience and matrix data for specimens have also been collected since 1980, which allows differentiation of faunules and inferences about faunal change (or the absence of change) over time. Here, we describe the generically and specifically identifiable herp- tile material collected at Saltville through 1984, including the division of this material into three radiocarbon-dated faunules. In addition, we dis- cuss the sampling function of the various depositional processes and comment on the paleoecological implications of these faunules. The herptile material reported here is the first to be described from the Salt- ville locality, and is also the first to be described from a stratified subae- rial, hydraulically deposited site in the middle Appalachians. This is, therefore, a contribution to the controlled chronostratigraphy of late Quaternary herptiles in this region, a contribution free of the collecting and preservation biases characteristic of herpetofaunas from karst or karstlike features in the middle Appalachians. STUDY AREA Saltville Valley lies some 525 m above sea level in the Valley and Ridge Physiographic Province in southwest Virginia (Fig. 1). The floor of this small valley slopes gradually to the north, converging on a water gap that leads to the nearby North Fork of the Holston River. The valley is bordered on the northeast and southeast by foothills of Walker Mountain, and on the northwest by low limestone hills. The herptiles described in this paper came from four sites on the valley bottom (Fig. 1). Most specimens were collected at SV-1 (the "musk ox" site: 36°52'19"N, 81°46'24"W), located near the south- west end of "The Flat" (McDonald and Bartlett 1983). Six specimens came from SV-2 (the "drug store" site: 36°52'52"N, 81°45'48"W), and one came from CSB-2A (36°52'29"N, 81°45'51"W). The anu- ran bone collected by VPI-SI in 1966 came from SI-1 (36°52'36"N, 81°46'01"W). SV-1 and CSB-2A are on the Glade Spring quadrangle, and SV-2 and SI-1 are on the Saltville quadrangle, USGS 7.5' series. Saltville Valley lies upon the Mississippian Maccrady Formation, a variable sequence of shales, siltstones, limestones, and dolomites con- taining substantial quantities of gypsum, anhydrite, and halite (Cooper Late Quaternary Herpetofauna 87 80° w SALTVILLE / ^ © Ul ro o jk y5 — 03 — c/i 5fl 03 03 03 03 £ S u u X 3C i so r- m rt r» 0 -* Tt m 0 rf CS ^~ S -<* . 1 •5 00 10 O Z CO D z CO D 0 z ^ t-« <* 0 s Q CO U z CO 3 O "5 D 3 2 z CO D -sl C ■8 •3- ^5 . 5 0 5! O . 1 1 cq m 00' m 1 O r-' "* m £T r-- ^ >n m O On* rn 0 •2 £5 co 2 1 fN CO 2 Z ^ 0 c r- z t-^ r- ti- Tf .^ Tf r-i "i- r^ '5 -Si "5 0 Z CO CO •5 > 0 0 z CO CO 0, 3 0 s z z CO D 03 c O 1 D 3 O 5h CO D ex <*> 0 <* s z -Si a 0 z 2! «<5 a, .a 2! 0 S3 .2 5 1 00 2 ^1 CO c a 1 6 D 03 g 0 D «5 g ; >^ -5 J3 "O u 03 O 0 03 s CU 00 cC V CQ T3 "5 > CQ 3 1> c § 03 O X> O SO m a, •B 2 > CX o ■s D J r o o ** ^ c .£ — 1 c <*-> 03 M •§ 8 « o «i p ^ Cu 00 ON O 0 "^ OS ex "* O CJ OO ON O 1c CX- O CS u 2 .SP (N "o3 C 0 T3 C 03 a 03 CB 0 O s 00 03 u '£ ^— * jr < u H 03 ^ 0 Late Quaternary Herpetofauna 9 1 fish and mammal remains. Most of the fish and herptile remains from Unit W3 are in good condition, suggesting that they have been subjected to relatively little fluvial abrasion, whereas the mammal remains range from unabraded to heavily abraded. Four herptile specimens were found in the lower several centimeters of Unit W2, associated with large numbers of mollusk, fish, and mammal remains. These remains do not show evidence of abrasion. Two herptile specimens were found in the lowest 5 cm of Unit Wl, a humus-rich mud that has preserved fluid- produced whorls at its contact with Unit W2. Herptiles have not been found to date at SV-1 in units PI, W4, the upper part of W2, H2, or HI (Fig. 2). Late in August 1983, the foundation of the old Olin Mathieson Chemical Corporation's company store was demolished and the area excavated with heavy machinery in preparation for construction of a new drug store. This excavation (site SV-2) exposed only artificial fill or otherwise disturbed sediments around most of the periphery and across the bottom, but a small section of undisturbed natural sediment was exposed along the southeast wall. Here, 225 cm of artificial fill was underlain by 13 cm of what appeared to be natural lacustrine clay, although this stratum did contain a few very small (< 3 mm) intrusive brick fragments. Beneath the clay was a layer of alluvium, consisting of medium sand to very fine gravel, numerous small pieces of wood, and bones and teeth. No intrusive material was found. This alluvium was separated from the overlying clay by a distinct boundary, and it lay unconformably upon well-scoured bedrock, indicating that it was depos- ited while the valley was still being drained by vigorously flowing water. This site is low and near the water gap leading to the Holston River; it is therefore unlikely that the alluvium could have been deposited after Lake Totten had formed unless the lake drained periodically. No radio- carbon date was obtained for this deposit, but we tentatively identify it as a member of Unit W3. Six herptile specimens were found in a 5- gallon (19-1) sample of this unit collected 3 September 1983. Also included in this sample was an abraded fragment of a mastodon (Mammut americanum) tooth and the unabraded crown of a superior molar of a cervid (Sangamona or Odocoileus). CSB-2A was excavated 28 and 30 October 1978, under the direc- tion of Charles S. Bartlett, Jr., as part of an effort to salvage paleonto- logical and archeological resources prior to construction of bleachers at the Saltville softball park. Bartlett reported finding many rounded fragments of large mammal bones and teeth, along with one fragment of turtle bone, in a "pebble zone" that we tentatively assign to Unit W4 (C. S. Bartlett, Jr., field notes, 28 and 30 October 1978; pers. comm.). The turtle bone (USNM 404721) does not, however, show signs of abrasion. Rather, its condition is similar to other remains found in units W2 and W3. Based on the condition of USNM 404721, we suspect that it might have come from the bottom, or from near the bottom, of Unit W2 instead of from within Unit W4, which typically contains noticeably abraded remains of large mammals only. Alternatively, Bartlett's "peb- ble zone" might have included, or consisted entirely of, Unit W3. 92 J. Alan Holman and Jerry N. McDonald SYSTEMATIC PALEONTOLOGY The classification used here follows Dowling and Duellman (1978). The common names used follow Collins et al. (1978). Ranges and notes on modern species follow Conant (1975) or personal observations by J. A. Holman. Numbers are those of the Department of Paleobiology, Division of Vertebrate Paleontology, U.S. National Museum, Washing- ton, D.C. (USNM). All measurements are in millimeters. Class Amphibia Order Caudata Family Cryptobranchidae Cryptobranchus alleganiensis (Daudin), Hellbender Material — Trunk vertebra: USNM 404722 (Fig. 3), from Unit W2. Remarks. — This vertebra is indistinguishable from those of modern Cryptobranchus alleganiensis. The Saltville fossil may be separated from the extinct species C. guildayi Holman of the late Kansan of Trout Cave, West Virginia, on the basis of vertebral ratios. The ratio of the greatest length through the zygapophyses divided into the greatest width through the posterior zygapophyses is .65 in the Saltville C. alleganien- sis and .56-. 65, mean .602, in 18 specimens of modern C. alleganiensis. This ratio was .69 in the single available vertebra of C. guildayi. The Hellbender occurs in the area today, and is found usually in rivers and large streams where shelter is available in the form of large rocks, snags, or debris. Family Salamandridae Notophthalmus cf. N. viridescens (Rafinesque), Eastern Newt Material. — Five trunk vertebrae: USNM 404723, from Unit W3, SV-1; USNM 404724, from Unit W3, SV-1; USNM 404725, from Unit W3, SV-2; USNM 404726, from Unit W3, SV-1; and USNM 404727 (Fig. 4), from Unit W3, SV-1. One femur: USNM 404728, from Unit W3, SV-2. One humerus: USNM 404729, from Unit W3, SV-1. Remarks. — The vertebrae of the genus Notophthalmus have a quite characteristic high, posteriorly thickened, posteriorly divided neural spine. These vertebrae appear to be identical to those of the Eastern Newt, Notophthalmus viridescens. The femur and the humerus also show no differences from the modern species. The Eastern Newt occurs in the area today, and the habitat of the aquatic stage is ponds, lakes, marshes, ditches, and other quiet bodies of unpolluted water. The ter- restrial stage usually hides under objects in forested areas, but at times individuals may be seen walking about in the open. We are unable to tell on the basis of osteological material whether the fossils represent the aquatic or the terrestrial stage. Late Quaternary Herpetofauna 93 Fig. 3. Trunk vertebra of Cryptobranchus alleganiensis (Daudin) (USNM 404722) from Unit W2. Upper left, ventral; upper right, dorsal; lower left, poste- rior; lower right, lateral. Line equals 5 mm and applies to all drawings. Order Anura Family Bufonidae Bufo woodhousei fowleri Hinckley, Fowler's Toad Material. — Left ilium: USNM 404730, from near (ca. 4 cm above) base of Unit Wl, SV-1. Two right ilia: USNM 404731, from Unit W2, SV-1; USNM 404732, from Unit W3, SV-2. Two tibiofibulae: USNM 404733, from Unit W3, SV-1; USNM 404734, from Unit W3, SV-1. Parasphenoid: USNM 404735, from Unit W2, SV-1. Remarks. — Holman (1967) and Wilson (1975) discussed characters of the ilial prominence that allow separation of Bufo woodhousei fowl- eri from the morphologically similar Bufo americanus. Bufo w. fowleri is easily separated from its western counterpart B. w. woodhousei on the basis of the much higher dorsal protuberance in the latter subspecies. Bufo w. fowleri occurs in the area today, and occurs chiefly in sandy areas around shores of lakes, or in river valleys. Family Ranidae Rana pipiens group, sp. indet. Material. — Right ilium: USNM 404736 (Fig. 5), from Unit W3, SV- 2. Two left humeri: USNM 404737, from Unit W3, SV-1; USNM 94 J. Alan Holman and Jerry N. McDonald 404738, from Unit W3, SV-1. Right humerus: USNM 404739, from Unit W3, SV-1. Remarks. — The small right ilium has a smooth vastus prominence and has the posterodorsal border of its ilial crest sloping gently into the dorsal acetabular expansion as in species of the Rana pipiens group such as R. pipiens, R. blairi, R. berlandieri, and R. utricularia. But we are unable to determine which of these species the ilium represents. The Southern Leopard Frog, Rana utricularia, occurs in the area today. This frog inhabits a wide variety of aquatic situations, and may move quite a distance from the water in summer where growing plants pro- vide shade and shelter. Class Reptilia Order Testudines Family Chelydridae Chelydra serpentina (Linnaeus), Snapping Turtle Material. — Partial nuchal bone: USNM 404740, from lowest part of Unit W2, SV-1. Scapulocoracoid: USNM 404741, from ca. 5 cm above base of Unit Wl, SV-1. Remarks. — These very characteristic bones represent a small Snap- ping Turtle. Preston (1979) gave some characteristics of chelydrid shell bones that allow identification of fragments. This species occurs in the area today. Snapping Turtles inhabit almost any body of water that is relatively slow moving and permanent (pers. observ.). Family Testudinidae Chrysemys picta Schneider, Painted Turtle Material. — Third right costal: USNM 404721 (Fig. 6), from Unit W2 (?) or W4 (?), CSB-2A. Remarks. — The smooth nature of the dorsal surface of this shell bone, and the position of the impression of the seam for the second epidermal shield, is diagnostic in Chrysemys picta. This turtle occurs in the area today and is an inhabitant of quiet, vegetation-choked bodies of water (pers. observ.). Order Squamata Family Colubridae Elaphe cf. E. obsoleta, Rat Snake Material. — Trunk vertebra: USNM 404742, from Unit W3, SV-2. Remarks. — Auffenberg (1963) gave vertebral characters of Elaphe obsoleta. The above trunk vertebra is from a moderately large speci- men. This snake occurs in the area today, and is a semiarboreal form that favors wooded areas and woodland edges (pers. observ.). Nerodia sipedon (Linnaeus), Northern Water Snake Material. — Trunk vertebra: USNM 404743, from Unit W3, SV-2. Late Quaternary Herpetofauna 95 Fig. 4. Trunk vertebra of Notophthalmus cf. N. viridescens (Rafinesque) (USNM 404727) from Unit W3. Upper left, ventral; upper right, dorsal; middle left, posterior; middle right, anterior; bottom, lateral. Line equals 2 mm and applies to all drawings. Remarks. — Holman (1967) gave vertebral characters that distinguish this species from others in the genus. The Northern Water Snake occurs in the area today and is found in many aquatic situations. Large popu- lations are often to be found where protective shelters occur near aqua- tic situations (pers. observ.). Storeria sp., Brown Snake or Red-bellied Snake Material — Trunk vertebra: USNM 404744,a from Unit W3, SV-1. Remarks. — Holman and Winkler (in press) discuss the separation of isolated vertebrae of the closely related genera Storeria and Virginia. We are unable to separate the vertebrae of the two species of Storeria; 96 J. Alan Holman and Jerry N. McDonald both S. dekayi and S. occipitomaculata occur in the Saltville area today. Thamnophis sp., Gartersnake or Ribbonsnake Material. — Trunk vertebra: USNM 404745, from Unit W3, SV-1. Remarks. — Brattstrom (1967) showed that the vertebrae of Tham- nophis are more elongate than those of the related genus Nerodia. It is almost impossible to separate isolated vertebrae of the two species of Thamnophis (T. sauritus and T sir talis) that occur in the vicinity of Saltville today. DISCUSSION The known herptile fauna from Saltville has been divided into three faunules on the basis of the depositional units from which the remains were recovered (Table 1). The taxonomic composition and chronology of these faunules can provide information about the duration of resi- dency of the taxa, the depositional environment in which each was best sampled, and the microhabitat of the respective taxa. Unit W3, the sorted stream channel bed load deposit found at SV-1 and SV-2, contained seven taxa including all identified specimens of Notophthalmus cf. viridescens, Rana pipiens group, Nerodia sipedon, Thamnophis sp., Elaphe cf. E. obsoleta, and Storeria sp. Only Bufo woodhousei fowleri is found in W3 and other depositional units. The stratigraphic nature of Unit W3 — silts, sands, and fine gravels, ranging from well sorted and laminated deposits to "unsorted" masses (perhaps mixed biogenically, as by trampling by large mammals) — indicates that the member deposits were laid down by moderately to slowly moving water, perhaps through several cycles of rise and fall. Fluctuations in stream stage would have permitted periodic integration of the remains of terrestrial vertebrates into the stream bed load, especially those taxa that inhabited or periodically used the riparian zone. This might explain the presence of terrestrial taxa, including most of the snakes, in the fluvial deposits. The large amount of woody plant remains of uniform size (< 50 mm) in Unit W3 at SV-2 strongly suggests fluvial sorting of "sediments" of terrestrial origin. Alternatively, semi-aquatic or avian predators or scavengers could have dropped the remains of terrestrial prey in or near the stream during feeding. The possibility that large mammals might have mixed units W2 and W3 at SV-1 while watering or feeding has been considered. However, in view of the fact that the composition of the herptile samples in Unit W3 at SV-1 and SV-2 is remarkably similar and that the composition of W2 and W3 at SV-1 are generally different, mixing of these two deposits must be considered unsubstantiated at present. The herptiles of Unit W3 may, therefore, be taken to represent a sampling of the Saltville Valley lotic and riparian herpetofauna as of ca. 14,500 to 14,000 B.P. Late Quaternary Herpetofauna 97 Fig. 5. Right ilium in lateral view of Rana pipiens group frog (USNM 404736) from Unit W3. Line equals 5 mm. frCTBBSjWgSM ■\'y-:M:Z {g^T»y/roWtoWa;y«^ Fig. 6. Third right costal in dorsal view of Chrysemys picta Schneider (USNM 404721) from Unit W2 (?) or W4 (?). Line equals 10 mm. Deposits associated with the early history of Lake Totten (ca. 14,000 to 12,000 B.P.) include ostracods, pelecypods, gastropods, fish, and mammal remains as well as those of Cryptobranchus alleganiensis, Bufo woodhousei fowled, Chelydra serpentina, and (?) Chrysemys picta. Most of the aquatic fauna of Lake Totten probably was residual from that of the Saltville River, although the change in local hydrology caused a shift in the dominant taxa and altered the collecting bias of the depositional environment. The kinds of turtles represented are compati- ble with the postulated lake environment, and the remains of Fowler's Toad could easily have been deposited following death in or alongside the lake. The environmental implication of the Hellbender is more equivocal; it could have occupied a spring-fed brook entering Lake Tot- ten near SV-1 (as does a small stream today), or it could represent feed- ing residue dropped by a predator or scavenger. The middle and upper parts of Unit W2 yield very few faunal remains. The reasons for this are unclear, but could include any or all of the following: change in water quality, water level fluctuation, and infilling of Lake Totten near SV-1. 98 J. Alan Holman and Jerry N. McDonald Fig. 7. Map showing the most northern area (crosshatched) where all members of the Saltville herpetofauna (dot) may be found living together today. Unit Wl, lying astride the Wisconsin-Holocene boundary (ca. 10,500 to 10,000 B.P.), consists of an organic-rich mud at SV-1 that contains remains of Bufo woodhousei fowleri and Chelydra serpentina. The boundary between W2 and Wl was distinct below where USNM 404730 and USNM 404741 were found, which suggests that these iso- lated remains were transported with the mud when — or deposited after — it moved, rather than being moved upward from the underlying lake deposit by bioturbation. Conceivably, the mud encompassing these specimens was a littoral deposit displaced by the downslope movement Late Quaternary Herpetofauna 99 of a larger wasting mass from the adjacent hills. Because only two iso- lated bones were found, it is unlikely that the mud slide killed and bur- ied the individuals from which these specimens came. Using this reason- ing, both Fowler's Toad and the Snapping Turtle appear to have been present throughout the first 4,000 years of Lake Totten's history. All of the herptile taxa present in the Saltville faunules can be found living in this area today. Based upon the presence of all 10 species in the herpetofauna in units W2 (lower) and W3, it is reasonable to conclude that this fauna has been in place for at least the last 13,500 to 15,000 years. Differences in the taxonomic composition of the faunules are probably attributable to microhabitat changes associated with hydro- logic changes in the valley and to different sampling biases of the var- ious depositional processes represented. The most northern area where all members of the Saltville herpeto- fauna may be found living together today is in extreme northeastern Pennsylvania (Fig. 7) (Conant 1975: maps 3, 22, 99, 116, 119, 127, 149, 188, 198, 265, and 303). The Saltville herpetofauna, therefore, clearly is not a "Boreal" herpetofauna. Boreal temperatures as we know them today would not provide enough warm days for the eggs of Chelydra serpentina, Chrysemys picta, and Elaphe cf. E. obsoleta to hatch. The summers of ca. 15,000 to 14,000 B.P., and those since, must have been warm enough for the eggs of these species to hatch (cf. Stuart 1979). ACKNOWLEDGMENTS.— The authors thank Clayton E. Ray for reviewing an earlier draft of this paper, and Rosemarie Attilio for draw- ing Figures 3, 4, 5, and 6. Support for this research was provided by the Town of Saltville and The National Geographic Society (grants 2512 and 2880). LITERATURE CITED Auffenburg, Walter. 1963. The fossil snakes of Florida. Tulane Stud. Zool. 10 (3):131-216. Boyd, Julian P. (editor). 1952. The Papers of Thomas Jefferson. Vol. 6. Prince- ton Univ. Press, Princeton. Brattstrom, Bayard H. 1967. A succession of Pliocene and Pleistocene snake faunas from the High Plains. Copeia 1967(1): 188-202. Collins, Joseph T., J. H. Huheey, J. L. Knight, and H. M. Smith. 1978. Standard Common and Scientific Names for North American Amphibians and Reptiles. Soc. Study Amphib. Reptiles Herpetol. Circ. No. 7. Conant, Roger. 1975. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Houghton Mifflin Co., Boston. Cooper, Byron N. 1966. Geology of the salt and gypsum deposits in the Salt- ville area, Smyth and Washington Counties, Virginia. Pages 11-34 in Second Symposium on Salt, J. L. Rau, editor. North. Ohio Geol. Surv., Cleveland. Vol. 1 . 100 J. Alan Holman and Jerry N. McDonald Dowling, Herndon G., and W. E. Duellman. 1978. Systematic Herpetology: A Synopsis of Families and Higher Categories. Hiss Publications, New York. Holman, J. Alan. 1967. A Pleistocene herpetofauna from Ladds, Georgia. Bull. Ga. Acad. Sci. 25:154-166. , and A. J. Winkler. In press. A mid-Pleistocene (Irvingtonian) herpetofauna from a cave in southcentral Texas. Pearce-Sellards Series, Texas Memorial Museum. Jefferson, Thomas. 1787. Notes on the State of Virginia. (W. Peden, editor, 1972). W. W. Norton and Co., New York. McDonald, Jerry N. 1984. The Saltville, Virginia, Locality: A Summary of Research and Field Trip Guide. Va. Div. Mineral Resour., Charlottesville. . 1985a. Valley-bottom stratigraphy of Saltville Valley, Virginia, and its paleoecological implications. Nat. Geogr. Soc. Res. Rep. 21:291-296. . 1985b. Late Quaternary deposits and paleohydrology of the Salt- ville Valley, southwest Virginia. Current Research, 2: 123-1 24. , and C. S. Bartlett, Jr. 1983. An associated musk ox skeleton from Saltville, Virginia. J. Vertebr. Paleontol. 2:453-470. Ogle, Douglas W. 1981. Long-distance dispersal of vascular halophytes: The marshes of Saltville, Virginia. Castanea 46:8-15. Peterson, O. A. 1917. A fossil-bearing alluvial deposit in Saltville Valley, Vir- ginia. Ann. Carnegie Mus. ll(3/4):469-474. Preston, Robert E. 1979. Late Pleistocene cold-blooded vertebrate faunas from the midcontinental United States: 1. Reptilia: Testudines, Crocodilia. Univ. Mich. Mus. Paleontol. Pap. Paleontol. 19:1-53. Ray, Clayton E., B. N. Cooper, and W. S. Benninghoff. 1967. Fossil mammals and pollen in a late Pleistocene deposit at Saltville, Virginia. J. Paleontol. 41(3):608-622. Stuart, Anthony J. 1979. Pleistocene occurrence of the European pond tortoise (Emys orbicularis L.) in Britain. Boreas 8:359-371. Wilson, Vincent V. 1975. The systematics and paleoecology of two Late Pleistocene herpetofaunas from the southeastern United States. Ph.D. dissert., Michigan State Univ., East Lansing. Accepted 6 May 1985 Discovery of Noturus eleutherus, Noturus stigmosus, and Percina peltata in West Virginia, with Discussions of Other Additions and Records of Fishes Dan A. Cincotta l, Robert L. Miles 2, Michael E. Hoeft 3, and Gerald E. Lewis4 Wildlife Resources Division, West Virginia Department of Natural Resources, Charleston, West Virginia ABSTRACT. — Reports on several West Virginia fishes regarded as part of the state's ichthyofauna or known to inhabit certain drainages are ambiguous. Much of the information is unverifiable, unpublished, or erroneous, and makes preparation of state faunal and endangered species lists problematic. This paper discusses the addition of Alosa sapidissima, Oncorhynchus nerka, Ctenopharyngodon idella, Notropis e. emiliae, Rhinichthys bower si, Noturus eleutherus, N. stigmosus, Lepomis microlophus, Cycleptus elongatus, Percina gymnocephala, P. p. peltata, P. shumardi, Cottus cognatus, and C. girardi to the state checklist. Problem data are also qualified for Ichthyomyzon unicuspis, Lampetra appendix, Hybognathus nuchalis, Notropis dorsalis, Miny- trema melanops, Noturus gyrinus, Etheostoma m. maculatum, and E. tippecanoe. Verifiable or reliable records are documented for all the fishes concerned. West Virginia waters, which include drainages from both sides of the Appalachian divide, contain a fairly unique and diverse ichthy- ofauna (Denoncourt et al. 1975). Although often analyzed as part of several drainages (Denoncourt et al. 1975, Jenkins et al. 1972, Stauffer et al. 1982), the fishes are most easily discussed as constituents of four distinct river systems (Miles 1971, Cincotta and Miles 1982). These are the Potomac and James rivers of the Atlantic slope, and the greater Ohio and New rivers of the Mississippi basin. The New River, techni- cally the upper Kanawha River (Ohio River drainage), is usually regarded as a separate drainage because of its unique faunal assemblage (Addair 1944, Jenkins et al. 1972, Stauffer et al. 1982). Historically, literature pertaining to the fishes of the state was meager and not readily available. The basis for information was dependent on the surveys of Osburn (1901), Goldsborough and Clark Present addresses: ' P.O. Box 67, Elkins, West Virginia 26241; 2 1800 Washington St. East, Charleston, WV 25305; 3 McClintic Wildlife Station, Point Pleasant, WV 25550; 4 P.O: Box 1930, Romney, WV 26757. Brimleyana No. 12:101-121, September 1986 101 102 Dan A. Cincotta, et al. (1908), and Addair (1944). Raney (1947) and Raney and Seaman (1950, cited in Denoncourt et al. 1975) consolidated West Virginia fishery data by discussing the known and expected fauna based on the literature, numerous collections by the West Virginia Conservation Commission, personal sampling, unpublished information, and museum specimens (particularly the re-examination of Goldsborough and Clark's mate- rials). These two checklists, which were designed as the basis for a Con- servation Commission sponsorsed book dedicated to the state ichthy- ofauna (E. A. Seaman, pers. comm.; Anon. 1947), remained internal documents and were not widely disseminated. Unfortunately, the pro- posed publication was not completed. Subsequent to E. C. Raney and E. A. Seaman's efforts and prior to 1970, numerous surveys were conducted in the state. The majority were performed by the Conservation Comission (e.g., W.Va. Wildl. Resour. Div. unpubl. records, Van Meter 1952, Menendez and Robinson 1964, Ross and Lewis 1969) and by F. J. Schwartz (e.g., in Core 1959; Schwartz 1958a, 1959, 1962, 1967). However, most of these data were unverifiable or unpublished. Following this period, several species were added to state faunal and drainage checklists (Miles 1971; Jenkins et al. 1972; Denoncourt et al. 1975; Stauffer et al. 1978, 1982), but were usu- ally reported in an ambiguous manner. Although the works of Ham- brick et al. (1973), Hocutt et al. (1978, 1979; in review), Stauffer et al. (1975, 1980; in press), Hardman et al. (1981), and Cincotta and Hoeft (in press) and certain systematic species reviews (e.g., Denoncourt 1969, Gilbert 1969, Jenkins 1970) clarify much data, distributional informa- tion is lacking for several species and drainages. The purpose of this paper is to add fourteen species to the state faunal list and to clarify several ambiguous fish records. These data were compiled primarily during the preparation of Cincotta and Miles (1982, i.e., revision of Miles 1971), thus reference to this document is omitted. MATERIALS AND METHODS The following species accounts are based on verifiable or reliable data. Confirmation of ambiguous data for discussed species was made via literature review, personal communications with regional investiga- tors, inspection of museum specimens, and examination of unpublished records of the West Virginia Department of Natural Resources, Wildlife Resources Division (WVWR; formerly the Conservation Commission, Fisheries Management Division). Materials from Cornell University (CU), Kentucky Department of Fish and Wildlife Resources (KFW), University of Louisville (UL), University of Michigan Museum of Zool- ogy (UMMZ), Ohio State University (OSU), and National Museum of West Virginia Fishes 103 Natural History (USNM) were used. Data regarding WVWR records and their deposition in the Department of Natural Resources fish museum at Elkins, are summarized in Table 1. Common and scientific names are from Robins et al. (1980). ADDITIONS TO WEST VIRGINIA CHECKLIST The following accounts discuss the addition of fourteen species to the West Virginia ichthyofauna, based on the checklist of Denoncourt et al. (1975). These additions are the result of recent collecting (Noturus eleutherus, Noturus stigmosus, Lepomis microlophus, Percina peltata, Percina schumardi), recent introduction (Ctenopharyngodon idella), data oversights (Alosa sapidissima, Oncorhynchus nerka, Notropis emi- liae, Cycleptus elongatus, Cottus cognatus), description {Percina gym- nocephala), and resurrection (Rhinichthys bowersi, Cottus girardi). Each species discussion is arranged in the order of listing in Robins et al. (1980), with emphasis given to those species collected by WVWR personnel (Table 1). The data presented herein, combined with the addition of Ammo- crypta asprella (Cincotta and Hoeft, in press) and the deletion of Per- cina phoxocephala (Hendricks et al. 1979; Thompson 1980; Stauffer et al., in press) and Notorus gyrinus (discussed in next section), increase the total number of West Virginia species to 164. It should be noted, however, that first West Virginia occurrence records reported by Pear- son and Krumholz (1984) for Lepisosteus platostomus, Notropis boops, N. heterolepis, Erimyzon sucetta, Fundulus notatus, and Etheostoma spectabile were not treated here. These unverified data (W. D. Pearson, pers. comm.) are suspect, based on the information of Trautman (1981), Cooper (1983), and WVWR (unpubl. records). Attempts to verify much of this information by one of the authors (DAC) resulted in either rede- terminations of incorrectly identified fishes or the inability to acquire voucher specimens. Alosa sapidissima (Wilson), American shad This anadromous clupeid is indigenous to Atlantic slope drainages of Canada and the United States (Burgess 1980). It was not reported as part of West Virginia's fauna by Goldsborough and Clark (1908), Raney (1947), Miles (1971), or Denoncourt et al. (1975). Although this shad is native to the lower Potomac River, it was introduced to the upper part (West Virginia and Maryland) of the drainage by the U.S. Fish Com- mission around the turn of the century (Kinney 1963). Omission of this species in past state checklists is attributed to either literature oversight or unsuccessful transplantation. Oncorhynchus nerka (Walbaum), sockeye salmon In North America, this species is native to Pacific slope drainages 104 Dan A. Cincotta, et al. 8°i „ 0> oS S3 *— ' O d U _ c W O ■5 a o „ 'JS c O g o U I* C3 O .55 U OS c « o OS c o c o 60 ."2 « £ H u 3 T3 (L> C - GO c •-> o o Pu O C 3 o u C 6? ON S3 «^ -H -H ON O O 0 r- OO OO 00 ON On ON ON P"M ^H ■*-» ^j > ^j Cu O O O a-> C/3 O 55 O 00 fN 0 ~- tN ^ o < C^ vo 1^- r- r- ON On ^^ <— 1 ^_, i^t a> a, O . J j D* S<3 g3 ^8 2 1 8-3 11 IS la ^ $ 1 S?*s*b ^ §6 t c/3 5 2 42 5 u 2 .2 5 o 2 I" IS S& • If fc^l? 2 2 5 2 5 2§ su aH |e gj 6 3 .;£ 3 U £ U .0 oh ^ ^g § g g, § ". g g 2 g g 2 3 U 2 Q .£5 ^ .5 ^ ^ ^ <^ G . >^ 'S.* "Q "a, *u *o .Si ^c ^ 6 g § S § I * o 9 S ^3 S II S 9 Si S S S.sp a S | d wu WUd ^^ CW "J •-> <. <. Z < OnOn On On NO NO ^O On u ex > O Q. Cu 0 0 < ^ in X/l O O O 0 NO ^ . 3 3 106 Dan A. Cincotta, et al. c cfl o4 1 s X Oj © „ o ii G ed °£ 2 » 2 £ 22 *■ 0-1 • ou > -c « G «-> t o .- M £ 6§ oo . oo H > o w 0 > 09 - at £ oo O Q 0< • O g a X G G S G O OS iu U 3 ° m £ p4 ■g 2 G O U of- * 2 Ih O J3 O on Ou Gfl is o -a o a> eg \o o ^ o 8| 6 Oh ~ G c/i Ad ,« oo J3 2 o ^ -a a 1*1 o °2 o !i o o ^ 2 £ 8 ■^2 «r> <^» S3 O ° «i c £ rt ^ MH o u O cG A< d oj d O 13° __ ♦-, ^ 3 c/3 £ ^ «j x ON G < a oo r-~ o r- r- oo o> On ON ^^ ^^ 00 x> ._ 3 o < G O CM in T* ON VO »n •^ Tt ON »— ' -H CO ro fS a, c/3 "^3 'So West Virginia Fishes 107 and has been stocked in numerous locations within the United States (Lee and Shute 1980). Although Kinney (1963) reported that "California and Pacific salmon" (species unknown) were stocked in the late 1800s in West Virginia waters (along with Alosa sapidissima), no salmon species have ever been included on past state ichthyofaunal checklists. Schwartz (in Jenkins et al. 1972), however, ambiguously indicated O. nerka in the Monongahela River drainage; this information is probably based on his Cheat River, West Virginia, record reported in Core (1959). During the 1950s the landlocked form of this species, the kokanee, was stocked by the WVWR in the Potomac (Stoney River Reservoir, Grant County; Cacapon Lake, Morgan County; Trout Pond, Hardy County), Monon- gahela (Spruce Knob Lake, Tucker County), and New (Watoga Lake, Pocahontas County) river drainages (Van Meter 1953). These records have probably been omitted from the state lists due to literature over- sight or failure of the introductions. Ctenopharyngodon idella Valenciennes, grass carp This species, a native of China, has been introduced throughout the United States for aquatic vegetation control (Guillory 1980). Guillory gave two unconfirmed Kanawha River drainage records. WVWR Di- vision personnel have verified the occurrence of this species in a Nicholas County pond, Gauley River drainage (B. F. Dowler, pers. comm.). Furthermore, some of the specimens from this introduction have sup- posedly been transferred to a pond in Wirt County, Little Kanawha River drainage. To date, there are no records of this species from lotic environments in the state. Notropis emiliae emiliae (Hay), pugnose minnow Gilbert and Bailey (1972) transferred this species from the mono- typic genus Oposopoeodus to Notropis and recognized the subspecies N. e. emiliae and N. e. peninsularis. The latter form is endemic to the Florida peninsula, while the former is found in Lake Erie, Mississippi, and southern Atlantic slope and Gulf coast drainages. Trautman (1981) noted three lower Muskingum River records collected between 1901 and 1938 a few kilometers from the Ohio River, West Virginia (i.e., main channel). He further indicated that, since the species had not been recently collected from this area, it had been extirpated. Apparently, two records for this species have been overlooked in past reviews of the state fauna, as it is not included in previous publications. It was col- lected from Big Run, Wood County, in 1949 (Gilbert and Bailey 1972; CU 21054), and from Oldtown Creek, Mason County, in 1958 (UL 10523, unpubl. data of Krumholz et al. 1962; W. D. Pearson, pers. comm.). These data indicate the presence of this species in the upper Ohio River subsequent to the period discussed by Trautman (1981), and support his contention that it once was more widespread and common. 108 Dan A. Cincotta, et al. Notropis e. emiliae is either extirpated or extremely rare in the upper Ohio River, as there are no recent published records from West Virginia or Ohio. Rhinichthys bowersi Goldsborough and Clark, Cheat minnow This controversial form was originallly described as a species by Goldsborough and Clark (1908), but was subsequently identified as a Nocomis micropogon x Rhinichthys cataractae hybrid by Raney (1940). The distribution of this minnow appears restricted to Lake Erie and Monongahela River drainages (Hendricks et al. 1979; Stauffer et al. 1979). Although Stauffer et al. (1979) indicated that this form qualified morphometrically and meristically as a species, they could not conclu- sively decide its validity. Recent electrophoretic data indicate it is a true species (Goodfellow et al. 1984). In West Virginia, R. bowersi is rare to common in the eastern Monongahela River tributaries (Stauffer et al. 1979; Goodfellow et al. 1984). WVWR personnel recently collected two specimens from Whiteday Creek (Marion/ Monongalia County; WVWR 350), which represents only the second time this minnow has been taken from western tributaries of the Monongahela River. C. H. Hocutt (pers. comm.) indicated that R. bowersi would be petitioned under provisions of the Endangered Species Act of 1973 as a threatened species. Cycleptus elongatus (Lesueur), blue sucker This sucker is usually found in the larger rivers of the Mississippi and Gulf slope drainages (Gilbert 1980a). In West Virginia, Trautman (1981) reported it in the main channel Ohio River. However, probably due to an absence of verifiable historical records (J. R. Stauffer, pers. comm.), Denoncourt et al. (1975) did not include the species on their state checklist. The authors, as did Pearson and Krumholz (1984), accepted the data of Trautman and recognize the species as part of the West Virginia ichthyofauna. Although this sucker has not been taken in numerous surveys in recent years on the West Virginia portion of the Ohio River (Trautman 1981, Preston and White 1978, WVWR unpubl. data), Trautman (1981) reported two records in Ohio near West Virgin- ia. Additionally, a specimen may have been captured (unconfirmed) from the Ohio River adjacent to Hancock County, West Virginia, in 1981 (Pearson and Krumholz 1984). These records are possibly attribu- table to migrating fish from the lower river where the population is improving (W. L. Davis, pers. comm.; Pearson and Krumholz 1984). Noturus eleutherus Jordan, mountain madtom The mountain madtom is found sporadically in southcentral Mis- sissippi River drainages within Oklahoma, Arkansas, and Missouri, and throughout the Ohio River to Pennsylvania (Taylor 1969, Rohde 1980b). In the vicinity of West Virginia, this madtom is known from the Levisa Fork of the Big Sandy River in Kentucky (Jenkins et al. 1972, Rohde West Virginia Fishes 109 1980b, Stauffer et al. 1982, and from tributaries immediately adjacent the main channel Ohio River in Ohio (Trautman 1981). The mountain madtom may have been collected by Krumholz et al. (1962) from the main channel Ohio River of West Virginia, but the specimens assigned UL 11461 and 11617 are missing (W. D. Pearson, pers. comm.). On 20 April 1978 and 16 November 1982, the species was collected from two locations in lower Elk River (Kanawha River drainage) during seining surveys (WVWR 135 verified by Hocutt, 278 by Jenkins). These WVWR records represent the first verifiable evidence of N. eleutherus in the state, and a distributional record for the lower Kanawha River. On each occasion, the mountain madtom was taken in swift riffles (ca. 50 cm depth) containing medium to large rubble. The river was ca. 30 m wide at both sites. Species associates common to both localities were: Etheostoma blennioides, E. camurum, E. tippecanoe, E. variatum, E. zonale, Percina copelandi, and P. macrocephala. Absence of N. eleutherus in past surveys is attributed to a lack of sampling in large rivers and their major tributaries. Noturus stigmosus Taylor, northern madtom Rohde (1980a) gave this madtom's range as tributaries of the Mis- sissippi River from the western margin of Tennessee, northeastward throughout much of the Ohio River basin to the western edge of Penn- sylvania; it also occurs within the western Lake Erie drainages in Ohio, Indiana, and Missouri. Relative to West Virginia, Clay (1975; KFW 1221) and Burr (1980) reported this species from the Levisa Fork of the Big Sandy River in Kentucky, Trautman (1981) reported it from the lower Muskingum River and a minor tributary near the main channel Ohio River in Ohio, and Cooper (1983) reported it from certain tribu- taries of the Allegheny River drainage in Pennsylvania. Denoncourt et al. (1975) expected it to occur within West Virginia waters. The follow- ing data represent the first verfication of the species in West Virginia (C. H. Hocutt, pers. comm.; Stauffer et al. 1982). Paucity of surveys from large rivers probably explains its exclusion from previous collections. In 1976, 1977, and 1981 N. stigmosus was taken from the Kanawha River at London, West Virginia, during lock rotenone surveys (WVWR 27, 48, 352; first two verified by Hocutt). In addition, two specimens were collected from the same area in 1977 by Virginia Polytechnic Insti- tute personnel (C. H. Hocutt, pers. comm.). On 7 October 1980 the fifth collection of this species occurred in Tug Fork River (Big Sandy drain- age) during a rotenone survey near Matewan, Mingo County, West Vir- ginia (WVWR 361). Species common to all WVWR samples were: Notropis volucellus, Moxostoma anisurum, M. macrolepidotum, Ictalu- rus punctatus, Noturus flavus, Pylodictis olivaris, Micropterus punctu- latus, and Percina caprodes. 110 Dan A. Cincotta, et al. Taylor (1969) and Rohde (1980a) reported that in the Ohio River drainage N. stigmosus prefers large creeks and rivers with bottoms of shifting sand and mud, and water varying from clear to turbid with moderate current. The 0.85 hahabitat sampled in the Tug Fork con- sisted primarily of riffles with boulders (30%) and rubble (70%) and a long pool of primarily sand bottom. The water was turbid, and flows in the 30.48-m-wide channel were 4.8 to 5.9 cm/ second. Water quality parameters recorded with a Hach kit at the time of the sampling were: pH (7.6), Fe (.18 mg/ 1), alkalinity (160 mg/ 1 as CaCo^, conductivity (68 micromhos/cm), and water temperature (14.4 °C). This area of the river is known to experience repeated load violations regarding organic sus- pended solids (i.e., domestic sewage) and iron (Steele and McCoy 1980). Lepomis microlophus (Giinther), redear sunfish Lee (1980) considered this species native to the Mississippi, south- ern Atlantic slope, and Gulf slope drainages from Florida to Texas. In the immediate vicinity of West Virginia, the redear sunfish was collected from the main channel Ohio River and the Big Sandy River in Ken- tucky (Clay 1975, Burr 1980, Lee 1980), and the Monongahela River in Pennsylvania (Jenkins et al. 1972, Lee 1980, Stauffer et al. 1982). Denoncourt et al. (1975) listed the redear sunfish as expected, but Miles (1971) regarded it as present in West Virginia based on WVWR records (Anon. 1950, Menendez and Robinson 1964). Other evidence support- ing its existence in the state comes from the Ohio River sampling sum- mary of Preston and White (1978; some L. microlophus specimens veri- fied by M. L. Trautman, pers. comm.) and Trautman (1981). These authors found the species generally infrequent in its introduced range in the upper Ohio River. Percina gymnocephala Beckham, Appalachia darter This endemic upper Kanawha River species was recently described by Beckham (1980). He discussed its relationship to P. maculata and P. peltata. The Appalachia darter appears to be more closely aligned with P. peltata, which is confined to Atlantic slope drainages. Percina gym- nocephala has been recently collected in West Virginia by Hocutt et al. (1978, 1979; in review), Stauffer et al. (1975, 1980), and WVWR (67, 70, 108, 156). These data indicate the species is widely distributed through- out the upper Kanawha River system in West Virginia, but is usually not abundant. Percina peltata peltata (Stauffer), shield darter This darter is known to inhabit streams of the Atlantic slope from New York to North Carolina (Malick 1980). Geographic variation in the species was reported in Raney and Suttkus (1948) as P. p. peltata West Virginia Fishes 1 1 1 from the James River, Virginia, to Hudson River, New York; as P. peltata nevisense from the Neuse and Tar rivers, North Carolina; and as P. p. subspp. from the upper Roanoke River. This percid was expected to occur in the West Virginia part of the Potomac and James rivers by Raney (1947) and Denoncourt et al. (1975). Stauffer et al. (1978) indi- cated that it was not known in the upper Potomac River west of the- Blue Ridge divide. On 15 July 1977 a single specimen of the shield dar- ter was collected from the Shenandoah River, West Virginia, during a boat electrofishing survey (WVWR 398, verified by Jenkins). This cap- ture represents an upstream distribution record, and an addition to the Shenandoah River (R. E. Jenkins, pers. comm.) and West Virginia fauna. Other species taken concurrently were: Anguilla rostrata, Cyprinus carpio, Catostomus commersoni, Hypentelium nigricans, Moxostoma sp., Ictalurus punctatus, Lepomis auritus, L. gibbosus, L. macrochirus, Micropterus dolomieui, and M. salmoides. The inability of past investigators to collect P. p. peltata in the Potomac River, West Virginia, suggests that it is either extremely rare or restricted to large- river habitat. Percina shumardi (Girard), river darter Gilbert (1980b) indicated that the river darter is broadly distributed throughout the Gulf slope, Mississippi basin, Lake Huron, Lake Erie, and Hudson Bay drainages of North America. It is sporadically distrib- uted and rare in the Ohio River basin, especially in the middle and upper reaches of the main channel (Trautman 1981, Clay 1975, Smith 1979, Burr 1980). Trautman (1957) reported it from only a few Ohio localities in the Ohio River drainage. He indicated it was definitely known from the Ohio River proper before 1900, and depicted three records (two in West Virginia) from this period. No new records in West Virginia were noted by Trautman (1981). Although Miles (1971) listed the species as known in the state, Raney (1947) and Denoncourt et al. (1975) reported it as an expected species (probably due to the absence of verifiable specimens). On 14 October 1980, one specimen of the river darter was found in a rotenone sample of an Ohio River backwater area (WVWR 367, verified by R. M. Bailey). This record represents the first report in over 80 years of P. shumardi in the Ohio River, West Virginia. In 1981 another individual was collected from the Ohio River adjacent to Mason County, West Virginia, by personnel of Geo-Marine Inc. (J. A. Pfeiffer, pers. comm.; specimen verified by Pearson). Cottus cognatus Richardson, slimy sculpin This sculpin is broadly distributed in Canada and the northern Uni- ted States. It is found in certain drainages west of the Rocky Moun- tains, the Great Lakes basin, and the north and central Atlantic slope 1 1 2 Dan A. Cincotta, et a 1 . (Wallace et al. 1980). Its southeastern range limit is the Potomac- Shenandoah drainage (R. E. Jenkins, pers. comm.), and the taxonomic status of this Potomac River population is uncertain. Strauss (1980) said that the Potomac River population represents an undescribed endemic species, genetically similar to Cottus girardi but morphometri- cally similar to C. bairdi. However, Jenkins (pers. comm.) indicates it may only be a subspecies of cognatus. For the purpose of this paper, the Potomac River population is recognized as Cottus cognatus. Until 1975, the slimy sculpin was regarded as part of the West Vir- ginia fauna by Raney (1947), Hubbs and Lagler (1958), and Miles (1971). Denoncourt et al. (1975) altered the occurrence status to antici- pated because of the absence of verifiable specimens (J. R. Stauffer, pers. comm.). The only published West Virginia record of this cottid was recently reported ambiguously by Wallace et al. (1980). This infor- mation, which may be in error (R. L. Wallace, pers. comm.), is proba- bly based on a missing UMMZ collection (75426) taken from South Branch Potomac River in 1939. Apparently the first records of this spe- cies in West Virginia were overlooked, as in 1909 E. L. Goldsborough collected it from two locations in the Opequon Creek drainage of the Potomac River, Berkeley County (USNM 64591, 64593; R. E. Strauss, pers. comm.). The only other records of this fish in the state were taken in 1975 and 1981 by WVWR personnel from two streams in Jefferson County, West Virginia (WVWR 256, 257, verified by Jenkins). Species common to both locations were Rhinichthys atratulus, Semotilus mar- garita, and Catostomus commersoni. Absence of C. cognatus from numerous past collections in the West Virginia part of the Potomac River suggests a sparse distribution or confusion with Cottus bairdi or C. girardi. Cottus girardi Robins, Potomac sculpin This species is currently known only from the Potomac, James, and Susquehanna river drainages of the Atlantic slope (Strauss 1977). Although originally described and aligned to the carolinae species group by Robins (1961), Savage (1962) considered it synonymous with Cottus bairdi. Its taxonomic status remained controversial (Jenkins et al. 1972, Mathews et al. 1978, Stauffer et al. 1978) until resurrected by Strauss (1977) and Mathews (1980). It may be fairly common in the upper Potomac River tributaries as suggested by data of Mathews et al. (1978), Jenkins et al. (1980), Goodfellow and Lebo (1981), and Cincotta et al. (ms.). The WVWR has only two verifiable records of this species to date (WVWR 345, 499, former verified by Jenkins). AMBIGUOUS RECORDS The first attempt to document fishes of West Virginia was made by Goldsborough and Clark (1908), but most of their data were collected West Virginia Fishes 1 13 from small waters. It was not until the extensive Kanawha River work of Addair (1944) and the annotated checklist of Raney (1947) that the occurrence and distribution of many species was generally understood. Although the recent drainage surveys by Hocutt et al. (1978, 1979; in review), Stauffer et al. (1975, 1980; in press), and Hardman et al. (1981) resulted in significant contributions in this regard, information relative to several species is lacking. Investigators have encountered difficulty in preparing state nongame or "endangered species" documents because much information relative to West Virginia's ichthyofauna is ambigu- ous, unverifiable, and /or unpublished. This section discusses the status of several species that are uncommon either statewide or in a particular drainage. New information collected by WVWR is noted (Table 1). Ichthyomyzon unicuspis Hubbs and Trautman, silver lamprey This parasitic lamprey is found in the Mississippi basin, primarily from Tennessee northward to the Great Lakes, St. Lawrence and Hud- son Bay drainages (Rohde and Lanteigne-Courchene 1980). It was not reported from West Virginia drainages by Raney (1947), Schwartz (1958b), Jenkins et al. (1972), or Stauffer et al. (1982); but Miles (1971), Denoncourt et al. (1975), and Stauffer (pers. comm.) considered it native on the basis of unpublished WVWR records. The earliest West Virginia record for the silver lamprey was that reported from the main channel Ohio River by Trautman (1957, OSU 11657). This record appears to have been overlooked by past investigators, probably due to the nearness of the site to the boundaries of Kentucky, Ohio, and West Virginia. Verifiable specimens have since been taken from four Ohio River locations (WVWR 113, 132, 153, 228, 390). These data suggest that the silver lamprey population in the upper Ohio River is increasing, rather than decreasing as theorized by Trautman (1981). Lampetra appendix (DeKay), American brook lamprey Lampetra appendix (= lamottei) is a nonparasitic lamprey of the subgenus Lethenteron. It is known from the Great Lakes and Atlantic slope drainages from Minnesota to Virginia, and throughout the middle and upper sections of the Mississippi River basin (Rohde 1980c). Raney (1947) reported this species in the state on the basis of the Monongahela River record of Gribble (1939). Rohde (1980c) did not show the Ameri- can brook lamprey in West Virginia, but indicated occurrence in the Ohio River drainage of Kentucky, Ohio, Pennsylvania, and New York. The species was noted as native to only the Little Kanawha River by Jenkins et al. (1972) and Stauffer et al. (1982). Stauffer (pers. comm.) indicated that there are no confirmable specimens from state waters. WVWR personnel recently collected L. appendix from Middle Island Creek of the Ohio River drainage (WVWR 83, 388). The WVWR vouchers and an uncatalogued Little Kanawha River specimen at the 1 14 Dan A. Cincotta, et al. USNM (F. C. Rohde, pers. comm.) are the only verifiable records of this lamprey from West Virginia. Hybognathus nuchalis Agassiz, Mississippi silvery minnow Pflieger (1980) indicated that H. nuchalis contains three nominal subspecies of uncertain relationships that probably qualify for specific designations due to their morphological distinctiveness and allopatric ranges. The two forms whose ranges encompass West Virginia are H. n. nuchalis, of the Mississippi River and Mobile Bay drainages, and H. n. regius, of the Lake Ontario, St. Lawrence, and Atlantic slope drainages south to Altamaha River, Georgia (Pflieger 1980). Lee et al. (1980) and Robins et al. (1980) recognized the specific distinctiveness of H. regius (see Hubbs and Lagler 1958 for characters). To date, there are no pub- lished records of Hybognathus regius from West Virginia (C. H. Hocutt, pers. comm.; Pflieger 1980). Hybognathus nuchalis was apparently first collected from the state in 1888 from the mouth of the Big Sandy River, Wayne County (Everman 1918). Raney (1947) confirmed the only other silvery minnow record from the Monongahela River drainage, based on a specimen misidentified as Notropis whipplei by Goldsborough and Clark (1908). The exclusion of these records in Jenkins et al. (1972), Pflieger (1980), and Stauffer et al. (1978, 1982) is attributed to either oversight or absence of verifiable materials. Absence of H. nuchalis from recent collections from the upper Ohio River drainages (Preston and White 1978, Trautman 1981) and H. regius from the upper Potomac River drainages (Mathews et al. 1978; Stauffer et al. 1978; Goodfellow and Lebo 1981; Cincotta et al., in ms.) suggests that both are either rare in or extirpated from these waters. Trautman (1957) attributed the silvery minnow's extirpation from Ohio to turbidity and siltation. Notropis dorsalis (Agassiz), bigmouth shiner The bigmouth shiner is found primarily in the upper Mississippi and Great Lakes (excluding Lake Huron) drainages (Gilbert and Bur- gess 1980a). It is discontinuously distributed in the eastern part of its range. Prior to Gilbert and Burgess (1980a), only Schwartz (in Jenkins et al. 1972) and Denoncourt et al. (1975) indicated its presence in West Virginia. Schwartz regarded the species native to the Little Kanawha River, but a lack of verifiable specimens led C. R. Gilbert (pers. comm.) and Stauffer et al. (1982) to doubt this assumption. Gilbert and Burgess (1980a) indicated a single Monongahela River drainage record for West Virginia (UMMZ 198279, Tygart River, collected and identified by C. L. Hubbs and M. B. Trautman). Omission of this shiner from past liter- ature on the Monongahela River is attributed to the obscurity of the record. Notropis dorsalis is probably extirpated from the state, as it has not been collected since 1932. West Virginia Fishes 1 15 Minytrema melanops (Rafinesque), spotted sucker This sucker is known from the lower Great Lakes (Erie, Huron and Michigan), throughout the Mississippi, and from the Gulf slope and southern Atlantic coastal basins (Gilbert and Burgess 1980b). Jenkins et al. (1972), Stauffer et al. (1978), and Hendricks et al. (1979) originally considered this species native to the Monogahela River drainage based on the Youghiogheny River record of Schwartz (1964), but the validity of this record is now questioned since no verifiable specimens exist (Gil- bert and Burgess 1980b, Stauffer et al. 1982). The spotted sucker was confirmed in the West Virginia section of this basin by Raney (1947), based on a specimen misidentified as Moxostoma macrolepidotum by Goldsborough and Clark (1908). Although these data support the record of Schwartz (1964), M. melanops has not been recently collected from the Monongahela River drainage. This sucker is still common in other Ohio River drainages of West Virginia (e.g., WVWR 29, 43, 50, 87). Noturus gyrinus (Mitchell), tadpole madtom The tadpole madtom is found throughout the Mississippi, Gulf coast, and Atlantic slope (including Great Lakes) drainages of North America (Rohde 1980d). Although it is widely distributed in Ohio (Trautman 1981) and is reported in the lower Potomac and James rivers (Stauffer et al. 1982), this species has never been verified from West Virginia waters. Raney (1947) anticipated its occurrence in West Virgin- ia, but Miles (1971) and Denoncourt et al. (1975) listed the species as part of the fauna. This madtom may have been collected from the main channel Ohio River of West Virginia by Krumholz et al. (1962), but no specimens are extant (W. D. Pearson, pers. comm.). The closest records of this species to West Virginia are those of Trautman (1981), only a few kilometers from the state border. Owing to the absence of confirmable records, C. H. Hocutt (pers. comm.) presently regards these species as expected to occur in the state. Etheostoma maculatum maculatum Kirtland, spotted darter Zorach and Raney (1967) reviewed the systematics and distribution of the three recognized subspecies that are restricted to the Ohio River drainage: E. m. maculatum, E. m. sanguifluum, and E. m. vulneratum Etnier (1980) noted that the nominate form exhibited a disjunct distri- bution pattern in the Ohio River basin from New York to Kentucky. Schwartz (in Jenkins et al. 1972) reported E. m. maculatum from lower Kanawha River (below Kanawha Falls), but did not substantiate the record. Based on these unverifiable data the species was listed as part of West Virginia's fauna (Miles 1971, Denoncourt et al. 1975). In 1978, WVWR personnel collected three spotted darters in a rotenone sample 1 16 Dan A. Cincotta, et al. on the Elk River (Kanawha River drainage; WVWR 85). These speci- mens* represent the only verifiable occurrence of this species from West Virginia (J. R. Stauffer, pers. comm.), and this record is depicted in the distributional review of Etnier (1980). Etheostoma tippecanoe Jordan and Everman, Tippecanoe darter This species is restricted to the Ohio River basin, where it is broadly but discontinuously distributed (Zorach 1969). It was first col- lected in West Virginia by WVWR personnel (unpubl. data, verified by Schwartz) in 1966 from Little Kanawha River and later in the same year by Schwartz from Elk River. Although these two unpublished records were overlooked by Zorach (1969), Schwartz ambiguously reported both in Jenkins et al. (1972). Hocutt (1980) depicted records for this percid in the state, but did not include detailed data. WVWR data (WVWR 9, 10, 11, 12, 13, 14, 278) suggest that the species, which is considered generally rare within its range (Kuehne and Barbour 1983), is common in the Little Kanawha and lower Elk rivers. ACKNOWLEDGMENTS.— We particularly wish to express our gratitude to R. M. Bailey, University of Michigan; T. M. Cavender, Ohio State University; R. F. Denoncourt, York College; C. R. Gilbert, Florida State Museum; C. H. Hocutt, University of Maryland; R. E. Jenkins, Roanoke College; W. D. Pearson, University of Louisville; J. Pfeiffer, Geo-Marine, Inc., Piano, Texas; H. R. Preston, U.S. Environ- mental Protection Agency, Wheeling; F. C. Rohde, Chas. T. Main, Inc., Boston; R. Schoknecht, Cornell University; J. R. Stauffer, Jr., Univer- sity of Maryland; R. E. Strauss, University of Michigan; and M. B. Trautman, Ohio State University, for verifying certain species and /or providing data. In addition, Jenkins, Hocutt, and Stauffer reviewed the manuscript and offered critical comments for its improvement. Our appreciation is also extended to present and former West Vir- ginia Wildlife Resources Division (WVWR) personnel who assisted in various aspects of data collection and /or manuscript preparation, par- ticularly K. Watson, T. Oldham, B. Dowler, W. Santonas, J. Rawson, D. Phares, J. Reed, F. Jernejcic, B. Pierce, C. Doerfer, D. Courtney, C. Heartwell, R. Menendez, E. A. Seaman, and S. Muth. Members of the Water Resources Division, West Virginia Department of Natural Resour- ces, Charleston; the U.S. Environmental Protection Agency, Wheeling Field Office; the U.S. Army Corps of Engineers, Huntington and Pitts- burgh Districts; and the Ohio River Valley Water Sanitation Commis- sion, Cincinnati, provided assistance and/ or partial funding for certain collections. Fishery collections were made primarily under Dingell-Johnson Federal Aid Project F-10-R. West Virginia Fishes 1 17 LITERATURE CITED Addair, John. 1944. The fishes of the Kanawha River system in West Virginia and some factors which influence their distribution. Ph.D. dissert., Ohio State Univ., Columbus. Anonymous. 1947. Experts engaged in classifying West Virginia fish. W.Va. Conservation 11(5): 13. . 1950. Commission discovers new West Virginia fish. W.Va. Conservation 14(6):6. Beckham, Eugene C. 1980. Percina gymnocephala, a New Percid Fish of the Subgenus Alvordius, from the New River in North Carolina, Virginia, and West Virginia. Occas. Pap. Mus. Zool. La. State Univ. 57. Burgess, George A. 1980. Alosa sapidissima (Wilson), American shad. Page 67 in D. S. Lee et al., editors. Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. Hist., Raleigh. Burr, Brooks M. 1980. A distributional checklist of the fishes of Kentucky. Brimleyana 3:58-84. Cincotta, Dan A., and M. E. Hoeft. In press. Rediscovery of the crystal darter, Ammocrypta asprella, in the Ohio River basin. Brimleyana 13. , and R. L. Miles. 1982. Checklist of West Virginia Fishes. W.Va. Dep. Nat. Resour. Wildl. Resour. Div., Charleston. Clay, William M. 1975. Fishes of Kentucky. Ky. Dep. Fish Wildl. Resour., Frankfort. Cooper, Edwin L. 1983. Fishes of Pennsylvania and the Northeastern United States. Pa. State Univ. Press, University Park. Core, Earl L. 1959. Biological investigations of Cheat Lake. W.Va. Univ., Morgantown. Denoncourt, Robert F. 1969. Systematic study of the gilt darter Percina evides (Jordan and Copeland) (Pisces, Percidae). Ph.D. dissert., Cornell Univ. , E. C. Raney, C. H. Hocutt, and J. R. Stauffer, Jr. 1975. A checklist of the fishes of West Virginia. Va. J. Sci. 26(3): 1 17-120. Etnier, David A. 1980. Etheostoma maculatum Kirtland, spotted darter. Page 664 in D. S. Lee et al., editors. Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. Hist., Raleigh. Everman, Barton W. 1918. The fishes of Kentucky and Tennessee: a distribu- tional catalogue of the known species. Bull. U.S. Bur. Fish. 35:295-368. Gilbert, Carter R. 1969. Systematics and distribution of the American cyprinid fishes Notropis ariommus and Notropis telescopus. Copeia 1969(3):474-492. . 1980a. Cycleptus elongatus (Lesueur). Page 396 in D. S. Lee et al., editors. Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. 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Etheostoma tippecanoe Jordan and Everman, Tip- pecanoe darter. Page 703 in D. S. Lee et al., editors. Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. Hist., Raleigh. , R. F. Denoncourt, and J. R. Stauffer, Jr. 1978. Fishes of the Greenbrier River, West Virginia, with drainage history of the central Appalachians. J. Biogeogr. 5:59-80. , ,and 1979. Fishes of the Gauley River, West Virginia. Brimleyana 1:47-80. , J. R. Stauffer, Jr., T. Y. Barila, and R. F. Denoncourt. In review. Fishes of the Monongahela National Forest, with a provisional key to the fishes of West Virginia. W.Va. Dep. Nat. Resour. Wildl. Resour. Div., Charleston. Hubbs, Carl L., and K. F. Lagler. 1958. Fishes of the Great Lakes Region. Univ. Mich. Press, Ann Arbor. Jenkins, Robert E. 1970. Systematic studies of the catostomid fish tribe Mox- ostomatini. Ph.D. dissert., Cornell Univ. , E. A. Lachner, and F. J. Schwartz. 1972. Fishes of the central Appalachian drainages: their distribution and dispersal. 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Lampreys of West Virginia. W.Va. Conservation 21(12):8-9. . 1959. Records of the Allegheny brook lamprey Ichthyomyzon greeleyi Hubbs and Trautman from West Virginia, with comments on its occurrence with Lampetra aepyptera (Abbott). Ohio J. Sci. 59:217-220. . 1962. Fishes of Cranesville Swamp. W.Va. Univ. Arboretum Newsletter (winter 1961-62) ll(4):l-5. . 1964. Several Maryland fishes are close to extinction. Md. Con- servationist 41(3):8-12. . 1967. Fishes of the headwaters of Shavers Fork. Pages 28-32 in Cass Scenic Railroad. W.Va. Univ., Morgantown. Smith, Philip W. 1979. The Fishes of Illinois. Univ. Illinois Press, Urbana. Stauffer, Jay R., Jr., R. F. Denoncourt, C. H. Hocutt, and R. L. Miles. In press. Fishes of the Guyandotte River, West Virginia. Nat. Hist. Misc. (Chic). , C. H. Hocutt, M. T. Masnik, and J. E. Reed, Jr. 1975. The longitudinal distribution of the fishes in East River, West Virginia- Virginia. Va. J. Sci. 26:121-125. , and and D. S. Lee. 1978. The zoogeography of the freshwater fishes of the Potomac River Basin. Pages 44-54 in K. C. Flynn West Virginia Fishes 121 and W. T. Mason, editors. The Freshwater Potomac: Aquatic Communities and Environmental Stresses. Proc. Interstate Comm. Potomac River Basin. , and and R. F. Denoncourt. 1979. Status and distribu- tion of the hybrid Nocomis micropogon x Rhinichthys cataractae, with a discussion of hybridization as a viable mode of vertebrate speciation. Am. Midi. Nat. 101(2):355-365. , and and S. L. Markham. 1980. Aquatic biological sur- vey of the New River, Virginia and West Virginia. Final Rep. submitted to U.S. Fish Wildl. Serv., Elkins, West Virginia. , B. M. Burr, C. H. Hocutt, and R. E. Jenkins. 1982. Checklist of the fishes of the central and northern Appalachian mountains. Proc. Biol. Soc. Wash. 95(l):27-47. Steele, B. Douglas, and L. E. McCoy. 1980. Water quality status assessment 1977-1979, W.Va. Dep. Nat. Resour. Wat. Resour. Div., Charleston. Strauss, Richard E. 1977. Morphometric and electrophoretic analysis of the systematic status of Coitus girardi Robins (Pisces:Cottidae). Unpubl. M.S. thesis, Pa. State Univ., University Park. , 1980. Genetic and morphometric variation and the systematic relationship of eastern North American sculpins. Ph.D. dissert., Pa. State Univ., University Park. Taylor, W. Ralph. 1969. A revision of the catfish genus Noturus Rafinesque with an analysis of higher groups of Ictaluridae. U.S. Nat. Mus. Bull. 282. Thompson, Bruce A. 1980. Percina phoxocephala (Nelson), slenderhead dar- ter. Page 737 in D. S. Lee et al., editors. Atlas of North American Fresh- water Fishes. N.C. State Mus. Nat. Hist., Raleigh. Trautman, Milton B. 1957. The Fishes of Ohio. Ohio State Univ. Press, Columbus. . 1981. The Fishes of Ohio. Ohio State Univ. Press, Columbus. Van Meter, Harry. 1952. West Virginia suffers horrid hangover, the bowfin. W.Va. Conservation 16(5):22-23. . 1953. Can we have Kokanee? W.Va. Conservation 17(5):4-5. Wallace, Richard L., D. E. McAllister, and M. Rankin. 1980. Cottus cognatus Richardson, slimy sculpin. Page 809 in D. S. Lee et al., editors. Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. Hist., Raleigh. Zorach, Timothy. 1969. Etheostoma jordani and E. tippecanoe, species of the subgenus Nothonotus (Pisces: Percidae). Am. Midi. Nat. 8 1(2):4 12-434. , and E. C. Raney. 1967. Systematics of the period fish, Etheos- toma maculatum Kirtland, and related species of the subgenus Nothono- tus. Am. Midi. Nat. 77(3):296-322. Accepted 27 December 1984 The Pre-Pliocene Tennessee River and Its Bearing on Crawfish Distribution (Decapoda: Cambaridae) J. F. Fitzpatrick, Jr. Department of Biological Sciences, University of South Alabama, Mobile, Alabama 36688 ABSTRACT.— Recent demand for fossil fuels has provided oppor- tunities for extensive and detailed examination of surface and subsur- face unfossiliferous clastic deposits of the Coastal Plain of the Gulf of Mexico. Among the new discoveries is an ancient outlet directly into the Gulf for the upper Tennessee River more than once during mid- and late Tertiary times. Also discovered is evidence that the intrusion of the Mississippi Embayment apparently occurred much later than implied by surface outcrops in Mississippi and Alabama. Many Cam- baridae distribution patterns show close associations with these Ter- tiary deposits; included are Cambarellus , Fallicambarus, Faxonella, Hobbseus, Procambarus (Acucauda), P. {Girardiella), P. (Leconticam- barus), P. (Pennides), and P. (Scapulicambarus). Some possible inter- pretations relating to these distributions are discussed, as is the pattern of Orconectes and Cambarus invasion. Much detailed study is badly needed, and potentially fruitful areas for investigation are indicated. The earliest attempts to explain the population of North America by cambarid crawfishes were based on the assumption of a Mexican epicenter, from which the major groups radiated to invade the United States and Canada, east of the continental divide. This was probably best articulated by Ortmann (1905). Subsequently, however, Hobbs has presented a cogent and compelling series of arguments in favor of an origin in the southeastern United States (1958, 1962a, 1967, 1969, 1981, 1984; Hobbs and Barr 1972). Probably his best statements appeared in his treatment of the Pictus Group of Procambarus (1958) and his mas- terly analysis of Cambarus (1969). He continued his strong contentions in a monograph of Georgia species (1981) and an analysis of the distri- bution of Procambarus (1984). Although a detailed analysis of phylogenetic relationships is inap- propriate here, it does seem worthwhile to review some of the major trends. Most of these are based on Hobbs. A ProcambarusAike ancestor is generally accepted, and indeed no one has taken issue with Hobbs's contention that the Pictus Group of the subgenus Ortmannicus, of all extant species, is most like the ancestral form (1958). He has, however, recently (1981, 1984) added that certain members of the subgenus Pen- nides are among the most primitive. Although he has somewhat revised his concepts of relationships (1972, 1981, 1984), Hobbs has retained Brimleyana No. 12:123-146, September 1986 123 124 J. F. Fitzpatrick, Jr. much of the phylogeny of Procambarus that he expressed in his review of the Blandingii Section (1962a). Two of his "Groups" in that paper were elevated to subgenera in 1972, with the remaining species of the Section being assigned to the subgenus Ortmannicus. It is important to reemphasize in this paper that certain members of the subgenus Pen- nides (formerly the Spiculifer Group of the Blandingii Section) possess many of the "primitive" characters assigned to the "ancestral procam- barid" (multiple cervical spines; short, broad areola; strongly acuminate rostrum; "striped saddle" pattern of coloration; male first pleopod with full complement of terminal elements, those elements relatively simply constructed; etc.). One must likewise keep in mind that in the Cambari- dae the male and female organs associated with sperm transfer are the most — and sometimes only — reliable taxonomic characters; one can develop good concepts of initial (i.e., early) plesiomorphies in other char- acters/structures, but they are all subject to considerable convergence or modification in response to environmental habits, making determina- tion of synapomorphies nearly impossible. Considerable data are accumulating to indicate that the "upper Tennessee" river had independent access to the Gulf of Mexico at least as recently as the early Pliocene. This new interpretation does not refute the phylogeny of the Cambaridae accepted by the more recent workers, but it does require reexamination of temporal assignments for events. Certain zoogeographic confusions are partially resolved. Alternate explanations to those currently accepted are proposed to (1) account for the distribution of the early-emerging Cambarellinae, (2) elucidate the existence of "primitive forms" of the subgenera Pennides and Ortman- nicus of Procambarus in their present geographic distribution, (3) sug- gest the origin of the subgenus Scapulicambarus as being in lower Georgia in pre-Miocene times, (4) propose that the spread of the genus Fallicambarus east of the Mississippi River is post-Miocene, (5) place the origin of the genus Faxonella in central Louisiana during the Eocene, (6) identify the origin of the genus Hobbseus as eastcentral Mississippi during the Eocene, and (7) suggest pre-Eocene origins for the genera Orconectes and Cambarus, with their spread into the area of the Mississippi Embayment occurring only relatively late in geologic time. PHYLETIC AND ZOOGEOGRAPHIC OVERVIEW The genera Barbie amb arus , Cambarus, Disto cambarus, Fallicam- barus, Faxonella, Hobbseus, Orconectes, and Troglocambarus have been demonstrated to be derivatives of the ancestral procambarid (Hobbs 1967, 1969, 1981). Hobbs, however, did not visualize a more or less lineal descent with a simple cladistic dichotomy. Instead, he postu- lated radiate evolution in which some Procambarus, principally eastern species, arose at one level of the tree and diversified, and a second, somewhat later in time, series of diversifications in one of the stem Tennessee River and Crawfish Distribution 125 stocks organized around an adorconectoid stock. (Mexican diversity, especially interesting to a zoogeographer, is outside the realm of this treatment.) From the former (earlier) populations we see today members of the subgenera Capillicambarus , Hagenides, Lonnbergius, Ortmanni- cus, Pennides, Scapulicambarus , Tenuicambarus, and Villalobosus, plus the genus Troglocambarus . From the adorconectoid line (later temporally), stocks developed that culminate in the procambarid subgenera Acucauda, Austrocambar- us, Girardiella, Leconticambarus , Mexicambarus, Paracambarus, Pro- cambarus, and Remoticambarus , plus the genera Barbicambarus , Cam- barus, Distocambarus, Fallicambarus, Faxonella, Mobbseus, and Orconectes. One of the more striking features of the latter assembly is that, except for Cambarus and Distocambarus, geographically they are more or less western (in relation to the proposed center of origin of the cambarines). Although not a complete "family tree" for the Cambari- dae, Figure 11 of Hobbs's Georgia monograph (1981) is adequate to demonstrate his ideas. He does not visualize polyphyletic origins; instead, he sees the non-procambarid genera as widely divergent stocks that orig- inated from divers stocks of Procambarus. (The groupings as I have made them fundamentally rest on Hobbs, as I have cited him; but if they prove to be non-congruent to his concepts, the fault is entirely misinterpretation on my part.) This latter adorconectoid line seemed to be the less conservative of the two main Procambarus stocks, as evi- denced by the extremes — recognized as genera — of apomorphies devel- oping in it. Another early divergence from the cambarine-procambarid stock resulted in the monogeneric Cambarellinae. Hobbs' last lengthy discus- sion of this phylogeny (1969) was concerned with establishing the rela- tionships between the Cambarinae and the Cambaroidinae, taking for granted an understanding of the close association of the former and the Cambarellinae. More recently, Fitzpatrick (1983) addressed the infrage- neric relationships of the members of Cambarellus and tried to establish their phylogenetic affinities with other Cambaridae. The dwarf craw- fishes are also basically western in distribution. The determination of these lineages did not, however, afford non- moot concepts and explanations of current distributions. Indeed there are many enigmas and paradoxes. Among these are the geographic ranges of those Cambarellus most like the ancestral form, and an expla- nation of why the culminations of an early offshoot of cambarine evolu- tion would be excluded from the proposed ancestral home. Yet they seem to be highly competitive and successful against advanced (and therefore, competitively selected) members of groups that emerged at a later date (Penn and Fitzpatrick 1962, 1963). Members of the subgenus Pennides have many characters attribu- table to the "ancestral procambarid": a full complement of simple ter- minal elements on the male pleopod; multiple carapace spines; a short, 126 J. F. Fitzpatrick, Jr. broad areola; the shape of the rostrum, chela, carapace and pereiopodal coxae; and color pattern. In the subgenus there are two principal assemblages, not formally recognized by Hobbs (1972). In one group, a full complement of terminal elements is present on the gonopod; in the other {gibbus, raneyi, spiculifer) the cephalic process is absent; in P. (Pe.) ouachitae Penn the cephalic process is also sometimes absent. I am not sure that Hobbs would still believe that P. (Pe.) vioscai Penn is the most "primitive" extant species of the subgenus, but there is no doubt that a reduction in terminal elements is apomorphic. Of the three species so disposed, all are found in the most eastern part of the range of the subgenus, while species with a full complement of terminal elements also found in that part of the range have quite specialized pleopods and annuli ventrales (Fig. 1). The more generalized species are found from Mississippi westward. Populations of P. (Pe.) ouachitae (or siblings) occur allopatrically in Arkansas and Mississippi. This species seems to be morphologically intermediate between species with the full-complement of terminal ele- ments and those with a short-complement. Further, the populations of P. (Pe.) vioscai that occur east of the Mississippi River have a much more modified cephalic process than those west of the river. They are sufficiently different that work I have in progress will probably result in my proposing subspecies categories for the two forms. The siblings, P. (Pe.) penni Hobbs and P. (Pe.) clemmeri Hobbs, are so distributed that the more eastern form is also the more remote (from the ancestral type) form (Fitzpatrick 1977a). The entire picture suggests an invasion of the lower Gulf Coastal Plain by an early offshoot of Procambarus stock, and subsequent reinvasion of the southeastern United States along cor- ridors located near the present coastline (Fig. 2). Except for the nearly unique subgenus Lacunicambarus , which Hobbs (1969:163) believed to have been "one of the earliest branching stocks," Cambarus is represented in the central Gulf area only by C. (Depressicambarus) striatus Hay. Bouchard (1978) assigned this species to the superspecific assemblage he considered the more advanced, yet one must remember that Hobbs (1969) believed Depressicambarus to represent a moderately early digression in cambarid evolution. Hobbs (1969:169) conceded that his proposed dispersal corridors to this region, especially for Lacunicambarus, are tenuous. The representatives of Orconectes in the area are all members of specialized and advanced Virilis and Palmeri Groups. Except for Falli- cambarus fodiens (Cottle) and F. uhleri (Faxon), all members of that genus occur on the Gulf Coastal Plain or in reasonable proximity to its central and western parts. Further, the most primitive species lie in southwestern Louisiana and southwestern Arkansas, "probably not far from the ancestral home of the genus" (Hobbs 1969:124). The most "primitive" Faxonella, Fx. creaseri Walls, is found in northcentral Loui- siana, while Hobbseus is confined to the middle and upper Tombigbee Tennessee River and Crawfish Distribution ,88> * Fig. 1. Distribution of Procambarus {Pennides). Arrow designates route of proposed "Miocene Tennessee River." Diagonal rulings = cephalic process pres- ent; horizontal rulings = cephalic process absent; vertical rulings = P. (Pe.) versutus. River drainage (proper) and the upper part of the Pearl River drainage (Fitzpatrick 1977b). Clearly, then, considerable diversity of cambarine crawfishes seems to have originated in a secondary center associated with the lower reaches of the Mississippi River and its environs, markedly distant from the "southeastern" primary center envisioned by Hobbs (loc. cit.). An enigma of how the several populations became established there presents itself. Since this is not a taxonomic paper, it seems improper to continue a discussion of detailed relationships; besides, Hobbs (1967, 1969, 1981, 1984) has explained well our current knowledge of phylogenies. Instead, I propose to examine geographic and geologic information, particularly some recently collected data, which could assist in resolving some of the apparent paradoxes of craw- fish distribution. GEOLOGIC CONSIDERATIONS Classical thinking by crawfish workers (and many others) estab- lishes a thesis that, during some pre-Pleistocene period, the upper and lower portions of the Tennessee River were separate. Faunal compari- sons certainly seem to indicate this. The upper basin is more intimately 128 J. F. Fitzpatrick, Jr. associated with the centers proposed for cambarine, cambaroid, orco- nectoid, graciloid, and mexicanoid stock emergences (Hobbs 1981, 1984). Although the exact routes followed in the past by waters now flow- ing in the Tennessee River do not meet with general agreements among geologists, their paths at specific times are critical to interpretations of crawfish evolution. Hobbs argued (1981:52-53) that the invasion of fresh waters by the cambarine stock occurred in late Cretaceous or early Cenozoic times. He placed them spatially in the tidewater areas of the extreme Southeast. Thus, the route(s) of major watercourses from the southern Appalachians becomes very significant in interpreting the invasion of North America. It is important, too, to recognize that use of the word "river" here designates a basin or drainage source. Rivers themselves have lives measured in thousands of years, not the millions of geologic times. Some geologists (Hayes and Campbell 1894, Hayes 1899) believed that the Appalachian segment of the Tennessee River flowed south- westward through the present Coosa-Alabama basin (or the Black War- rior). They envisioned a capture near Chattanooga at the close of the Tertiary, which led to the present transection of the Walden Ridge. Zoogeographically, this would seem to be supported. A major faunal break seems to be associated with the Walden Gorge. Some geologists (Johnson 1905, Wright 1936) believed otherwise. They insisted that the present route of the Tennessee River has existed at least since the Schooley (dissection of the peneplain ending probably in the Miocene). The geological evidence to support this thesis is of equal strength as that supporting the one of Hayes and some subsequent authors. The Tennessee remains a difficult problem. A good review is in Thornbury (1965:124-126). Sedimentary analysis of Mississippi "Eocene" deposits by Grim (1936), however, provided compelling data to indicate the delta of a sizeable river in eastcentral Mississippi. The Midway alluvial deposits (Paleocene) (Fig. 3) indicate that a significant river had a delta in the vicinity of the Chickasaw-Clay counties area near the juncture of the Porter's Creek and Clayton formations. The succeeding Wilcox deposits (early Eocene) (Fig. 4) demonstrate the continuance of this river into the Choctaw-Montgomery- Webster counties area. Grim (p. 208) attributed both the Midway and Wilcox deposits to a "complex of ancient rocks located in the present Piedmont Plateau." The Claiborne deposits (mid- Eocene), in contrast, suggest that "many streams" (p. 214) rather than one contributed to them. Similarly, the post-Claiborne Jackson Forma- tion (late Eocene) indicates the major "Appalachian [= Tennessee] River" was not a controlling depositional factor in Mississippi. Brown (1967) was concerned over an apparent inconsistency of the major streams of southern Mississippi. Contrary to other Recent drain- age patterns, they flow at a decided angle to the dip and strike of the Tennessee River and Crawfish Distribution ,SsoUr/ Fig. 2. Distribution of western species of Procambarus (Pennides). Arrow as in Figure 1. Solid vertical rulings = P. (Pe.) ablusus; broken vertical rulings = P. (Pe.) lylei; solid horizontal rulings = P. (Pe.) ouachitae; broken horizontal rul- ings = P. (Pe.) clemmeri; cross-hatching = P. (Pe.)penni; stippling = P. (Pe.) la- gniappe; enclosed by open circles (2) = P. (Pe.) elegans. "Miocene" belt. Northeast trending fluvial ridges, which form a drain- age divide, readily explain the disparity (Fig. 5). The underlying depos- its that defend the ridges are mapped as Citronelle Formation (Pliocene- Pleistocene). (It should be noted that many geologists question the accuracy of equating the Mississippi-Alabama Citronelle with the for- mation of the same name farther to the east in Florida and to the west in Louisiana.) Brown's analysis of the gravels led him to postulate the existence of a "very large river flowing southwestward" (p. 82), the gravels forming a part of that river's bed. New studies, using different and more modern techniques, have helped resolve some of these problems. An important aspect of contem- porary geology, especially along the Gulf Coastal Plain, is the greatly expanded search for fossil fuels. Geologists are no longer confined to outcrops as sources of stratigraphic data. Indeed, the economic consid- erations of the petroleum industry have mandated an intensive study of subsurface formations and expanded drilling activities. The masses of new information have transformed the study of the Coastal Plain into a rapidly evolving, incessantly refined activity. Along with this have come many reevaluations of the relationships between stratigraphic series, 130 J. F. Fitzpatrick, Jr. Fig. 3. Midway deposits in Mississippi. (Redrawn from Grim 1936.) Vertical rulings = Porter's Creek Formation; stippling = Clayton Formation. Tennessee River and Crawfish Distribution 131 Fig. 4. Wilcox, loess, and river alluvium in Mississippi. (Redrawn from Grim 1936). Vertical rulings = recent river alluvium; stippling = loess, loam, gravel, etc.; other lines delimit the several formations of the Wilcox deposit. 132 J. F. Fitzpatrick, Jr. interpretations of the implications of the materials that compose them, and clearer understandings of the events and periods of deposition. No longer is the zoogeographer able to rely on a few well-established stud- ies and assume a stability of concept. As the geologic knowledge pro-4 gresses, so the zoogeographic interpretations must follow. And signifi- cant modification of age, stratigraphic relationships, sources of deposits, and biological responses is to be expected as the essentially unfossilifer- ous elastics of the Gulf Coastal alluvia are examined. Isphording (1981), working with drill cores from southwestern Mis- sissippi and especially in the Hattiesburg Formation (Miocene), amassed considerable, nearly irrefutable mineralogical data establishing the existence in Miocene times of a river that entered the Gulf somewhere near Hattiesburg (Fig. 6). Further, these data tie the sediments to the eastern Piedmont and southern Appalachians rather than to the "local" source areas (Isphording 1983). The mineralogical suites encountered are incompatible with weathering from the Mississippi Embayment to the north of the collecting sites or the more remote Rocky Mountains or Central Interior, which had been suggested as sources of the alluvium of the central Embayment by earlier writers (Storm 1945, Murray 1955, MacNeil 1966). Such a river, if not the Tennessee, requires the discovery of yet another river of equal magnitude draining from the same Appa- lachian source area. No geological evidence exists to support such a thesis. Even more data are available to support the contribution of the southern Appalachians to the Embayment. Todd and Folk (1957), working with sediments from Bastrop County, Texas (lower Claiborne), reported that they encountered a kyanite-saurolite suite that they felt could come only from the southern Appalachians, which suite they called "diagnostic" (p. 2560). Isphording (1981) and Brown (1967) implied that the "Eocene" deposits of Grim (1936) were possibly misleading in dating the demise of the last Tennessee outlet directly into the Gulf. Working with geophysi- cal logs and elastics, subsurface and surface, and mapped outcrop pat- terns, May (1981:29) independently reached the same conclusions: "Miocene outcrop patterns should be extended further landward into the Embayment," in Mississippi. Analyses from drillings in northcentral Mississippi led Murphey and Grissinger (1981) to believe that the mate- rials under the Pleistocene loess mantle as far south as Holmes County suggest an erosion surface, frequently out of phase with modern sur- faces. They placed the age, from paleomagnetic data, at earlier than 700,000 B.P. (late Pliocene-early Pleistocene) and postulated a general "Citronelle" age for these deposits. None of these hypotheses seems to be incompatible with Alt's (1974) ideas that modern stream drainage patterns (on the Atlantic coast) began in post-Miocene times. But Mur- phey and Grissinger's (1981) conclusions indicated clearly that modern drainage patterns in the upper Embayment are unreliable indicators of history before the late Pleistocene. Tennessee River and Crawfish Distribution 133 Fig. 5. Proposed Miocene "Tennessee River." (After Brown 1967.) Stippling gravel-defended ridges; broken arrows = proposed route of river. 134 J. F. Fitzpatrick, Jr. It appears, then, that there is considerable evidence to counterindi- cate Smith- Vaniz's (1968:122-124) contention that the present zoogeo- graphic pattern of aquatics (specifically Alabama fishes) must be inter- preted on the basis of the Tennessee occupying its present course at least since Cretaceous times. The only question seems to be when did the connection of the upper Tennessee directly into the Gulf of Mexico become replaced by the indirect Ohio River outlet. Isphording (1981) claimed Miocene or early Pliocene; May (1981) argued Miocene; and Brown (1967) and Murphey and Grissinger (1981) said Pliocene. Grim's Eocene datings (1936) seem possibly compromised, but his stratigraphic relationships remain valid. Equally, one must recognize that nothing in the geologic record requires continuous discharge through a particular basin, and intermit- tent flow remains a viable hypothesis. Indeed, Grim's interpretation of Claiborne sediments seems to indicate this. A river could easily have accounted for Grim's deposits, found another outlet during late Eocene, and reestablished a direct Gulf outlet during Miocene times. It is gener- ally recognized that Miocene is the date of a significant uplift of eastern North America. Even the Citronelle Formation in southern Alabama exhibits a "tilt" to reflect the magnitude of this change (Isphording, pers. comm.). Isphording and Flowers (1980) reexamined the Citronelle in Alabama and Mississippi and suggested that it represents the rework- ing, largely as a result of this uplift, of older deposits. And regardless of precise interpretations, the Miocene uplift surely had profound effects on the directions and flow rates of the then-extant watercourses. Equally, the uplift would have had significant impact on the nature of the gravels and patterns of their deposition. Alt's (1974) opinions on drainages and the Miocene in general were given considerable weight when Hobbs (1981) speculated about phylo- geny. In reviewing the development of the Cambaridae, Hobbs over- looked, possibly deliberately, an important part of Alt's thesis: an arid Miocene. An arid climate would reduce flow of streams and promote emergence of forms adapted to lentic situations. Contrarily, however, the same climate would impede dispersal of crawfishes still adapted to lotic situations. Reduced stream flow would produce a saline intrusion into estuaries. Procambarus (Ortmannicus) acutus acutus (Girard) and P. {Scapulicambarus) clarkii (Girard) are among the very few species with any saline tolerances; thus, the dispersal of cambarines would be effectively blocked in tidewater areas. The overland route would like- wise be impaired, leaving only stream capture as a mechanism for invad- ing new river systems. Fortunately, however, Alt's thesis can be seriously questioned. Isphording (1970) noted that epidote, garnet, and hornblende, although present only a short distance away, are absent from the Kirkwood For- mation and Cohansey Sand of the Middle and Upper Miocene in New Jersey. Otherwise, he found that the remaining heavy mineral species, Tennessee River and Crawfish Distribution 135 MISSOURI 1 / I ilSfHHL Gulf of Mexico Fig. 7. Distribution of non-Mexican Cambarellus. (After Fitzpatrick 1983.) Arrow as in Figure 1. Horizontal ruling = subgenus Dirigicambarus; vertical rulings = subgenus Pandicambarus; crosses indicate small allopatric, probably introduced, populations of Cs. (D.) shufeldtii. entirely different fauna in these habitats, and the latter two faunae are more closely related to each other than either is to the lower Coastal Plain species. Procambarus {Capillicambarus) and most of Fallicambarus are west of the area in question. Procambarus (C) hinei (Ortmann) occurs as far east as the Florida Parishes of Louisiana, but most of the distri- bution of the subgenus is in Louisiana and Texas. The range of the more primitive Fallicambarus suggests origin of the genus west of the Mississippi River with expansion from there. Fallicambarus fodiens is widespread, occurring from lower Ontario to Arkansas and Alabama. Fallicambarus uhleri is a species of the Atlantic Coastal Plain, and F. hortoni Hobbs and Fitzpatrick is apparently of restricted distribution north of the lower Gulf Coastal Plain (Fig. 8). Fallicambarus hedgpethi (Hobbs) scarcely crosses to the east bank of the Mississippi River above the delta region, but it can be found in relatively recent deposits all the way to southwestern Georgia. The latter species and F. fodiens require thorough taxonomic study before firm conclusions about their distribu- tions can be made. 38 J. F. Fitzpatrick, Jr. Mi. ou rL *. ^o^t^ sr Gulf of Mexico Fig. 8. Distribution of Fallicambarus (excluding F. uhleri). Arrow as in Figure 1. Horizontal rulings = F. byersi; vertical rulings = F. oryktes; stippling = F. danielae; enclosed by open circle = F. hortoni. Fallicambarus oryktes (Penn and Marlow) is found in the Florida Parishes of Louisiana and along the Mississippi coast. Its eastern limits abut the western limits of the morphologically and ecologically distinc- tive F. byersi (Hobbs). The latter taxon probably represents more than one species, but this does not interfere with the geographic interpreta- tions; the populations occur as far east as the Yellow River basin in Florida. As does F. oryktes, it (they) occurs in the immediate vicinity of the coast, rarely penetrating more than 100 km inland. Fallicambarus danielae Hobbs is similarly distributed, but apparently it is geographi- cally sympatric with the respective extremes of the two earlier-mentioned species in the central part of the coast. Thus, the spread of these taxa seems to be an event of the late Pliocene or early Pleistocene (Fig. 8). I am not prepared here to discuss the factors that led to establishment of other species of the genus, except to note that the genus and at least some species probably are the result of pre-Pliocene events. Faxonella probably began in the environs of central Louisiana, where one finds the greatest diversity and the apparently most primitive forms. Indeed, only Fx. clypeata (Hay) is widely distributed, and it is found restricted to post-Eocene areas of Alabama and Mississippi in Tennessee River and Crawfish Distribution SsoUr, 139 Fig. 9. Distribution of Faxonella. Arrow as in Figure 1. Horizontal rulings = Fx. clypeata. that part of its range (Fig. 9). Other, apparently later-differentiating species of other taxa, which have similar environmental habits and cohabit successfully with Faxonella elsewhere, are not so widely distrib- uted. Thus, such a distribution as exhibited by Fx. clypeata, a relatively advanced member of the genus, argues for an Eocene origin for the genus. Hobbseus orconectoides Fitzpatrick and Payne, the most primitive member of that genus, occurs in streams associated with Midway (Paleocene) deposits (Fig. 10). The other species occur up and down the Tombigbee drainage, except for one just across the divide in the head- waters of the Pearl drainage. As May (1981) and Murphey and Grissin- ger (1981) suggested that surface materials analyzed by Grim (1936) represent post-Eocene alluvium rather than primary deposits, the above areas could easily be considerably younger than proposed. One cannot escape the close relationship between H. orconectoides habitat and the delta of Grim's (1936) "river of considerable size" or "late Eocene" (probably Miocene). The intimate association of the genus with the Tombigbee drainage makes one suspect that some members of the archiorconectoid stock became isolated in the lower reaches of the river 140 J. F. Fitzpatrick, Jr. during Miocene times and expanded and diversified as the more south- ern lands emerged from the sea and new drainages developed. Procambarus , the largest of the crawfish genera, is expectedly the most complex. And no significiant argument can be made against the supposition that among its members are the species most like the ances- tral Cambarinae. Equally, those species are certain members of P. (Pen- nides) and of the Pictus Group of Ortmannicus. Here an interesting geographic dichotomy occurs. The Pictus Group is unquestionably associated with the Atlantic Coastal Plain, whereas Pennides is found in the Atlantic drainage and the Gulf drainages as far west as Texas (plus an isolate in northern Mexico). The two "groups" within Pennides have been noted, as have been the geographic relationships (Fig. 1, 2). I suggest a very early isolation of the ancestral procambarid stock into eastern and western populations, possibly in the vicinity of northeastern Alabama or northwestern Georgia. Not long afterward, possibly by the large Midway river of Grim, the proto-Pennides were divided. Fitzpatrick and Hobbs III (1968) noted the absence of members of the subgenus from the alluvial plain of the Mississippi River and suggested that such a feature, which denies proper environmental situa- tions, is as effective a barrier as if a dry-land bridge were interposed. Perhaps such a barrier acted to isolate a primitive stock of Pennides. During Miocene times the western stock retained the cephalic process but diversified into a complex of species. Significantly, most widespread members are west of the Mississippi River, but P. (Pe.) vioscai and P. (Pe.) ouachitae have variants on the east side. Recently, Hobbs, Jr., and I have discovered what appears to be a population of P. {Pe.) elegans Hobbs on the east side, but that species seems to be of limited distribu- tion on both sides of the river. Procambarus (Pe. ) ablusus Penn is essen- tially isolated in western Tennessee. The siblings, P. (Pe.) clemmeri and P. (Pe.) penni, are found south of the "river" of Brown (1967), indicat- ing their divergence and spread occurred no earlier than the Pliocene. The other Mississippi species, P. (Pe. ) lagniappe Black and P. (Pe. ) lylei Fitzpatrick and Hobbs, seem to be very restricted, regional isolates. Farther eastward are the species of Pennides that lack a cephalic process. For these, Hobbs's (1981:36-38, 53-54) arguments seem valid. The two enigmas to me are P. (Pe.) petersi Hobbs and P. (Pe.) versutus (Hagen), both of which have a cephalic process. Otherwise, P. (Pe.) petersi is close to P. (Pe.) raneyi Hobbs, morphologically and geo- graphically. Perhaps this is indicative that the eastern proto-Pennides retained for a short while the cephalic process, but most populations lost it early. Surely the most difficult to interpret is P. (Pe.) versutus. Hobbs (1981:38) said, "Considering the Georgia representatives of Pen- nides alone, clearly the most disjunct of the five is Procambarus versu- tus . ..." I concur, but add that it is different from all other Pennides, too. It shares many characteristics with the highly restricted P. (Pe.) lylei. Both have a distinct shoulder on the cephalic surface of the male Tennessee River and Crawfish Distribution 141 M'ssoUr, / Gulf of Mexico Fig. 10. Distribution of Hobbseus. Arrow as in Figure 1. Stippling = H. orconectoides. pleopod; the appendage in each has an attenuated tip; and both have a carinate rostrum. Several western species have caudal projections of the sternite just anterior to the annulus, which partially obscure the recepta- cle, but none is developed in the same way or to the degree as is the case in P. (Pe.) versutus. It is unique in the subgenus in retaining a strong spine on the basis of the cheliped. Despite considerable geographic vari- ation, the species stands alone. It is confined to areas younger than Grim's (1936) "Eocene." Does it represent a third line of proto-Pennides descendants, is it a Miocene phenomenon, or is it both of the preceding? Moving to a second subgenus of Procambarus, Scapulicambarus, another pattern related to post-Miocene development can be seen. Only P. (S.) clarkii (and one other, below) is found significantly outside the southern Atlantic Coastal Plain or the Flint-Chattahoochie basin (Fig. 11). The easternmost limit of this species is in Escambia County, Florida, and where it traverses the coast it is in post-Miocene areas. Again, its dispersal seems to be a post-Miocene event. As its relatives are all in the extreme southeastern United States, an origin in that area is not unreasonable. Equally, a post-Miocene origin is feasible. But since the species has spread as far as Mexico (Hobbs 1962b) in such a 142 J. F. Fitzpatrick, Jr. short time, it becomes a very interesting subject for dispersal and competition studies. The question is complicated by the presence of the relatively primitive P. (S.) strenthi Hobbs (1977) in San Luis Potosl, Mexico. Numerous other problems exist in the undiscussed subgenera of Procambarus. But the purpose of this treatment is not to attempt an exhaustive resolution of zoogeographical situations of North America. Instead, it is to emphasize that more sophisticated knowledge of the geology of an evolutionary critical area can and does require careful reflection on prior conclusions with respect to the phylogeny of the animals, and especially the temporal assignments of events. Thus, the specific answers are best left to other studies. The discussion would not be complete, however, without some mention of the genera Cambarus and Orconectes. As noted above, they both are poorly represented in the area of the old Mississippi Embayment. Until more is known of the precise relationships of the several populations of Cambarus {Lacunicambarus) almost nothing can be said of their history. This was recognized by Hobbs (1969), and the only progress thus far has been the description of two restricted, peripheral species (Fitzpatrick 1978, Hobbs 1981), leaving all the principal questions still unanswered. Otherwise, only C. (Depressicambarus) striatus, an "advanced" member of a "relatively primitive" group, invades to the Mississippi River. Particularly important here are the habits of this species. I have observed individuals moving across open ground when the humidity is only moderately high, and I have found their bur- rows on hillsides somewhat removed from flowing or standing surface water. Surely, this species is not as restricted in its dispersal as are many others. Orconectes is represented by no relatively primitive species. Although the exact relationships of the taxa are presently undetermined, I am sufficiently progressed in a monographic study of the genus to be com- fortable with the concept that the area in question is populated by rela- tively advanced forms. Many are members of the Palmeri Group; they probably represent an invasion from the west. Most of the remainder are Virilis Section species, which probably represent an eastern assem- blage expansion. The striking feature is the absence of simple, less advanced forms. Hobbs's (1967, 1981, 1983) arguments in favor of an early diver- gence of procambarid-like stocks are quite sound. Equally, his ideas of the emergence of proto-Cambarus and proto-Orconectes cannot be faulted. The paradox exists in the geologic data that suggest a large Midway-time river from the southern Appalachians, entering the Mis- sissippi Embayment in the area near the headwaters of the present Pearl River (Grim 1936). Another strong river reworked the "Citronelle" depos- its and emptied just north of Lake Pontchartrain (Brown 1967). Current dating would place these events in late Miocene or early Pliocene. Tennessee River and Crawfish Distribution 143 Fig. 1 1. Distribution of Procambarus {Scapulicambarus). Arrow as in Figure 1 Horizontal rulings = P. (S.) clarkii. Mineralogic data argue strongly that the southern Appalachian high- lands had a significant role in contributing to sediments of the central Gulf Coastal Plain, probably via a major river — the "upper" Tennessee — until late Pliocene times (Isphording 1983). It is difficult to imagine that a vigorous Cambarus and Orconectes stock established in the southern Appalachians or on the Cumberland Plateau would not exploit this route (or routes) for the invasion of the newly emerging habitats. Thus, either the two genera were well estab- lished and diversified by the end of the Miocene or they did not emerge until Pliocene times. Logic favors the former thesis. Otherwise, craw- fishes would be undergoing speciation at a rate not supported by any other evidence. A Miocene intrusion in Mississippi to within 50 km of the Tennes- see boundary (May 1981, Murphey and Grissinger 1981) is a signifi- cantly different situation than previously assumed. As Murphey and Grissinger (1981) indicated, the Eocene (and probably subsequent) drainage patterns have been buried. Surely, the influential Miocene uplift had profound effects on the freshwater drainage. A very fruitful area for study exists in Alabama and Mississippi. Detailed analysis of 144 J. F. Fitzpatrick, Jr. the specifics of microdistribution patterns should reveal much of the geologic, as well as the faunistic, history of the eastern Mississippi Embayment. Correlation of these results with reinterpretation, based on the more recent datings of "Citronelle" deposits, of faunistic patterns to the east or west of the Embayment promises to illuminate the manner in which aquatics populated the southern part of the North American continent. ACKNOWLEDGMENTS.— As always, I benefited greatly from conversations and exchanges of ideas with Horton H. Hobbs, Jr. He and Perry C. Holt read an early draft of this manuscript, and their criticisms contributed much to the development of my thinking. Wayne C. Isphording is thanked for helping me to interpret the complex and often very technical geological literature, and he provided Figure 6 (from Isphording 1983, Fig. 10); George M. Lamb helped with some of the geomorphological concepts. LITERATURE CITED Alt, David. 1974. Arid climate control of Miocene sedimentation and origin of modern drainage, southeastern United States. Pages 21-29 in R. Q. Oakes, Jr., and J. R. DuBar, editors. Post-Miocene Stratigraphy, Central and Southern Atlantic Coastal Plain. Utah State Univ. Press, Logan. Bouchard, Raymond W. 1978. Taxonomy, ecology and phylogeny of the sub- genus Depressicambarus, with the description of a new species from Flor- ida and redescriptions of Cambarus graysoni, Cambarus latimanus, and Cambarus striatus (Decapoda: Cambaridae). Bull. Ala. Mus. Nat. Hist. 3:27-60. Brown, Bahngrell. 1967. A Pliocene Tennessee River hypothesis for Mississippi. Southeast. Geol. 8:81-84. Dorf, Erling. 1960. Climatic changes of the past and present. Am. Sci. 48:341-364. Fitzpatrick, J. F., Jr. 1977a. Distribution of the subgenus Pennides of the crawfish genus Procambarus in Mississippi (Decapoda, Cambaridae). ASB (Assoc. Southeast. Biol.) Bull. 24:51. . 1977b. A new crawfish of the genus Hobbseus from northeast Mississippi, with notes on the origin of the genus. Proc. Biol. Soc. Wash. 90:367-374. . 1978. A new burrowing crawfish of the genus Cambarus from southwest Alabama. Proc. Biol. Soc. Wash. 91:748-755. . 1983. A revision of the dwarf crawfishes (Cambaridae, Cambarel- linae). J. Crustacean Biol. 3(2):266-277. , and H. H. Hobbs III. 1968. The Mississippi River as a barrier to crawfish dispersal. Am. Zool. 8:807. Grim, Ralph E. 1936. The Eocene Sediments of Mississippi. Miss. State Geol. Sur. Bull. 30. Hayes, Charles W. 1899. Physiography of the Chattanooga District. U. S. Geol. Sur. 19th Annu. Rep., Part 2:63-126. Tennessee River and Crawfish Distribution 145 , and M. R. Campbell. 1894. Geomorphology of the southern Appalachians. Natl. Geogr. Mag. 6:63-126. Hobbs, Horton H., Jr. 1958. The evolutionary history of the Pictus Group of the crayfish genus Procambarus (Decapoda, Astacidae). Q. J. Fla. Acad. Sci. 21:71-91. . 1962a. Notes on the affinities of the members of the Blandingii Section of the crayfish genus Procambarus (Decapoda, Astacidae). Tulane Stud. Zool. 9:273-293. . 1962b. La presencia de Procambarus clarkii (Girard) en los estados de Chihuahua y Sonora, Mexico (Decapoda, Astacidae). An. Inst. Biol. Univ. Nac. Auton. Mex. 33:273-276. . 1967. A new crayfish from Alabama caves with notes on the origin of the genera Orconectes and Cambarus (Decapoda: Astacidae). Proc. U. S. Natl. Mus. 123(3621): 1-17. . 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Pages 93-178 in P. C Holt, R. L. Hoffman, and C. W. Hart, Jr., editors. The Distributional History of the Biota of the Southern Appalachians. Part I: Invertebrates. Res. Div. Monogr. 1, Va. Polytech. Inst., Blacksburg. . 1972. The Subgenera of the Crayfish Genus Procambarus (Deca- poda, Astacidae). Smithson. Contrib. Zool. 117. . 1977. A new crayfish (Decapoda: Cambaridae) from San Luis Potosi, Mexico. Proc. Biol. Soc. Wash. 90:412-419. . 1981. The Crayfishes of Georgia. Smithson. Contrib. Zool. 318. . 1984. On the distribution of the crayfish genus Procambarus (Decapoda: Cambaridae). J. Crustacean Biol. 4(1): 12-24. , and T. C. Barr, Jr. 1972. Origins and Affinities of the Troglobitic Crayfishes of North America (Decapoda: Astacidae). II. Genus Orconectes. Smithson. Contrib. Zool. 105. Isphording, Wayne C 1970. Late Tertiary paleoclimate of eastern United States. Am. Assoc. Pet. Geol. Bull. 54:334-343. . 1981. Mineralogical evidence for a Miocene Gulf of Mexico outlet for the ancestral Tennessee River. Abstracts with Program, Southeast. Sect. Geol. Soc. Am. 13(1): 10. . 1983. Interpretive mineralogy: Examples from Miocene Coastal Plain sediments. Trans. Gulf Coast Assoc. Geol. Soc. 33:295-305. , and G. C. Flowers. 1983. Differentiation of unfossiliferous clastic sediments: Solutions from the southern portion of the Alabama-Mississippi Coastal Plain. Tulane Stud. Geol. Paleontol. 17(3):59-83. Johnson, Douglas W. 1905. Tertiary history of the Tennessee River. J. Geol. 13:194-231. MacNeil, F. Stearns. 1966. Middle Tertiary sedimentary regimen of Gulf Coast region. Bull. Am. Assoc. Pet. Geol. 50:2344-2365. May, James E. 1981. The updip limit of Miocene sediments in Mississippi. Abstracts with Program, Southeast. Sec. Geol. Soc. Am. 13(1):29. Murphey, Joseph B., and E. M. Grissinger. 1981. Post-Eocene alluvial materials in north-central Mississippi. Abstracts with Program, Southeast. Sec. Geol. Soc. Am. 13(1):31. 146 J. F. Fitzpatrick, Jr. Murray, Grover E. 1955. Midway Stage, Sabine Stage and Wilcox Group. Am. Assoc. Pet. Geol. Bull. 39:671-696. Ortmann, A. E. 1905. The mutual affinities of the genus Cambarus, and their dispersal over the United States. Proc. Am. Philos. Soc. 44:91-136. Penn, George H., and J. F. Fitzpatrick, Jr. 1962. Interspecific competition between crawfishes Am. Zool. 2:436. , and 1963. Interspecific competition between two sym- patric species of dwarf crawfishes. Ecology 44:793-797. Smith- Vaniz, William F. 1968. Freshwater Fishes of Alabama. Auburn Univ. Agric. Exp. Sta., Auburn. Storm, L. W. 1945. Resume of facts and opinions on sedimentation in the Gulf Coast region of Texas and Louisiana. Am. Assoc. Pet. Geol. Bull. 29:1304-1335. Thornbury, William D. 1965. Regional Geomorphology of the United States. John Wiley and Sons, Inc., New York. Todd, Thomas W., and R. L. Folk. 1957. Basal Claiborne of Texas, record of Appalachian tectonism during Eocene. Am. Assoc. Pet. Geol. Bull. 41:2545-2566. Wright, F. J. 1936. The new Appalachians of the South, Part II. Denison Univ. Sci. Labs 31:93-142. Accepted 13 May 1985 147 To John E. Cooper, with Appreciation In August 1985, John E. Cooper resigned from the staff of the North Carolina State Museum of Natural History and from the editor- ship of Brimleyana. His accomplishments during his 1 1 years here are many. Dr. Cooper nourished Brimleyana from an idea to 1 1 thick issues published between March 1979 and October 1985. Because of his voluminous correspondence with colleagues throughout the country, his broad background as a museum curator and population ecologist, and his special skills as a writer and scientific illustrator, John was unusually well qualified to found and edit an interdisciplinary journal devoted to the zoology and ecology of the Southeast. A native of Maryland, Cooper graduated from Johns Hopkins University and obtained M.S. and Ph.D. degrees from the University of Kentucky at Lexington. Prior to joining the staff of the North Carolina State Museum in September 1974, he lived and taught in Baltimore, where he was a strong, constructive force in the Maryland Natural His- tory Society and the principal editor of Maryland Naturalist. His par- ticular interests are herpetology, crayfish biology, and cave life. At the N.C. State Museum, John organized the Research and Collections Sec- tion and was for a time the assistant director in addition to his service as editor of the journal. Although biologists are supposed to remain detached and analyti- cal in regard to the organisms they study, most of us develop a strong sense of stewardship for them. John is no exception. Well known for his expertise in the biology of cave systems, he deserves credit for the development of the biological and conservation ethics of the National Speleological Society. During his tenure at the museum, he organized the 1975 Symposium on the Endangered and Threatened Plants and Animals of North Carolina, edited the proceedings, and participated in similar symposia in other states. One example is his keynote presenta- tion at the Symposium on Threatened and Endangered Plants and Animals of Maryland. Entitled "Vanishing Species: The Dilemma of Resources Without Price Tags," this is one of the most recent in a long series of scientific contributions dating back to a boyhood interest in biology. John Cooper served this museum well, and when he resigned, he did so in typical Cooper style. He departed just as Brimleyana 1 1 was going to press and after copy for the present issue was ready for typeset- ting. He did everything possible to ensure a smooth transition of 148 responsibility to the managing editor, and now acting editor of the journal, Eloise F. Potter. "Coop," we who worked with you at the museum and the contribu- tors to Brimleyana wish you well in your future endeavors. We will do our best to maintain the high standards you set. JOHN B. FUNDERBURG Director, N.C. State Museum Editor-in-Chief, Brimleyana 149 MANUSCRIPT REVIEWERS The editor and editorial staff are indebted to the following biologists who kindly reviewed manuscripts for Brimleyana Nos. 7 through 10 (1982-1985): Rudolf G. Arndt, Stockton State College Ray E. Ashton, Jr., International Expeditions, Inc. James R. Baker, North Carolina State University Thomas C. Barr, Jr., University of Kentucky Ernest F. Benfield, Virginia Polytechnic Institute and State University William Birkhead, Columbus College Alvin L. Braswell, North Carolina State Museum Richard C. Bruce, Highlands Biological Station Brooks M. Burr, Southern Illinois University at Carbondale Archie Carr, University of Florida Arthur R. Clarke, Ecosearch, Inc. Richard N. Conner, United States Forest Service, USDA E. J. Crossman, Royal Ontario Museum Victor E. Diersing, Museum of Natural History, University of Illinois Philip D. Doerr, North Carolina State University David A. Etnier, University of Tennessee at Knoxville George A. Feldhamer, Appalachian Environmental Laboratory George W. Folkerts, Auburn University Dorothea D. Franzen, Illinois Wesleyan University Thomas W. French, The Nature Conservancy Thomas Fritts, United States Fish and Wildlife Service Samuel L. H. Fuller, Academy of Natural Sciences of Philadelphia John B. Funderburg, North Carolina State Museum J. Edward Gates, Appalachian Environmental Laboratory W. Douglas Harned, Tennessee Valley Authority Julian R. Harrison, III, College of Charleston Richard Hoffman, Radford University Eugene P. Keferl, Brunswick Junior College David S. Lee, North Carolina State Museum Etienne Magnin, Universite'de Montreal William J. Matthews, University of Oklahoma John C. Morse, Clemson University William B. Muchmore, University of Rochester Jerry W. Nagel, East Tennessee State University H. H. Neunzig, North Carolina State University Dan Osterburg, State University of New York at Potsdam William M. Palmer, North Carolina State Museum Peter W. Parmalee, University of Tennessee at Knoxville Peter W. Price, Northern Arizona University Peter C. H. Pritchard, Florida Audubon Society Selwyn S. Roback, Academy of Natural Sciences of Philadelphia 50 Fred C. Rohde, North Carolina Division of Marine Fisheries Robert K. Rose, Old Dominion University C. Robert Shoop, University of Rhode Island Jon D. Standing, University of California at Berkeley William Threlfall, Memorial University of Newfoundland Amy S. VanDevender, Boone, North Carolina J. Reese Voshell, Jr., Virginia Polytechnic Institute and State University S. David Webb, Florida State Museum Paul Yokley, Northern Alabama University DATE OF MAILING Brimleyana No. 1 1 was mailed on 25 November 1985. ACKNOWLEDGMENTS The acting editor is grateful to John E. Cooper, who compiled the list of manuscript reviewers and the index that appear elsewhere in this issue. 151 TABLE OF CONTENTS Number 9 Cicerello, Ronald R. (see Warren, Melvin L., Jr.) 97 Hair, Jay D. (see King, Anne M.) Ill Highton, Richard. A New Species of Woodland Salamander of the Plethodon glutinosus Group from the Southern Appalachian Mountains 1 King, Anne M., Richard A. Lancia, S. Douglas Miller, and Jay D. Hair. Winter Food Habits of Bobcats in North Carolina Ill Lancia, Richard A. (see King, Anne M.) Ill Lenat, David R. Benthic Macroinvertebrates of Cane Creek, North Carolina, and Comparisons with Other Southeastern Streams 53 Manooch, Charles S., Ill, and Diane L. Mason. Comparative Food Studies of Yellowfin Tuna, Thunnus albacares, and Blackfin Tuna, Thunnus atlanticus (Pisces: Scombridae) from the Southeastern and Gulf Coasts of the United States 33 Mason, Diane L. (see Manooch, Charles S., Ill) 33 Mayes, Carol H. (see Shields, Mark A.) . 141 McBride, Steven I., and Donald Tarter. Foods and Feeding Behavior of Sauger, Stizostedion canadense (Smith) (Pisces: Percidae), from Gallipolis Locks and Dam, Ohio River 123 McComb, William C, and Robert L. Rumsey. Bird Density and Habitat Use in Forest Openings Created by Herbicides and Clearcutting in the Central Appalachians 83 Miller, S. Douglas (see King, Anne M.) Ill Nicoletto, Paul F. (see VanDevender, Robert Wayne) 21 Rumsey, Robert L. (see McComb, William C.) 83 Shields, Mark A., and Carol H. Mayes. Occurrence and Habitat Preference of Fundulus luciae (Baird) (Pisces: Cyprinodontidae) on a Southeastern North Carolina Salt Marsh 141 Smith, Charles K. Notes on Breeding Period, Incubation Period, and Egg Masses of Ambystoma jeffersonianum (Green) (Amphibia: Caudata) from the Southern Limits of its Range 135 Tarter, Donald (see McBride, Steven I.) 123 VanDevender, Robert Wayne, and Paul F. Nicoletto. Lower Wilson Creek, Caldwell County, North Carolina: A Thermal Refugium for Reptiles? 21 Warren, Melvin L., Jr., and Ronald R. Cicerello. Drainage Records and Conservation Status Evaluations for Thirteen Kentucky Fishes 97 Wilkins, Kenneth T. Pleistocene Mammals from the Rock Springs Local Fauna, Central Florida 69 Woodward, David K. (see King, Anne M.) Ill 152 Number 10 Ashton, Ray E., Jr. (see Cooper, John E.) 1 Ashton, Ray E., Jr. (see Braswell, Alvin L.) 13 Ashton, Ray E., Jr. Field and Laboratory Observations on Microhabitat Selection, Movements, and Home Range of Necturus lewisi (Brimley) 83 Ashton, Ray E., Jr., (see Hall, Russell J.) 107 Brandon, Ronald A., and James E. Huheey. Salamander Skin Toxins, with Special Reference to Necturus lewisi (Brimley) 75 Braswell, Alvin L., and Ray E Ashton, Jr. Distribution, Ecology, and Feeding Habitats of Necturus lewisi (Brimley) 13 Callard, Gloria V. (see Pudney, Jeffrey) 53 Canick, Jacob A. (see Pudney, Jeffrey) 53 Cooper, John E., and Ray E. Ashton, Jr. The Necturus lewisi Study: Introduction, Selected Literature Review, and Comments on the Hydrologic Units and Their Faunas 1 Hall, Russell J., Ray E. Ashton, Jr., and Richard M. Prouty. Pesticide and PCB Residues in Necturus lewisi (Brimley) 107 Huheey, James E. (see Brandon, Ronald A.) 75 Prouty, Richard M. (see Hall, Russell J.) 107 Pudney, Jeffrey, Jacob A. Canick, and Gloria V. Callard. The Testis and Reproduction in Male Necturus, with Emphasis on N. lewisi (Brimley) 53 Sessions, Stanley K., and John E. Wiley. Chromosome Evolution in the Genus Necturus 37 Wiley, John E. (see Sessions, Stanley K.) 37 153 INDEX TO SCIENTIFIC NAMES (New names in italics) Numbers 9 and 10 New Name Plethodon aureolus 9: 1-20 Ablabesmyia mallochi 9:60 ornata 9:60 parajanta 9:60 Acanthurus 9:38 pomotis 10:92 Acer ssp. 9:22 Acris gryllus 10:92 Acroneuria abnormis 9:56 evoluta 9:56 Agkistrodon contortrix 9:24 Allocapnia spp. 9:56 Aluterus sp. 9:39 Ambystoma 10:79 jeffersonianum 9: 135-140 opacum 10:78 texanum9:134; 10:78 Ameletus lineatus 9:56 Amphinemura sp. 9:56,62 Amphiuma means 10:92 sp. 10:76 Anchytarsus bicolor 9:58 Ancronyx variegata 9:58 Andrias japonicus 10:79 Aneides lugubris 10:76 Anguilla 10:24 rostrata 10:92 Anolis carolinensis 9:21,24,28 Anopheles punctipennis 9:59 Antocha sp. 9:59 Aphredoderus sayanus 10:92 Aplodinotus grunniens 9:127 Argia moesta 9:57 sedula 9:57 spp. 9:57 tibialis 9:57 translata 9:57 Argonauta argo 9:40,43 Aulodrilus pigueti 9:61 pluriseta 9:61 Auxis sp. 9:39 Baesiaeschna janata 9:57 Baetis amplus 9:56 flavistriga 9:56 intercalaris 9:56 pluto 9:56 propinquus 9:56 Baetisca Carolina 9:62 Batrachoseps attenuatus 10:76 Belastoma fluminea 9:57 Bison 9:78 antiquus 9:78 bison 9:78 latifrons 9:78 sp. 9:70,78 Blarina9:71 brevicauda 9:71 carolinensis 9:71 cf. carolinensis 9:70,71 hylophaga 9:71 Bonasa umbellus 9:87,89,1 17 Boyeria vinosa 9:57 Brachycentrus sp. 9:58 Branchiura sowerbyi 9:61 Brillia spp. 9:60 Bufo terrestris 10:92 Buteo jamaicensis 9:87 Caenis cf. diminuta 9:56 Callibaetis sp. 9:56 Calopteryx sp. 9:57 154 Cambarus acuminatus 9:61 (Depressicambarus) latimanus 10:9 reduncus 10:9 (Lacunicambarus) diogenes diogenes 10:9 (Puncticambarus) acuminatus 10:9 Campeloma decisum 9:61 cf. Canis dirus 9:70,75 "Canthyria" 10:8 Caranx crysos 9:38,42 Cardinalis cardinalis 9:87,91 Cardiocladius sp. 9:60 Carphophis amoenus 9:24; 10:26 Carunculina pulla 10:9 Carya spp. 9:84 Castor 10:87 canadensis 9:70,74,1 16 Cathartes aura 9:87 Catostomus commersoni 10:8 Centropristis sp. 9:42 Centroptilum sp. 9:56 Ceraclea ancylus 9:58 tarsipunctata 9:58 Cerataspis monstrosa 9:40,44,48 petila 9:40,44,48 Cernotina sp. 9:62 Certhia familiaris 9:87,89 Chaoborus punctipennis 9:59 Chauliodes pectinicornis 9:57 Cheumatopsyche spp. 9:57 Chilomycterus sp. 9:39 Chimarra cf. aterrima 9:58 Chironomus sp. 9:59 Chrysops sp. 9:59 Cladotanytarsus spp. 9:59 Climacia sp. 9:57 Clinotanypus pinquis 9:60 Cloeon alamance 9:56 Coccyzus americanus 9:89 Colaptes auratus 9:87 Colinus virginianus 9: 1 17 Coluber constrictor 9:24,1 17 Conchapelopia group 9:60 Constempellina sp. 9:59 Contopus virens 9:89 Copelatus glyphicus 9:58 Cordulegaster sayi 9:57 Corydalus cornutus 9:57 Corynoneura spp. 9:60 Cottus carolinae 9:98 Crangonyx spp. 9:61 Cratogeomys 9:80 Cricotopus/Orthocladius gr. 9:60 (C.) bicinctus 9:60 tremulus gr. sp. 1 9:60 sp. 2 9:60 infuscatus 9:60 cf. cylindraceus 9:60 Crotalus giganteus 9:67 horridus 9:23 Cryptobranchus alleganiensis 10:79 Cryptochironomus blarina 9:59 fulvus gr. 9:59 Culex restuans 9:59 Cura foremanii 9:62 Cyanocitta cristata 9:87 Cynops 10:76 Cynoscion sp. 9:42 Dactylopterus volitans 9:39 Dardanus sp. 9:41 Dasypus bellus 9:70,72 Decapterus punctatus 9:38 Dendroica cerulea 9:89 petechia 9:89 virens 9:88,91 Demicryptochironomus sp. 9:59 Desmognathus 10:76 brimleyorum 10:78 fuscus 10:92 monticola 10:92 Dibusa angata 9:55,58 Dicranota sp. 9:59 Dicrotendipes neomodestus 9:59 nervosus 9:59 155 Didelphis virginianus 9:115,116 Didymops transversa 9:57 Dineutes sp. 9:58 Diodon sp. 9:39 Diplectrona modesta 9:57 Diplocladius cultriger 9:60 Dixa sp. 9:59 Dorosoma cepedianum 9:1 17,125,127 Dromogomphus spinosus 9:57 Drymarchon corais 9:69 Dryocopus pileatus 9:87,89 Dubiraphia quadrinotata 9:58 Dugesia tigrina 9:62 Ectopria nervosa 9:58 Elaphe obsoleta 9:23 Elimia sp. 9:61 Elliptio camplanata 9:61 (Canthyria) steinstansana 10:8 icterina 9:61 Empidonax virescens 9:89 Enallagma divergens 9:62 spp. 9:57 Ensatina eschscholtzii 10:76 Ephemerella 9:62 (Antenella) attenuata 9:56 (Danella) simplex 9:56 (E.) catawba 9:56 (Eurylophella) bicolor 9:56 funeralis 9:56 temporalis 9:56 (Seratella) deficiens 9:56 Epitheca cynosura 9:57 Equus 9:79 caballus 9:77 sp. 9:70,77 Erimyzon oblongus 10:92 Esox americanus 10:49,50,92 Etheostoma 10:97,98 camurum 9:103,104 (Catonotus) 9:102 maculatum 9:104 (Nanostoma) 9:102 (Nothonotus) 9:102,104 olmstedi 10:92 tippecanoe 9:104 vitreum 10:92 rufilineatum 9:99 Etrumeus teres 9:42 Eukiefferiella claripenis gr. 9:60 Eumeces fasciatus 9:24 inexpectatus 9:23,24,28 laticeps 9:24 Eunapius sp. 9:62 Eupera cubensis 9:61 Eurycea 10:97 bislineata 10:92 longicauda 10:78 lucifuga 10:78 Fallicambarus (Creaserinus) uhleri 10:9,10 Felis amnicola 9:69,70,75,76 concolor coryi 9:76 rufus 9:1 11-122 sp. 9:76 yagouaroundi 9:76 Ferrissia rivularis 9:61 Fredericella sultana 9:62 Fundulus catenatus 9:102 chrysotus 9:103 luciae 9: 141-144 Geomys 9:73,80,81 bursarius 9:80 pinetis 9:70 Gambusia affinis 10:92 Gerris remigis 9:57 Glaucomys volans 9: 1 16 cf. Glossotherium 9:70,72 Gomphus spp. 9:57 Gyraulus sp. 9:61 Gyrinophilus 10:98 porphyriticus 10:78 Gyrinus sp. 9:58 Hagenius brevistylus 9:57 Hastaperla brevis 9:56 Helichus fastigiatus 9:58 156 Heliosoma anceps 9:61 Helmitheros vermivorus 9:89 Helobdella elongata 9:61 Helocordulia selysii 9:57 Helophorus sp. 9:58 Heptagenia aphrodite 9:56 Heterotrissocladius marcidus 9:60 sp. 9:62 Hexagenia munda 9:56 Hexatoma sp. 9:59 Hippocampus sp. 9:38,42 Holmesina septentrionalis 9:70,72 Hyallela azteca 9:61 Hybognathus hayi 9:100 Hybopsis insignis 9:101 Hydatophylax argus 9:58 Hydrobaenus spp. 9:60 Hydrolimax grisea 9:55,62 Hydroporus sp. 9:58 spp. 9:58 Hydropsyche betteni 9:57 Hylocichla mustelina 9:89 Ictalurus natalis 10:92 Illex 9:46 Ilyodrilus templetoni 9:61 Ischnura spp. 9:57 Isonychia bicolor 9:62 spp. 9:56 Isoperla clio 9:56 namata 9:56 Junco hyemalis 9:87 Juncus roemerianus 9:141,143 Katsuwonus pelamis 9:46 Kiefferulus dux 9:59 Labrundinia neopilosella 9:60 nr. virescens 9:60 Laccophilus sp. 9:58 Lampetra aepyptera 10:15 appendix 9:98,99 lamottei 9:98 Lampropeltis triangulum 9:24 "Lampsilis" ochracea 10:9 Lanthus parvulus 9:57 Larsia sp. 9:60 Lepidostoma sp. 9:58 Lepisoteus oculatus 9:99 Lepomis auritus 10:92 cyanellus 10:92 gulosus 10:92 macrochirus 10:92 marginatus 9:103 Leptophlebia sp. 9:56 Leuctra sp. 9:56 Libellula sp. 9:57 Limnodrilus hoffmeisteri 9:61 Limnogonus sp. 9:57 Limonia sp. 9:59 Liquidambar 9:22 Lirceus sp. 9:61 Liriodendron tulipifera 9:84 Loligo 9:46 Lype diversa 9:58 Macromia allegheniensis 9:57 Macronema Carolina 9:57 Macronychus glabratus 9:58 Magnolia acuminata 9:84 Mammut americanum 9:70,76 Mammuthus sp. 9:70,76 Melanerpes carolinus 9:86,87,89 Mesovelia mulsanti 9:57 Metrobates hesperius 9:57 Micropsectra sp. 9:59 Micropterus salmoides 10:92 Microtendipes pedellus 9:59 nr. rydalensis 9:59 Microtus pennsylvanicus 9: 1 19 pinetorum 9:119 spp. 9:112,116 Microvelia americana 9:57 Mniotilta varia 9:88 Molanna blenda 9:58 157 Molothrus ater 9:117 Monacanthus hispidus 9:39 sp. 9:39,43 Mormoops 9:70,72,79,80 blainvilli 9:79 megalophylla 9:69,70,72,79 Mooreobdella tetragon 9:62 Morone chrysops 9:128 Mugil sp. 9:42 Mus musculus 9:116 Mylohyus 9:79 nasutus 9:70,77 Myotis 9:80 austroriparius 9:70,72,90 grisescens 9:72 Mystacides alafinbriata 9:55,58 Nais bretscheri 9:61 variabilis 9:61 Nanocladius genus nr. 9:60 spp. 9:60 Natarsia sp. 9:60 Nectopsyche sp. 9:57 Necturus 10: 1,3-5, 15,17,18,3 1 ,35,37-74, 79,83,103-105 alabamensis 10:4,5,38-46,48,49 beyeri 10:3-5,29,38,39,41,42,44, 48-50,84,103,104 alabamensis 10:38 lateralis 10:13 lewisi 10:1-35,38-44,46,48,56-59, 66-68,71,72,76,77,79,83-109 maculosus 10:2-6,13-15,38-44,46, 48-50,55-57,59,63,65-67,70,72,76, 83,84,103-105 lewisi 10:3,14,38 louisianensis 10:29 maculosus 10:1,3,38 punctatus 10:4-6,14,15,19,23,26,27,30, 38-46,48,49,92,103,105 alabamensis 10:38 beyeri 10:38 punctatus 10:1,38 Neophylax cf. oligius 9:58 Neotoma floridana 9:116 Nerodia sipedon 9:24; 10:92 Nigronia serricornis 9:57 Nilotanypus sp. 9:60 Nocomis effusus9:102 sp. 10:92 Norocordulia obsoleta 9:57 Notophthalmus 10:76 viridescens 10:76,78 Notropis 10:97,98 altipinnis 10:92 amoenus 10:92 ariommus 9:101,102 atherinoides 9:125,127 chrysocephalus 9:125,127 leuciodus 9:102 procne 10:92 sp. 10:92 telescopus 9:102,103 Noturus furiosus 10:8 insignis 10:92 Odocoileus 9:79 virginianus 9:70,78,1 11,1 16 Oecetis cf. cinerascens 9:62 spp. 9:58 Ondatra 10:87 zibethica9:116 Oporornis formosus 9:88,92 Optioservus ovalis 9:58 Orconectes 10:9 sp. A 10:9 Orthocladius (O.) nr. clarkei 9:60 nr. dorenus 9:60 cf. nigritus 9:60 cf. obumbratus 9:60 roback; 9:60 (Euorthocladius) sp. 1&2 9:60 Oryzomys palustris 9: 1 16 Oulimnius latiusculus 9:58 Oxydendrum arboreum 9:84 158 Palaeomonetes paludosus 9:61; 10:9 Paleolama mirifica 9:70,78 Palpomyla (complex) 9:59 Papogeomys 9:80 Parachaetocladius sp. 9:61 Paracricotopus sp. 9:61 Parakiefferiella sp. 1&3 9:61 nr. triquetra 9:61 Paraleptophlebia sp. 9:56 Paramesotriton 10:76 Parametriocnemus 9:62 Paraphaenocladius sp. 1 9:61 Paratanytarsus sp. 9:60 Paratendipes albimanus 9:59 Parula americana 9:89 Parus bicolor 9:87,88 carolinensis 9:86-88 Passerina cyanea 9:89 Peloscolex variegatus 9:61 Penaeopsis goodei 9:44 Peprilus burti 9:43 triacanthus 9:39,43 Percina macrocephala 9:104,105 peltata 10:92 phoxocephala 9:105 roanoka 10:92 shumardi 9:106 Percopsis omiscomaycus 9:127 Perithemis tenera 9:57 Perlesta placida 9:56 Peromyscus spp. 9: 1 1 6, 1 20 Phaenopsectra flavipes 9:59 sp. 9:59 Philohela minor 9:89 Phtheirichthys lineatus 9:38 Phylocentropus sp, 9:58 Physella sp. 9:61 Picoides pubescens 9:87,89 villosus 9:86,88 Pinus echinata 9:84 rigida 9:84 virginiana 8:22 Pipilo erythrophthalmus 9:87,89,91 Piranga olivacea 9:89 rubra 9:89 Pisidium spp. 9:61 Placobdella multilineata 9:61 papillifera 9:62 Plethodon 10:76 aureolus 9:1-20 caddoensis 9:2 cinereus 10:109 fourchensis 9:2 glutinosus 9:1-4,6-19; 10:78 jordani 9:2,4,6-16,19; 10:76,78 teyahalee 9:7 kentucki 9:2,3,16 ouachitae 9:2 teyahalee 9:4,6-19 websteri 9:2 yonahlossee 9:2 Plumatella repens 9:62 Polioptila caerulea 9:89 Polycentropus spp. 9:58 Polypedilum aviceps 9:59 convictum 9:59 fallax 9:59 illinoense 9:59 scalaenum 9:59 Pomoxis 9:127 Porichthys porossimus 9:42 Portunus sayi 9:41,44,48 sp. 9:41,44 spinicarpus 9:41,48 Prionotus sp. 9:43 Pristigenys alta 9:38 Procambarus 10:29 acutus 9:61 (Ortmannicus) acutus acutus 10:9,10 medialis 10:9,10 plumimanus 10:10 159 Procladius bellus 9:60 sublettei 9:60 Procyon lotor 9:1 15,1 16 Progomphus obscurus 9:62 Prolasmidonta heterodon 10:9 Prosimilium mixtum 9:59 rhizophorum 9:59 Prostoma graecens 9:62,66 Proteus 10:105 anguinus 10:37,84,105 Psectrocladius sp. 9:61 Psectrotanypus dyari 9:60 Psephenus herricki 9:58 Pseudemys concinna 10:92 Pseudocloeon spp. 9:56 Pseudolimnophila sp. 9:59 Pseudorthocladius 9:62 Pseudosmittia sp. 9:61 Pseudotriton 10:79 montanus 10:76,78 ruber 10:76,78 Psilotreta sp. 9:58 Ptilostomis sp. 9:58 Pycnopsyche guttifer 9:58 gentilis 9:58 Quercus alba 9:84 coccinea 9:84 prinus 9:84 rubra 9:84 ssp. 9:22 velutina 9:84 Rana catesbeiana 10:92 clamitans 10:92 palustris 10:92 Regulus calendula 9:87 satrapa 9:87 Reithrodontomys humulis 9:1 16 Remora remora 9:38 Rhagovilia obesa 9:57 Rhantus sp. 9:58 Rheocricotopus cf. robacki 9:60 Rheotanytarsus spp. 9:60 Rheumatobates palosi 9:57 Rhyacophila acutiloba 9:58 Carolina 9:58 ledra 9:58 Salamandra salamandra 10:76 Sargassum 9:36,46 sp. 9:41,44 Sceloporus undulatus 9:24 Scincella laterale 9:23,24,29 Sciurus carolinensis 9: 1 1 2, 1 1 6 Scomber spp. 9:33 Scomberomorus spp. 9:33 Seiurus aurocapillus 9:88,92 Sepioteuthis 9:46 Seriola zonata 9:42 Setophaga ruticilla 9:89 Sialis 9:57 Sicyonia brevirostris 9:40,44,48 sp. 9:44 Sigara spp. 8:57 Sigmodon bakeri 9:74 hispidus 9:74,1 11,1 16 cf. Sigmodon 9:70,74 Simulium vittatum 9:59 Siphloplectron basale 9:56 Siren lacertina 10:76 Sitta carolinensis 9:86,87,89 Slavinia appendiculata 9:61 Somatogyrus sp. 9:61 Spartina 9:46 alterniflora 9:141,143 sp. 9:41 Sphaerium simile 9:61 Sphoeroides sp. 9:39 Sphyrapicus varius 9:87 Squilla empusa 9:40,43 Stactobiella sp. 9:58 Stagnicola sp. 9:61 Stenacron interpunctatum 9:56 pallidum 9:56 Stenelmis spp. 9:58 STATE LIBRARY OF NORTH CAROLINA 3 3091 00748 4884 160 Stenochironomus sp. 9:59 Stenonema (femoratum) 9:56 modestum 9:56 smithae 9:56 vicarium 9:56 Stenotomus caprinus 9:42 Sternotherus odoratus 10:92 Stictochironomus 9:59 Stizostedion canadense 9:123-134 Strix varia 9:80 Strophitus undulatus 9:61 Strophoteryx fasciata 9:56 Stylaria lacustris 9:61 Stylogomphus albistylus 9:57 Sylvilagus aquaticus 9:73 floridanus 9:73 palustris 9:73 sp. 9:70,73 spp. 9:111,116 Sympotthastia sp. 9:60 Synodus sp. 9:42 Tabanus sp. 9:59 Taeniopteryx burksi 9:56 metaqui 9:56 Tamias striatus 9:1 16 Tantilla coronata 9:23,24,29 Tanytarsus glabrescens 9:60 nr. glabrescens 9:60 guerlus gr. 9:60 spp. 9:60 Tapirus copei 9:77 veroensis 9:70,77 Taricha 10:76 granulosa 10:76,78 torosa 10:76,78 Terrapene Carolina 9:24 Thalassia 9:46 testudinum 9:41 Thamnophis sirtalis 9:24 Thienemaniella sp. 9:61 Thomomys 9:70,73,74,79-81 bottae 9:80 cf. orientalis 9:70,73,74 Thunnus 9:33 alalunga 9:47 albacares 9:33 atlanticus 9:33 thunnus 9:47 Tipula abdominalis 9:59 sp. 9:59 Termarctos 9:75 floridanus 9:70,75 Tretenia bavarida gr. 9:60 discoloripes gr. 9:60 Triaenodes injustus 9:58 cf. sp. b 9:58 tardus 9:58 Tribelos jucundus 9:59 Trichechus 9:79 manatus 9:70,76 Trichiurus lepturus 9:39,43 Triturus 10:76 cristatus 10:76 Tropisternus sp. 9:58 cf. Tursiops 9:70,75 Umbra 10:97 limi 9:99,100 pygmaea 10:92 Urocyon cinereoargenteus 9:70,75 Ursus 9:75 americanus 9:70,75,76 Vireo gilvus 9:89 griseus 9:89 olivaceus 9:86,88 Vomer setapinnis 9:42 Wilsonia citrina 9:86,88,91 Wormaldia sp. 9:58 Xenochironomus xenolabis 9:59 Xylotopus par 9:60 Zavrelia sp. 9:60 Zavrelimyia sp. 9:60 Zonotrichia albicollis 9:87 Zostrea marina 9:41,44 sp. 9:41 INFORMATION FOR CONTRIBUTORS Submit original and two copies of manuscripts to Editor, Bnmleyana, North Carolina State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. In the case of multiple authorship, indicate correspondent. Manuscripts submitted for publication in this journal should not also be submitted elsewhere. Preparation of manuscript. Adhere generally to the Council of Biology Editors Style Manual, Fourth Edition. Use medium-weight bond paper, 8V2 X 11", and leave at least an inch margin on all sides. Double space all typewritten material. 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On papers with more than one author, it will be the responsibility of the correspondent to assure that other authors have an opportunity to obtain reprints. Proofs are to be corrected, signed and returned to the Managing Editor within 48 hours. Changes in proofs other than type corrections will be charged to the author. CONTENTS Notes on Turtle Egg Predation by Lampropeltis getulus (Linnaeus) (Reptilia: Colubridae) on the Savannah River Plant, South Carolina. James L. Knight and Raymond K. Loraine 1 Observations on the Social Behavior of the Southern Cricket Frog, Acris gryllus (Anura: Hylidae). Don C. Forester and Richard Daniel 5 Core Temperatures of Non-nesting Western Atlantic Seabirds. Steven P. Platania, Gilbert S. Grant, and David S. Lee 13 Spider Mites and False Spider Mites (Acari: Tetranychidae and Tenuipalpidae) Recorded from or Expected to Occur in North Carolina. Michael K. Hennessey, Daivd L. Stephan, and Maurice H. Farrier 19 Life History of the Wood Frog, Rana sylvatica LeConte (Amphibia: Ranidae), in Alabama. Mark S. Davis and George W. Folkerts 29 Notes on the Eastern Hognose Snake, Heterodon platyrhinos Latreille (Squamata: Colubridae), on a Virginia Barrier Island. David Scott 51 A Study of Variation in Eastern Timber Rattlesnakes, Crotalus horridus Linnae (Serpentes: Viperidae). Christopher W. Brown and Carl H. Ernst 57 Seasonal, Thermal, and Zonal Distribution of Ocean Sunfish, Mola mola (Linnaeus), off the North Carolina Coast. David S. Lee 75 A Late Quaternary Herpetofauna from Saltville, Virginia. J. Alan Holman and Jerry N. McDonald 85 Discovery of Noturus eleutherus, Noturus stigmosus, and Percina peltata in West Virginia, with Discussions of Other Additions and Records of Fishes. Dan A. Cincotta, Robert L. Miles, Michael E. Hoeft, and Gerald E. Lewis .... 101 The Pre-Pliocene Tennessee River and Its Bearing on Crawfish Distribution (Decapoda: Cambaridae). J. F. Fitzpatrick, Jr 123 To John E. Cooper, with Appreciation 147 Manuscript Reviewers 149 Miscellany 1 50 Table of Contents, No. 9 (1983) and No. 10 (1985) 151 Index to Scientific Names, No. 9 (1983) and No. 10 (1985) 153