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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.

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.

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)

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.

(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

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.

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*

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

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

LA4

NC

FL

NY

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

N.C. Spider Mites and False Spider Mites

25

Table 1. Continued.

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

* - 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.

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

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44 Mark S. Davis and George W. Folkerts

stage 21 (cornea transparent), and average 10.7 mm (R = 9.8 to 11.0, SD

= .34, N = 10) in length. Meeks and Nagel (1973) found that hatchling

size in eastern Tennessee averaged 8.0 mm, but did not indicate devel-

opmental stage at hatching. Herreid and Kinney (1967) found that

hatching occurred at stage 20 in Alaskan populations.

An easily observed size difference between hatchling R. sylvatica

and R. sphenocephala allowed us to monitor larval development of

these two species at one pond during 1979. Leopard frog tadpoles hatch

at slightly earlier stages, usually stage 19 (heartbeat) or stage 20, but are

considerably smaller than R. sylvatica tadpoles, averaging 6.6 mm (R =

6.2 to 6.8 mm, SD = .17, N = 10) in length. The results obtained from

the samples are summarized in Figure 3.

Wood frog tadpoles grew more rapidly than leopard frog tadpoles

until about 15 April, at which time the tadpoles of both species were

approximately the same length (R. sylvatica: x- 40.9 mm, N = 10; R.

sphenocephala: x = 40.7 mm, N = 10); however, wood frog tadpoles at

this time were 5 to 1 1 developmental stages beyond leopard frog tad-

poles. Leopard frog tadpoles then continued to increase in length, where-

as wood frog tadpoles began to decrease as a result of initial tail resorp-

tion with the onset of metamorphic climax. Wood frog tadpoles were

last collected in the pond on 29 April, at which time most were in stage

42 (both forelimbs erupted) and averaged 40.5 mm in length. Leopard

frog tadpoles averaged 53.6 mm in length at this time, with most indi-

viduals in stage 35 (toes 1 and 2 joined, others separate). Leopard frog

tadpoles collected between 29 April and 5 May revealed that tadpoles of

this species continue to grow. Mean length of R. sphenocephala tad-

poles collected on 5 May (not shown on graph) was 56.0 mm, at which

time larvae were in stage 38 (metatarsal tubercle formation).

It has been shown that wood frog and leopard frog tadpoles may

behave as ecological equals (DeBenedictus 1974). The larger size and

more advanced stage of development at hatching may give R. sylvatica

tadpoles some initial competitive advantage over those of R. spheno-

cephala. The more rapid development of R. sylvatica probably repre-

sents an adaptation to breeding in temporary pools and might result in

some resource partitioning on those infrequent occasions when these

two species use the same breeding ponds. Alford and Crump (1982)

found size class segregations in R. sphenocephala tadpoles, both in

laboratory experiments and field situations, and felt that the negative

correlation between large (and/ or older) and small (and/ or younger)

tadpoles indicated habitat partitioning.

Since newly transformed R. sylvatica froglets were not collected in

the field, the exact amount of time from oviposition until transforma-

tion is not known. No wood frog tadpoles were collected on 5 May,

Wood Frog Life History

45

60

50

40

30

- 20

10

S36

□ R.sylvatica

• R.sphenocephala

date

Fig. 3. Larval development of Rana sylvatica and Rana sphenocephala in Pas-

ture Pond, Tallapoosa County, Alabama. Numerals preceded by the letter S

indicate the developmental stage in tadpoles at the time of collection.

however, indicating that all larvae were transformed at this time. This

would give a maximum transformation period of 73 days in the field.

Most individuals had probably transformed by 29 April (or somewhat

earlier); only four tadpoles were collected in the pond at that time. This

would indicate a transformation time of about 66 days.

Various vertebrates and invertebrates preyed on wood frog larvae,

though not all predators were present at each breeding pond. All inver-

tebrate predators were insects. Adult back swimmers (Notonectidae)

and predaceous diving beetles (Dytiscidae) were often seen preying on

small tadpoles. Nymphal notonectids and larval dytiscids also probably

preyed on wood frog larvae, as observed by Dickerson (1906). Herreid

and Kinney (1966) noted extensive predation on wood frog larvae by

Dytiscus spp. in Alaska. Formanowicz and Brodie (1982) found no sur-

vival of stage 42 and younger wood frog tadpoles when subjected to

predation by larval Dytiscus verticalis in the laboratory. Increased sur-

vivorship in older tadpoles and froglets (stage 42 to 46) was attributed

to unpalatability, a result of the development of active granular glands

during later stages of metamorphosis. Other potential insect predators

46 Mark S. Davis and George W. Folkerts

present were odonate naiads (Libellulidae, Lestidae, Coenagrionidae)

and nymphal and adult giant water bugs (Belostomatidae).

The most significant vertebrate predator on R. sylvatica tadpoles

appeared to be the larvae of Ambystoma opacum. These salamanders

were always present in the breeding ponds and were usually seen in

close proximity to wood frog egg masses prior to hatching. Salamander

larvae probably were attracted to unhatched eggs by movements of

developing embryos. Walters (1975) stated that eggs and larvae of R.

sylvatica were readily eaten by adult newts and marbled salamanders,

but neither of these was noted as a predator during our study. Fish were

usually absent from the breeding ponds, but one potential tadpole pred-

ator, Lepomis cyanellus, was encountered in two ponds. No predation

by this species was observed during our study.

Adult Food Habits

We examined stomach and intestinal contents of 42 adult R.

sylvatica from Alabama. Because only 14 of these specimens contained

identifiable food items (Table 3), this analysis is useful only for generali-

zation. Alabama wood frogs appear to be opportunistic terrestrial feed-

ers. Insects, spiders, earthworms, and snails were the major food items

present. A scarab beetle, Eutheola rugiceps, was discovered in the coe-

lom of one specimen. A large hole in the stomach indicated that the

beetle had torn its way through the stomach wall after being ingested

(Davis and Folkerts 1980). Neither males nor females collected in breed-

ing ponds contained food. Since all were immediately preserved, we

assume that neither sex feeds at this time. Adult wood frogs may not

require food during such a short, explosive breeding period.

ACKNOWLEDGMENTS. — For companionship during many late-

night collecting trips, and a multitude of favors, we thank Tom Jones.

Dan Combs, Lee Elliot, Win Seyle, and Dan Warren provided assist-

ance in the field. Sonny Eiland and Pete Lahanas helped identify food

items and larval insects. Winston Baker provided the locality of a new

breeding population. Tom Yarbrough gave advice on making trips to

Mt. Cheaha, and Tom Johnson provided unpublished information on

Missouri wood frogs. Robert Mount, Wayne Clark, and John Cooper

reviewed early drafts of the manuscript and made numerous helpful

suggestions. Sue Kahre drew the range maps; Sharna King and Cheryl

James kindly typed the manuscript on short notice. Several people at

the University of Missouri provided assistance in some form and to

them we extend our heartfelt thanks: Richard Daniel, Carl Gerhardt,

Brian Miller, and Robert Witcher. This paper represents part of a thesis

Wood Frog.Life History 47

Table 3. Composition of stomach contents of 14 adult Rana sylvatica collected

in Alabama.

Food items Percent of stomachs

containing item

Arachnida

Tetragnathidae 43

Insecta

Homoptera

Membracidae 7

Orthoptera

Blattellidae 7

Gryllacrididae 29

Plecoptera 7

Coleoptera*

Elateridae 14

Carabidae 7

Staphylinidae 14

Scarabaeidae 7

Annelida

Lumbridae 21

Mollusca

Gastropoda 21

* Several beetles were not identifiable.

submitted by Davis to the Department of Zoology-Entomology, Auburn

University, in partial fulfillment of the requirements for the M.S.

degree.

LITERATURE CITED

Alford, Ross A., and M. L. Crump. 1982. Habitat partitioning among size

classes of larval southern leopard frogs, Rana utricularia. Copeia

1982(2):367-373.

Anderson, James D., D. D. Hassinger, and G. H. Dalrymple. 1971. Natural

mortality of eggs and larvae of Ambystoma t. tigrinum. Ecology

52:1107-1112.

Banta, Arthur M. 1914. Sex recognition and mating behavior of the wood

frog, Rana sylvatica. Bio. Bull. 26:171-183.

Behler, John L., and F. W. King. 1979. The Audubon Society Field Guide to

North American Reptiles and Amphibians. Alfred A. Knopf, Inc., New

York.

Bellis, Edward D. 1957. An ecological study of the wood frog, Rana sylvatica

48 Mark S. Davis and George W. Folkerts

LeConte. Unpubl. Ph.D. dissert., Univ. Minnesota, Minneapolis.

Berven, Keith A. 1981. Mate choice in the wood frog, Rana sylvatica. Evolu-

tion 35(4):707-722.

. 1982a. The genetic basis of altitudinal variation in the wood frog,

Rana sylvatica. I. An experimental analysis of life history traits. Evolution

36(5):962-983.

. 1982b. The genetic basis of altitudinal variation in the wood frog,

Rana sylvatica. II. An experimental analysis of larval development. Oeco-

logia 52:360-369.

, and D. E. Gill. 1983. Interpreting geographic variation in life

history traits. Am. Zool. 23(l):85-97.

Collins, James P. 1975. A comparative study of the life history strategies in a

community of frogs. Unpubl. Ph.D. dissert., Univ. Michigan, Ann Arbor.

, and H. M. Wilbur. 1979. Breeding Habits and Habitats of the

Amphibians of the Edwin S. George Reserve, Michigan, with Notes on the

Local Distribution of Fishes. Occas. Pap. Mus. Zool. Univ. Mich. No. 686.

Cory, Lawrence, and J. J. Manion. 1953. Predation on eggs of the wood frog,

Rana sylvatica, by leeches. Copeia 1953(1):66.

Davis, Mark S., and G. W. Folkerts. 1980. A beetle (Eutheola rugiceps LeC:

Scarabaeidae) penetrates the stomach wall of its predator {Rana sylvatica

LeC: Amphibia). Coleopt. Bull. 34(4):396.

DeBenedictus, Paul A. 1974. Interspecific competition between tadpoles of

Rana pipiens and Rana sylvatica: An experimental field study. Ecol.

Monogr. 44:129-151.

DeGraaf, Richard M., and D. D. Rudis. 1983. Amphibians and Reptiles of

New England. Univ. Mass. Press, Amherst.

Dickerson, Mary C. 1906. The Frog Book. Doubleday, Page and Co., Garden

City.

Formanowicz, Daniel R., and E. D. Brodie. 1982. Relative payabilities of

members of a larval amphibian community. Copeia 1982(l):91-97.

Gatz, A. John. 1973. Algal entry into the eggs of Amby stoma maculatum. J.

Herpetol. 7:137-138.

Gilbert, Perry W. 1942. Observations on the eggs of Amby stoma maculatum

with especial reference to the green algae found within the egg envelopes.

Ecology 23(2):215-227.

. 1944. The alga-egg relationship in A mbystoma maculatum, a case

of symbiosis. Ecology 25(3):366-369.

Golden, M. S. 1968. Oak-hickory components of the forests of the Alabama

Piedmont. Unpubl. M.S. thesis, Auburn Univ., Auburn.

Gosner, Kenneth L. 1960. A simplified table for staging anuran embryos and

larvae with notes on identification. Herpetologica 15(1 4): 203-2 10.

Green, N. B. 1952. A key to the eggs of West Virginia Salientia. Proc. W. Va.

Acad. Sci. 24:36-38.

Hammen, Carl S., and V. H. Hutchinson. 1962. Carbon dioxide assimilation

in the symbiosis of the salamander, Ambystoma maculatum, and the alga,

Oophila amblystomatis. Life Sci. 10:527-532.

Harper, Roland M. 1943. Forests of Alabama. Geol. Surv. Ala. Monogr. 10.

Wood Frog Life History 49

Herreid, Clyde F., and S. Kinney. 1966. Survival of Alaskan wood frog (Rana

sylvatica) larvae. Ecology 47(6): 1039-1041.

, and 1967. Temperature and development of the wood

frog, Rana sylvatica, in Alaska. Ecology 48(4):579-590.

Hinckley, Mary H. 1882. Notes on the development of Rana sylvatica

LeConte. Proc. Bost. Nat. Hist. Soc. 22:85-95.

Hodgkins, E. J. (editor). 1965. Southeastern Forest Habitat Regions Based on

Physiography. Auburn Univ. Agric. Exp. Stn. For. Dep. Series No. 2.

Howard, Richard D. 1980. Mating behavior and mating success in wood frogs,

Rana sylvatica. Anim. Behav. 28:705-716.

Hudson, Robert G. 1954. An annotated checklist of the reptiles and amphibi-

ans of the Unami Valley, Pennsylvania. Herpetologica 10:67-72.

Hutchinson, Victor H., and C. S. Hammen. 1958. Oxygen utilization in the

symbiosis of the embryos of the salamander, Amby stoma maculatum, and

the alga, Oophila amblystomatis. Biol. Bull. 115:483-491.

Johnson, Evert W., and L. R. Sellman. 1975. Forest Cover Photointerpreta-

tions Key for the Mountain Forest Habitat Region in Alabama. Auburn

Univ. Agric. Exp. Stn. For. Dep. Series No. 7.

Martof, Bernard S. 1970. Rana sylvatica. Cat. Amer. Amphib. Rept.: 86.1-86.4.

, and R. L. Humphries. 1959. Geographic variation in the wood

frog, Rana sylvatica. Am. Midi. Nat. 61(2):350-389.

Meeks, David E., and J. W. Nagel. 1973. Reproduction and development of

the wood frog, Rana sylvatica, in eastern Tennessee. Herpetologica

29:188-191.

Minton, Sherman A. 1972. Amphibians and Reptiles of Indiana. Indiana

Acad. Sci. Monogr. No. 3.

Moore, John A. 1949. Patterns of evolution in the genus Rana. Pages 315-338

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in frogs and toads. Adv. Genet. 7:139-182.

Mount, Robert H. 1975. The Reptiles and Amphibians of Alabama. Auburn

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Nelson, Craig E. 1971. Breakdown of ethological isolation between Rana

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Noble, G. Kingsley, and E. J. Farris. 1929. The method of sex recognition in

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Savage, R. Maxwell. 1961. The Ecology and Life History of the Common

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50 Mark S. Davis and George W. Folkerts

Smith, Philip W. 1961. The Amphibians and Reptiles of Illinois. Bull. 111. Nat.

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Wright, Albert H. 1914. North America Anura. Life Histories of the Anura of

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Accepted 18 June 1985

Notes on the Eastern Hognose Snake,

Heterodon platyrhinos Latreille (Squamata:

Colubridae), on a Virginia Barrier Island

David Scott

Savannah River Ecology Laboratory,

P.O. Drawer E, Aiken, South Carolina 29801

ABSTRACT. — An unusually high population density of the eastern

hognose snake, Heterodon platyrhinos Latreille, is reported from a

Virginia barrier island. The average snout-vent length of females in

this population is significantly greater than the average SVL of males,

but individuals of equal SVL do not differ in body mass. Females also

differ from males in seasonal activity patterns, with males most active

in early summer and females in late summer. The number of dorsal

blotches of the island population differs significantly from that reported

for a mainland population on the Delmarva Peninsula.

The eastern hognose snake, Heterodon platyrhinos Latreille, is

found over much of the eastern United States (Conant 1975). Within

this range it uses a great diversity of hatitats, but like other members of

the genus Heterodon it prefers dry, relatively open, sandy areas in which

it can burrow easily (Corrington 1929, Lynn 1936, Duellman and

Schwartz 1958, Piatt 1969). This type of habitat is abundant on Tom's

Cove Hook, a fast-growing recurved spit on the southern tip of Assa-

teague Island, Virginia, and H. platyrhinos is especially conspicuous at

this site.

Three factors may account for the apparently high density of hog-

nose snakes. First, Assateague hosts a depauperate snake fauna, as do

many Atlantic Coast barrier islands (Gibbons and Coker 1976). Only 6

species occur on the island (Lee 1972), compared to 17 on the adjacent

Delmarva Peninsula (Martof 1980). The absence of other species may

promote increased numbers of H. platyrhinos. Second, the sparsely

vegetated dunes that form the interior of the spit provide ideal foraging

habitat. Fowler's toads, Bufo woodhousei fowleri, breed in freshwater

ponds between the dune lines and are abundant on the dunes. Hognose

snakes were observed several times in the act of hunting and capturing

toads buried in the sand. Third, it is possible that hognose snakes on

Assateague are no more abundant than on the mainland, but are simply

easier to census because of the open habitat. Given the apparent abun-

dance of H. platyrhinos, the objective of this paper is to present infor-

mation that is ordinarily difficult to obtain for a single population.

Brimleyana No. 12:51-55, September 1986 5 1

52 David Scott

STUDY AREA AND METHODS

I observed active hognose snakes while on a preliminary visit to

Assateague Island on 18 April 1981. The vegetation of Tom's Cove

Hook consists of discrete zones of sand-dune, shrub, grassland, and

salt-marsh habitats. Few snakes were observed in shrub and grassland

areas during preliminary sampling, and none were seen in the salt

marsh. Therefore, systematic sampling was confined to the sand-dune

habitat on the inland dune ridges that form a nexus on the spit. These

ridges were searched two or three times per week (22 days) from 1 1 June

to 12 September 1981. The sequence in which the dunes were searched

was varied. The first five searches lasted from 0800 to 1930. No snakes

were observed during midafternoon hours, and subsequent sampling

was confined to morning and late afternoon. On five occasions searches

were conducted for a 3-hour period after sunset.

The following data were recorded for each capture: date, time of

capture, exact location on the dune lines, temperature of the substrate

(52 captures only), color of snake, snout-vent length (SVL) to the near-

est 0.5 cm, body mass (g), number of dorsal blotches, and number of

ventral and subcaudal scales. Each individual was marked with a unique

identification code (ID) by clipping two subcaudal and three ventral

scales. For recaptured individuals, the linear distance traveled between

captures was estimated from an aerial photograph of the site. Sex (Fig. 1)

was determined by an analysis of ventral and subcaudal scale counts

(Edgren 1961). Five of the snakes that were classified as males according

to scale count were noted to have everted hemipenes during handling,

supporting the assumption that sex can be determined by scale count.

RESULTS AND DISCUSSION

A total of 66 individuals were captured, with 6 individuals (9.1%)

recaptured an average of 17.5 days after initial capture. More males (N

= 38) were captured than females (N = 28), but the ratio did not differ

from 1:1 (x2= 1.52, df = 1, p >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

TCB

SVL

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

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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.

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.

3.00;

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FACTOR 2

3.75

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FACTOR 2

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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

-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

-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

1.5

1.0

F .50

A

C 0.0

T

O-.50

R

4-1.0

-1.5

-2.0

■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.

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

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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

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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

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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

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

Fig. 1. The Saltville, Virginia, locality, showing location of sites within Saltville

Valley that have produced herptile specimens mentioned in text.

1966). This bedrock has been scoured and incised in places by Quater-

nary stream action, and is now overlain by up to 3 m of late Quaternary

sediments.

The Quaternary sediments result from multiple depositional epi-

sodes dating from the ?Sangamonian interglaciation through the Holo-

cene (Fig. 2, Table 1). Terrace-like deposits above 530 m in elevation

occur at several places around the edge of the valley; these are consid-

ered tentatively to date from Sangamon time. Most of the sediments

88 J. Alan Holman and Jerry N. McDonald

lying below 530 m elevation are late Wisconsinan or Holocene in age

(McDonald 1984, 1985a). The older of these late Wisconsinan sedi-

ments are fluvial deposits, laid down alongside or in the channel of the

extinct Saltville River before its capture by upstream piracy around

14,000 B.P. (McDonald and Bartlett 1983). The oldest and most exten-

sive of these fluvial deposits is a sheet of rounded gravel (Unit W4: peb-

bles to cobbles), containing numerous bones and teeth of large mam-

mals and a few lag boulders, that occurs over much of the valley

bottom. This unit is considered to have been deposited by one or more

floods between ca. 27,000 and 14,500 B.P. Within the channel of the

Saltville River are finer grained, in places well and differentially sorted,

fluvial sediments (Unit W3). These sediments apparently were laid down

over a relatively short period as bed load from moderate fluctuations in

stream stage/ transport capacity, just prior to the piracy of the Saltville

River.

A shallow lake — Lake Totten (McDonald 1985b) — formed fol-

lowing the loss of the Saltville River, and persisted as the dominant

hydrologic feature of the valley throughout most or all of the subse-

quent 14,000 years. Some small streams, mostly spring fed, also proba-

bly entered the valley during this period. As a consequence of this

changed hydrology, fine lacustrine and marsh sediments, primarily mas-

sive clays (Units W2, H2), occur over all of the valley below 530 m in

elevation and extend over an undetermined part of the upper valley

lying above 530 m. The only significant interruption of this clay

sequence is a mud-soil-peat mosaic that formed or was deposited over

part of the valley around 10,500 to 10,000 B.P., at a time when the

water table was lowered in the middle (and upper?) valley. No evidence

of an equally lowered water table has been found in the lower valley. A

shallow lake and marsh of some 200 acres (about 80 square hectometers)

existed in the valley when the first land patents to European settlers

were issued late in the 18th Century (Ogle 1981).

STRATIGRAPHY OF THE SITES

SV-1 lies directly over the southeast side of the Saltville River chan-

nel and, as a result, it contains all of the primary Wisconsinan-Holocene

stratigraphic units recognized to date (Fig. 2). Since excavations began

at this site in October 1980, more than 3,000 vertebrate specimens have

been removed from some 200 m of excavated area. Vertebrate remains

have been found in all stratigraphic units. Most fossils, however, have

been found near the bottom of the lower lake clay (Unit W2: ca. 13,500

B.P.), in the sand and fine gravel deposits of the channel bottom (Unit

W3: ca. 14,500 to 13,500 B.P.), and in the coarser gravel sheet (Unit

W4: ca. 27,000 to 14,500 B.P.).

Most (18 of 26) of the herptile specimens described in this paper

came from SV-1. Twelve specimens were recovered from the channel

bottom deposits (Unit W3), which have also yielded large numbers of

Depth

Surface— I — — —

100cm —

200cm

300 cm

Late Quaternary Herpetofauna

SV-1 SV-2

1

,10,050± 110

M0,690± 130

,13,130 ± 330

14,480 ± 300

J m$ |^J

ffff

UNIT HI

UNIT H2

UNIT Wl

UNIT W2

Fig. 2. Stratigraphic profiles for the three sites that have produced generically

or specifically identified herptile specimens. Radiocarbon dates on Unit Wl are

from samples collected near, but not at, SV-1; those on units W2 and W3 are

from samples collected at SV-1.

90

J. Alan Holman and Jerry N. McDonald

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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.

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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

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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

Fig. 6. " 'Ancestral' Tennessee River" (arrows) of Isphording (1983). (Repro-

duction of his Fig. 10, p. 303.) Stippling delimits Miocene outcrop.

136 J. F. Fitzpatrick, Jr.

less susceptible to weathering, were present in expected amounts. This,

plus other mineralogical considerations, led to an hypothesis that the

period was characterized by a warm, moist climate. A similar suite in

the comparable Pascagoula-Hattiesburg Formation indicates that this

area, too, was far from arid (Isphording 1983). Florida presents a

somewhat different and contradictory picture, but it is outside the con-

siderations of this paper; presumably Florida conditions influenced Alt's

thinking. The conclusions of Isphording, however, are compatible with

the position of Dorf (1960) who envisioned a subtropical or tropical

climate on the Gulf Coast throughout the Tertiary and during intergla-

cial stages of the Pleistocene.

ZOOGEOGRAPHIC IMPLICATIONS

Turning now to animal distribution, we find certain enigmatic fea-

tures. One of these is the Cambarellinae. Every evidence indicates an

early divergence from cambarine stock. Yet the more primitive members

of the genus are found associated with the marginal areas of the Gulf

Coastal Plain. Fitzpatrick (1983) noted that almost every site from

which the genus has been collected in Mississippi (and Florida/ Georgia)

is south and east of Brown's (1967) ridges or on the Mississippi River

flood plain (Fig. 7). Although not as pertinent to this discussion, a sim-

ilar restriction to geologically recent areas of the Coastal Plain in Loui-

siana and Texas exists, with deep inland areas being invaded only in

Mexico.

Fitzpatrick (1983) believed the ancestral cambarellid was most like

Cambarellus puer Hobbs and its relatives; but among the species he

considered as candidates for this status, all are outside the site of origin

for the Cambaridae proposed by Hobbs. Quite clearly, the dwarf craw-

fishes arose from a stock that became established in the lower Missis-

sippi River lowlands shortly after the emergence of the subfamily and

before much diversification of populations began. A temporal assign-

ment of this event is difficult, but it could easily have occurred when

proposed by Hobbs (late Cretaceous or early Cenozoic). Their subse-

quent diversification and expansion east of the Mississippi River delta,

however, could not have occurred before Miocene times. If, as proposed

by Isphording and Flowers (1983), Brown's (1967) ridges represent a

reworking of Miocene deposits, rather than primary deposits, then the

eastward expansion is post-Miocene, probably late Pliocene. Further,

their distributions give a relatively clear indication that no easy access to

lentic habitats of the upper Coastal Plain existed.

On the lower Gulf Coastal Plain, the temporary bodies of water are

dominated by Cambarellus, Faxonella, Procambarus (Capillicambarus),

P. (Scapulicambarus) clarkii, and the ubiquitous, probably multi-species

taxon, P. (Ortmannicus) acutus acutus. All are tertiary burrowers. They

are complemented, often sympatrically, by primary burrowers of Falli-

cambarus, Cambarus (Lacunicambarus), Procambarus (Acucauda), and

P. {Hagenides). The upper Coastal Plain and inland areas have an

Tennessee River and Crawfish Distribution

137

<?>

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

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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