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North Carotina State Library

w. a

number 7

July 1982

EDITORIAL STAFF

John E. Cooper, Editor Alexa C. Williams, Managing Editor John B. Funderburg, Editor-in-Chief

Board

Alvin L. Braswell, Curator of David S. Lee, Chief Curator

Lower Vertebrates, N.C of Birds and Mammals, N.C.

State Museum State Museum

John C. Clamp, Associate Curator William M. Palmer, Chief Curator

(Invertebrates), N.C. of Lower Vertebrates, N.C.

State Museum State Museum

Martha R. Cooper, Associate Rowland M. Shelley, Chief

Curator (Crustaceans), N.C. Curator of Invertebrates, N.C.

State Museum State Museum

James W. Hardin, Department of Botany, N.C. State University

Bnmleyana, 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 in- vertebrate 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 Editorial Board, and ap- propriate specialists will review each paper adjudged suitable. Final ac- ceptability will be decided by the Editor. Address manuscripts and all cor- respondence (except that relating to subscriptions and exchange) to Editor, Bnmleyana, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611.

In citations please use the full name — Bnmleyana.

North Carolina State Museum of Natural History

North Carolina Department of Agriculture

James A. Graham, Commissioner

CODN BRIMD 7 ISSN 0193-4406

Terrestrial Drift Fences With Pitfall Traps:

An Effective Technique for Quantitative

Sampling of Animal Populations

J. Whitfield Gibbons and Raymond D. Semlitsch

Savannah River Ecology Laboratory,

Drawer E, Aiken, South Carolina 29801

ABSTRACT. — The terrestrial drift fence with pitfall traps is a com- monly used technique to collect and quantitatively sample populations of certain vertebrate and invertebrate species. However, a variety of limitations, advantages, biases, and contingencies must be considered to use the method most effectively. The best materials to use for these fences and traps have been aluminum flashing and plastic 20-liter buckets. Aluminum flashing is rigid and does not deteriorate with age. Large plastic buckets permit the capture of many species that can escape from small can traps. Maintenance, such as filling cracks or holes along the fence, bailing water from traps, and mowing vegetation alongside fences, are necessary for continued effectiveness. Initial cost of construction is high, both in time and money; however, drift fences are cost effective for most ecological studies. Biases result primarily from variation in morphology, ecology, and behavior of species, or as a consequence of design and the manner in which the drift fence is checked and maintained.

INTRODUCTION

The terrestrial drift fence and pitfall trap technique has been used for many years for field sampling a variety of vertebrate and inverte- brate species (e.g. Imler 1945; Gloyd 1947; Woodbury 195 1, 1953; Storm and Pimentel 1954; Packer 1960; Husting 1965; Shoop 1968; Hurlbert 1969; Gibbons 1970; Gibbons and Bennett 1974; Briese and Smith 1974; Randolph et al. 1976; Collins and Wilbur 1979; Bennett et al. 1980; Brown 1981; Wygoda 1981). The use of pitfall traps (without drift fences) in the study of invertebrates was reviewed by Mitchell (1963) and Greenslade (1964); however, no thorough assessment of the drift fence technique has been presented (but see Storm and Pimentel 1954). Our purpose is to discuss the advantages as well as the limitations of the approach, using examples from 13 years of data taken on reptiles and amphibians collected on the U. S. Department of Energy's Savan- nah River Plant (SRP) near Aiken, South Carolina.

CONSTRUCTION OF DRIFT FENCES

Basic design for a terrestrial drift fence is a straight fence extending slightly below ground and up to 50 cm high, with pitfall traps placed

Brimleyana No. 7:1-16. July 1981. 1

2 J. Whitfield Gibbons and Raymond D. Semlitsch

alongside the fence and buried flush with the ground at prescribed intervals (Fig. 1 A). The intent is to intercept animals traveling overland so that upon encountering the drift fence they turn left or right and continue along the fence until they fall into a trap.

Fences and traps used on the SRP have undergone a 13-year evolu- tion of construction materials and design (Fig. 1B,C,D). The earliest drift fences, constructed in 1968 of chicken wire, were intended only for cap- ture of turtles moving overland. Twenty-liter (5-gallon) metal paint buckets served as pitfall traps. From 1969 to 1971, hardware cloth (!4- inch mesh) was used for fencing material. We have subsequently found the most effective material to be 50 cm high aluminum flashing, of which approximately 10 cm is placed below the surface of the ground. This has the advantage of preventing small animals from passing under or through the fence, or larger ones from using the mesh to climb over. The flashing is also considered to be superior to various plastic fencing materials used by other investigators (Storm and Pimentel 1954; Packer 1960; Husting 1965; Shoop 1974; Gill 1978; Wygoda 1979; Douglas 1979; Collins and Wilbur 1979) in that it is not easily torn or pushed down by larger vertebrates such as turtles, alligators, feral pigs, or deer. Furthermore, aluminum flashing does not rust or deteriorate with age as do many other commonly used fence materials.

Twenty-liter plastic buckets have proved to be the most effective pitfall traps. These containers are relatively permanent, whereas metal buckets begin to deteriorate within two years, making them less useful for long-term studies. Although smaller volume traps (# 8 cans) have been used effectively for certain species (Shoop 1965; Gill 1978; Douglas 1979), larger traps permit the capture of many species that can easily escape from a shallow can.

MAINTENANCE

Once drift fences and pitfall traps have been constructed, mainten- ance is required for continued effectiveness. Vegetation growing along- side the fence should be mowed or cut to prevent animals from using it to climb over, as well as to assure visibility in checking traps and fence margins. Mowing chores can be reduced in some situations by placing a heavy layer of sand alongside the fence, extending out 5-10 cm.

Cracks or crevasses may develop along the fence or around buckets following construction, particularly after heavy rainfalls. They should be filled to prevent animals from using them as tunnels under the fence. Pitfall traps often fill and overflow with water after heavy rains or from rises in groundwater. Holes drilled in the bottom or sides will prevent water accumulation in some instances, although on certain occasions bailing may be necessary.

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In addition to the hazards of too much water, traps must be checked at frequent intervals to avoid desiccation, predation, and other problems that can arise. In sampling amphibians, desiccation can be avoided at most times of the year by a daily checking schedule. A more frequent checking routine, or use of wet sponges in the bottom of traps, can prevent desiccation or lessen heat stress during summer. Reptiles and mammals are typically more resistant to desiccation and can remain in pitfall traps for longer periods of time than amphibians.

The probability of predation on animals in pitfall traps increases the longer they go unchecked, hence checking at several day intervals would not be desirable in most instances. Daily checking is recom- mended under most circumstances, although more frequent checking (i.e., two or three times a day) may be necessary during mass migration of amphibians from breeding ponds, or when prolonged disruption of the animals' activity is detrimental. Use of an electric wire system designed to safeguard each pitfall trap is effective in deterring most mammalian predators except shrews (C. R. Shoop, pers. comm.).

TIME, COST, AND EFFORT TO CONDUCT A STUDY Drift fences with pitfall traps yield a wealth of biological informa- tion, often providing ecological perspectives that could be obtained in no other manner. However, the cost in labor and materials can be great. The time investment in an effective drift fence operation can be parti- tioned into construction, maintenance, and operation (Table 1). Initial cost of construction is high, both for materials and in time required for installation. After construction, the cost for materials is small; however, the time investment can become onerous if the traps are checked in an effective manner, such as daily (Table 1).

Although the time and effort put into drift fence construction, maintenance, and operation are high, data accumulation is often super- ior to any other form of collecting for a wide variety of terrestrial anim- als, particularly amphibians (Fig. 2; Table 2). The technique is highly cost effective once the critical investment level has been reached. This is especially true for long-term ecological studies where continuous daily hand sampling is impractical.

INTERPRETATION OF RESULTS Drift fences with pitfall traps yield large amounts of data on numbers (often total population sizes), seasonality, migration patterns, diversity, and distribution patterns of many animals (see references in Introduction; Table 2). Some species are collected in high numbers that closely represent the actual population size, whereas the proportional capture of others is below that of their actual abundance. However, as

Terrestrial Drift Fences with Pitfall Traps 5

Table 1. Categorization of time, labor, and expenses for materials for various

stages of drift fence construction, maintenance and operation for two

study sites on the Savannah River Plant.

Category

Sites Rainbow Bay Sun Bay

September	1978	February 1979
440		450
88		90
168		119
$660		$682
0		0
0		0
$10		$10
$80		$80

Date constructed Circumference (m of fencing) Number of pitfall traps Construction costs:

total labor (man hours)

aluminum flashing (@$22/roll)

buckets (obtained at no cost)

stakes (180/ fence obtained at no cost)

plastic cable ties (400/ fence)

shovels, axes, sledge hammer

Maintenance costs (hr/yr):

cut grass around fence 5 5

check and fill cracks and holes,

replace sponges, renumber pitfall

traps, and other routine

maintenance 4 4

Operation (hr/yr):

daily checking of pitfall traps

and processing animals (not

including transportation) 365 365

with any sampling technique, certain biases and limitations must be taken into account in the interpretation of data. Biases are primarily due to variation in morphology, ecology, and behavior of species, or are a consequence of fence design and the manner in which it is checked and maintained.

A species' morphology is an obvious factor in determining the effectiveness of the technique in capturing certain animals. The large body size of some snakes and mammals permits ready escape from the pitfall traps, as does an ability to climb or jump over the fence. Climb- ing or burrowing adaptations, such as toepads on treefrogs or the dig- ging limbs of moles, can reduce the proportion of the population that is actually sampled.

Behavior can also influence the capture of certain species. For example, although many specimens of the eastern box turtle, Terrapene

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Fig. 2. Comparison between hand collecting and drift fence methods. X-axis represents cumulative man-hours (time investment in fence construction not included). The dashed curve for hand collecting indicates cumulative number of specimens of either reptiles or amphibians (specimens/ hour yields were similar). The dashed curve is based on field notes of "best-case" general collecting by experienced herpetologists in typical southeastern Coastal Plain terrestrial habi- tats during spring or summer. Data points on drift fence curves indicate cumula- tive numbers obtained at monthly intervals (approximately 35 hours each) during a complete year of sampling, beginning January 1979 at Rainbow Bay. These data, in constrast to hand collecting, include winter days when reptiles would not ordinarily be sought by hand collectors.

Carolina, have been collected alongside drift fences, few adults have actually fallen into the traps. This is presumably the result of this highly terrestrial species' awareness of topographic relief and an avoidance of natural pitfalls. When some specimens of the Appalachian woodland salamander, Plethodon jordani, encounter a drift fence, they will turn

Terrestrial Drift Fences with Pitfall Traps 11

around and return to their original retreat, and subsequent forays might be in other directions (C. R. Shoop, pers. comm.). Such behavior could bias interpretations of direction preferences.

Ecology of a species is probably the most important factor influenc- ing the rates and patterns of capture. Home range size and migratory movements are critical in certain species. For example, the mole sala- mander, Amby stoma talpoidewn, has a life cycle in which adults, under most conditions, characteristically migrate to a breeding pond during winter and return to land in early spring. Juveniles exit the pond a few months later (Patterson 1978; Semlitsch 1981). Therefore, any fence placed parallel to the edge of the breeding pond will capture most, if not all, salamanders moving through the area sampled.

Southeastern crowned snakes, Tantilla coronata, or scarlet snakes, Cemophora coccinea, may occur in the same habitat as A. talpoidewn. However, individual home ranges in these two species of snakes are independent of the orientation and distance to water, so that movement primarily represents daily activity patterns. Although either species of snake may be captured in relatively large numbers in pitfall traps (Nelson and Gibbons 1972; Semlitsch et al. 1981), the drift fence will only reveal that part of a population whose home ranges overlap the fence system. Thus, whether or not a study species has a congregating focal point as part of its life cycle will influence the effectiveness of the technique in assessing population size.

Similarly, use of drift fences around an aquatic habitat to monitor terrestrial movement of turtles will result in capture of a higher propor- tion of some species than others. The propensity of eastern mud turtles, Kinosternon subrubrwn, to overwinter on land (Bennett et al. 1970) means that they are more likely to migrate through the land-water inter- face than is a more aquatic species such as the stinkpot, Sternotherus odoratus. Undoubtedly, there are other subtle, important ecological dif- ferences among species that affect their respective rates of capture in similar manners.

Certain false impressions about abundance, diversity or behavior of animals in an area can be given by factors related to design of the tap- ping system, in combination with the ecology of the species involved. One of the most important design factors may be the spatial arrange- ment of the drift fences (Fig. 1B,C,D). Distance of a fence from a critical habitat, such as an aquatic breeding site or a terrestial denning or nest- ing area, can greatly influence the number of captures of certain species. The key factor is whether the fence intercepts the path of migration, or other movement, of animals from one site to another. A potential impact of fence placement can be readily seen in the disparity in numbers of certain species that leave or enter from particular directions

12 J. Whitfield Gibbons and Raymond D. Semlitsch

in a habitat. For example, a partial fence could lead to misinterpreta- tions about the numbers of some species (e. g., K. subrubrum) that leave or enter the aquatic area. Extrapolation errors would be less likely to result from other species (e.g., S. odoratus) that appear to use the perimeter in a more uniform manner (Table 3).

Table 3. Directional disparities that could result from drift fence placement if partial fencing is used in or around a habitat. Numbers are based on total captures of semi-aquatic turtles in pitfall traps on either side of the fence encircling Ellenton Bay, South Carolina, from 1975 to 1981. The perimeter was arbitrarily partitioned into the four compass directions for Chi-square contingency analysis.

Species	North	East	South	West	Chi-square value
Chelydra serpentina	28	34	24	24	1.29
Sternotherus odoratus	43	40	52	47	0.98
Pseudemys floridana	33	58	44	95	10.86*
Deirochelys reticularia	101	249	119	273	60.34 **
Pseudemys scripta	162	365	269	228	80.87 **
Kinosternon subrubrum	801	498	204	253	305.48 **
TOTAL	1168	1244	712	920	77.69 **

* P<.05 ** P< .01

The temporal aspect is also critical, not only at the seasonal level but in some instances on a daily basis (e. g. Hurlbert 1969; Gibbons 1970; Semlitsch et al. 1981). Long-term studies reveal that annual dis- parities can be great enough to provide the potential for improper inter- pretations if drift fences are used to sample habitats for short periods of time (Gibbons and Bennett 1974: Table 4).

Because of the factors discussed above, a well-constructed and maintained drift fence with pitfall traps will be effective in capturing most individuals of certain species in an area, and none of others. The outstanding number of captures of A. talpoideum, as well as the high recapture rate, suggest that the method is highly effective for this pond- breeding, migratory species (Table 5). On the other hand, relatively few adult black racers, Coluber constrictor (Table 2), have been captured in pitfall traps on the SRP, although the species is very abundant in the areas under study (Gibbons and Patterson 1978). Not surprisingly, these and other large snakes easily escape from the traps. The same is true of

Terrestrial Drift Fences with Pitfall Traps

13

Table 4. Annual and local variation in total captures of adult amphibian and reptile species commonly sampled in drift fences on the Savannah River Plant, South Carolina. Each sampling year began in September and continued through August.

Site and		Sampling year
species	1978-79	1979-80	1980-81
Rainbow Bay
Notophthalmus vihdescens	1,271	1,058	772
Scaphiopus holbrooki	51	10	45
Rana utricular ia	508	346	475
Kinosternon subrubrum	53	24	23
Cnemidophorus sexlineatus	1	1	3
Tantilla coronata	15	28	8

Sun Bay

Notophthalmus viridescens Scaphiopus holbrooki Rana utricularia Kinosternon subrubrum Cnemidophorus sexlineatus Tantilla coronata

1,757 1,271

2,745 756

728 12

99*	79	35
87*	16	19
19*	29	12
40*	50	10

* Minimum estimate

certain species of treefrogs (Hyla), which can climb the sides of a bucket or a fence (Gibbons and Bennett 1974). For many large mammals (e. g. raccoon, opossum), no adults have ever been captured in the traps. However, those species for which the technique is either always or never effective are not the primary problems. The major difficulty in interpre- tation and analysis of data from drift fences results from those species whose captures only partly reflect the numbers of individuals that actu- ally encounter the fence or live in the vicinity. Unless the effectiveness or sampling rate is known, certain conclusions relating to population size or absolute abundance should be drawn with caution. However, the potential for using the technique to estimate larval survivorship, immi- gration and emigration rates, genetic exchange, and other difficult-to- obtain data, has been demonstrated (Gibbons 1970; Shoop 1974; Gill 1978; Semlitsch and McMillan 1980; Semlitsch 1981) and should not be underestimated.

Merely revealing the presence of a rare species can be a contribu- tion to an understanding of its basic biology. Star-nosed moles, Con- dylura cristata, have been infrequently captured in pitfall traps on the

14 J. Whitfield Gibbons and Raymond D. Semlitsch

Table 5. The number of drift fence captures and recaptures of adult Ambystoma talpoideum entering and exiting Rainbow Bay, South Carolina. The percentages indicate the increasing effectiveness of the technique for this species as a greater proportion of the population is collected.

Sampling year 1978-79 1979-80 1980-81

Total number entering (marked and unmarked) 459 2,133 450

Total number exiting

Already marked 193 836 264

Unmarked 70 200 19

Percent of already marked individuals

of those entering 42% 39% 59%

Percent of unmarked individuals exiting

of those entering 15% 9% 4%

Sampling error (based on assumption that 100%

of specimens on inside of fence were marked) 27% 19% 7%

SRP. However, the 14 specimens captured represent a major sample compared to those previously reported from either South Carolina or Georgia in the previous century (Golley 1962, 1966). The findings sug- gest that the species is not necessarily rare or restricted in its geographic range or habitat preference, but is merely difficult to capture by conven- tional trapping methods.

Another form of bias is that adults of certain species may not be captured in pitfall traps, although the smaller juveniles may be suscepti- ble and reveal an unexpected abundance. This phenomenon was wit- nessed with the rainbow snake, Farancia erytrogramma, in which more than 100 subadult animals were captured in an area where large adults have never been seen (Gibbons et al. 1977). Such captures must be interpreted cautiously, but their value for certain purposes is obvious.

CONCLUSIONS

Drift fences are capable of collecting large amounts of data on a daily basis over long periods of time (> 5 years). For some species they provide the only effective sampling technique, and for many it is highly cost effective. However, variation in morphology, ecology, and behavior of each species must be considered. If the limitations and biases of the drift fence and pitfall trap technique are considered, population sizes, seasonality, migration patterns, diversity, and distribution of many spe- cies of animals can be effectively determined.

ACKNOWLEDGMENTS.— We are grateful to the more than 400 individuals who have worked at or visited the Savannah River Ecology

Terrestrial Drift Fences with Pitfall Traps 15

Laboratory and helped install, maintain, and check drift fences and pit- fall traps. Special debts of gratitude go to L. B. Wright, D. H. Bennett, D. H. Nelson, J. W. Coker, T. M. Murphy, K. L. Brown, L. A. Briese, S. H. Bennett, J. L. Greene, K. K. Patterson, G. B. Moran, C. A. Shoe- maker, J. P. Caldwell, and many others. Data for comparing the drift fence technique with that of conventional hand-collecting methods was made possible through field notes contributed by R. D. Aldridge, J. R. Harrison III, D. W. Herman, D. W. Tinkle, and L. J. Vitt, as well as our own. We thank S. H. Bennett, K. L. Brown, A. C. Lamb III, and C. R. Shoop for constructive comments on the manuscript.

Research was conducted under contract EY-76-C-09-0819 between the U. S. Department of Energy and the University of Georgia, and under NSF Grant NO. DEB 7904758.

LITERATURE CITED Bennett, David H., J. W. Gibbons and J. C. Franson. 1970. Terrestrial activity

of aquatic turtles. Ecology 5/(4):738-740. Bennett, Stephen H., J. Glanville and J. W. Gibbons. 1980. Terrestrial activity,

abundance, and diversity of amphibians in differently managed forest types.

Am. Midi. Nat. /0J(2):412-416. Briese, Linda A., and M. H. Smith. 1974. Seasonal abundance and movement

of nine species of small mammals. J. Mammal. 55(3):6 15-629. Brown, Kent L. 1981. An analysis of species diversity along a temporal gradient

of loblolly pine stands in South Carolina. Unpubl. M.S. Thesis, Texas

Christian Univ., Fort Worth. 44 pp. Collins, James P., 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. 686A-34. Douglas, Michael E. 1979. Migration and sexual selection in Ambystoma jef-

fersonianum. Can. J. Zool. 57(1 2):2303-23 10. Gibbons, J. Whitfield. 1970. Terrestrial activity and the population dynamics of

aquatic turtles. Am. Midi. Nat. #5(2):404-414. , and D. H. Bennett. 1974. Determination of anuran terrestial activity

patterns by a drift fence method. Copeia 1974(1 ):236-243. , and K. K. Patterson. 1978. The reptiles and amphibians of the

Savannah River Plant. SRO-NERP-2, Aiken, SC. 24 pp. , J. W. Coker and T. M. Murphy, Jr. 1977. Selected aspects of the life

history of the rainbow snake {Farancia erytrogramma). Herpetologica

53(3):276-281. Gill, Douglas E. 1978. The metapopulation ecology of the red-spotted newt,

Notophthalmus viridescens (Rafinesque). Ecol. Monogr. 48(2): 145-166. Gloyd, Howard K. 1947. Some rattlesnake dens of South Dakota. Chicago Nat.

9:87-97. Golley, Frank B. 1962. Mammals of Georgia. Univ. Georgia Press, Athens. 218

pp. 1966. South Carolina Mammals. Charleston Museum, Charleston.

181 pp.

16 J. Whitfield Gibbons and Raymond D. Semlitsch

Greenslade, P. J. M. 1964. Pitfall trapping as a method for studying populations

of Carabidae (Coleoptera). J. Anim. Ecol. 3J(2):301-310. Hurlbert, Stuart H. 1969. The breeding migrations and interhabitat wandering

of the vermilion-spotted newt Notophthalmus viridescens (Rafinesque).

Ecol. Monogr. 39(4):465-488. Husting, E. L. 1965. Survival and breeding structure in a population of

Ambystoma maculatum. Copeia 1965(3):352-362. Imler, Ralph H. 1945. Bullsnakes and their control on a Nebraska wildlife

refuge. J. Wildl. Manage. 9(4):265-273. Mitchell, B. 1963. Ecology of two carabid beetles, Bembidion lampros (Herbst)

and Trechus quadristriatus (Schrank). II. Studies on populations of adults

in the field, with special reference to the technique of pitfall trapping. J.

Anim. Ecol. 52(3):377-392. Nelson, David H., and J. W. Gibbons. 1972. Ecology, abundance, and seasonal

activity of the scarlet snake, Cemophora coccinea. Copeia 1972(3):582-584. Packer, Wayne C. 1960. Bioclimatic influences on the breeding migration of

Taricha rivularis. Ecology 4/(3):509-5 17. Patterson, Karen K. 1978. Life history aspects of paedogenic populations of the

mole salamander, Ambystoma talpoideum. Copeia 1978(4):649-655. Randolph, J. Collier, P. A. Randolph and C. A. Barlow. 1976. Variation in

energy content of some carabid beetles in eastern Canada. Can. J. Zool.

54(1):10-18. Semlitsch, Raymond D. 1981. Terrestrial activity and summer home range of

the mole salamander (Ambystoma talpoideum). Can. J. Zool. 59(2):3 15-322. , and M. A. McMillan. 1980. Breeding migrations, population size

structure, and reproduction of the dwarf salamander, Eurycea quadridigi-

tata, in South Carolina. Brimleyana 3:97-105. K. L. Brown and J. P. Caldwell. 1981. Habitat utilization, seasonal

activity, and population size structure of the southeastern crowned snake

Tantilla coronata. Herpetologica 57(l):40-46. Shoop, C. Robert. 1965. Orientation of Ambystoma maculatum: movements to

and from breeding ponds. Science 749(3683):558-559. 1968. Migratory orientation of Ambystoma maculatum: movements

near breeding ponds and displacements of migrating individuals. Biol. Bull.

735:230-238. 1974. Yearly variation in larval survival of Ambystoma maculatum.

Ecology 55(2):440-444. Storm, Robert M., and R. A. Pimentel. 1954. A method for studying amphibian

breeding populations. Herpetologica 10(3): 161-166. Woodbury, Angus M. 1951. Symposium: a snake den in Tooele County, Utah.

Introduction — a ten year study. Herpetologica 7(1):4-14.

1953. Methods of field study in reptiles. Herpetologica 9(2):87-92.

Wygoda, Mark L. 1979. Terrestrial activity of striped mud turtles, Kinosternon

baurii (Reptilia, Testudines, Kinosternidae) in west-central Florida. J.

Herpetol. /5(4):469-480. 1981. Notes on terrestrial activity of the crayfish Procambarus alleni

(Faxon) in west-central Florida. Fla. Sci. 44:56-59.

Accepted 22 April 1982

Marine and Freshwater Fishes of the Cape Fear

Estuary, North Carolina, and Their Distribution in

Relation to Environmental Factors

Frank J. Schwartz

Institute of Marine Sciences,

University of North Carolina,

Morehead City, North Carolina 28557

William T. Hogarth

Carolina Power and Light Company, New Hill, North Carolina 27562

AND

Michael P. Weinstein

Department of Biology, Virginia Commonwealth University, Richmond, Virginia 23284

ABSTRACT.— A survey of the saline lower Cape Fear River watershed, conducted from 1973 through 1980, yielded 12,612,022 fishes of 249 species in 85 families. This list is supplemented by an additional three families and seven species reported in the literature. A mixture of Virginian, Carolinian, Caribbean, resident, transient, and estuarine-dependent fishes seasonally occupied the study area, which included parts of the Northeast Cape Fear River, the Cape Fear River downstream of Lock 1, and the adjacent Atlantic Ocean off Baldhead and Oak islands. Collections were made with beach seine, rotenone, gill net, plankton net, otter trawl, and traveling screens of a nuclear steam electric power plant located 2.4 km upstream of Southport, North Carolina. Fish presence and abundance were recorded and related to water temperature, dissolved oxygen, and salinity. Resident or seasonal status of each species was also related to spatial habitat distribution, substrate preference, freshwater and marine intrusion, or geographic province origin. The river serves as a nursery area for many estuarine-dependent fishes. Although industrial and domestic pollution enter the river, primarily at Wilmington, North Carolina, they have not as yet seriously affected its vitality for fish survival.

INTRODUCTION

Early ichthyological studies by Lawson (1714), Catesby (1771), and Smith (1907) largely ignored the marine fish fauna of the lower Cape Fear River, North Carolina. By contrast, the freshwater fishes of the system have been well documented by Fowler (1945) and Menhinick et al. (1974). Brimley (1935) and Gudger (1948) reported strandings and

Brimleyana No. 7:17-37. July 1981. 17

18 Frank J. Schwartz, William T. Hogarth, Michael P. Weinstein

occurrences of the whale shark. Rhincodon typhus, and the basking shark, Cetorhinus maximus, respectively, from the river near its mouth. Attempts to survey the estuarine tributaries to the lower river were made by Bayless (1963) and Louder (1963). Lawler (1975) sampled the river to near Campbell Island (Buoy 42) and others have reported spo- radic occurrences of marine or freshwater fishes within the estuary (Lindquist et al. 1977; Lindquist et al. 1978; Martin and Shipp 1971; Tucker and Hodson 1976; Wade 1962). Myers (1925) described the freshwater minnow Notropis cummingsae from the Cape Fear River at Wilmington. Commercially important species, such as the shads, were studied by Davis and Cheek (1967), Nichols and Louder (1970), and Walburg and Nichols (1967), usually in relation to Lock 1 on the main Cape Fear River northwest of Wilmington. Huish and Benedict (1977) tracked the movements of several sonic tagged dusky sharks, Carchar- hinus obscurus, near Buoy 19 in the lower river. Jenkins (1970) cor- rected the systematics and occurrences of suckers reported by Bayless (1963) for the lower river. Copeland and Birkhead (1973) and Birkhead et al. (1977, 1979) focused their studies primarily in the Dutchman Creek area west of Southport, North Carolina.

Environmental impact studies associated with the operation of a nuclear steam electric power plant on the Cape Fear River near South- port permitted the first extensive survey of the marine and freshwater fish fauna of the saline portions of the Cape Fear watershed. The sur- vey was conducted between 1973 and 1980. Some results were docu- mented by Schwartz and Dahlberg (1978), Schwartz et al. (1979, 1979), Sulak et al. (1979), and Weinstein et al. (1980, 1980). Additional results are reported here. Fish presence and abundance were related to water temperature, dissolved oxygen, and salinity (see Schwartz et al. 1979, 1979). Additional data on tidal movements, periodicity, and flow strength, which would be experienced by fishes once they are in or near the system, were documented by Carpenter and Yonts (1979), Schwartz and Chestnut (1973), and Welch and Parker (1979). We review and cor- rect information on occurrences of several fishes in the system (Table 1).

STUDY AREA

The Cape Fear River lies entirely within North Carolina and is the largest Atlantic river drainage system in the state. It is about 528 km long with a drainage basin of about 14,553 km2. The estuarine portion accounts for 880 km2 of the system. Spring runoff varies from lows of 0.5 to highs of 3.1 m/sec (Schwartz and Dahlberg 1978; Schwartz et al. 1979, 1979) and is often influenced by tropical hurricanes, inland storms, rain, and snow melt from inland rivers. All or most of the estu- ary is subject to tidal excursions of ±2 m, as a result of strong tides and

Fishes of Cape Fear Estuary 19

southwest winds during nine months of the year.

The estuary is 0.3 km wide at Wilmington, widens to 2.1 km at Snows Cut and 2.0 km at its mouth. River discharge, which once passed through Corn Cake Inlet, has been diverted via a dredged channel to enter the sea between Baldhead Island and Caswell Beach, just east of Southport (Carpenter and Yonts 1979). The present dredged river chan- nel varies from 7.6 to 13.6 m deep between the estuary mouth and Wil- mington. North and west of Wilmington, the Northeast Cape Fear and Cape Fear rivers shoal to 3.7 and 3.6 m, respectively. The main ship channel has a mud-clay substrate throughout most of its length, with silting occurring north of Wilmington. Shoals exist on either side of the ship channel. From Wilmington south to Buoy 42 the narrow east and west shoals, on either side of the channel, are composed of sandy-mud. South of Snows Cut eastern shoals are typically sandy while mud domi- nates western shoals. Extensive mud flats are present in the vicinity of the mouth of the river at Caswell Beach. A Pleistocene sandstone and the porous Castle Hayne formation creates a series of rock ledges that pass from southwest to northeast across the river at Buoy 18 and pro- duce a sill on the east side of the river near the buoy.

Our study area included all major tidal tributaries and the North- east Cape Fear River from 36 km north of Wilmington to near Buoy 46; the main Cape Fear River from Lock 1 (63 km northwest of Wilming- ton); and the main river south of Wilmington (including the man-made Snows Cut channel), 24 km south of Wilmington on the east side of the river to its junction with the Atlantic Ocean (Fig. 1). The south- western boundary of the study area was Dutchman's Creek, 3.6 km west of the Intracoastal Waterway at Southport. The seaward boundary was the Atlantic Ocean and encompassed an area from the eastern tip of Baldhead (Smith) Island on the east, westward to Long Beach, North Carolina, and for 8 km offshore to depths of 12 m.

MATERIALS AND METHODS

Schwartz et al. (1979, 1979) employed 91.4 m gill nets to sample pelagic species during January through November 1973-1980. Otter trawls of 7.6 and 12.2 m were used to sample all other fishes at shoal, channel, power plant intake canal, and ocean stations during the same period. Weinstein et al. (1980b) employed rotenone and beach seines at tidal creek stations throughout the saline portion of the Cape Fear. Details of sampling methodology were cited in Weinstein et al. (1979). One excellent and efficient continuous sampling device was the traveling screens located at the head of an intake canal at the power plant, which were cleaned several times daily. Three species were obtained by North Carolina State University (Copeland et al. 1979) and Carolina Power

20 Frank J. Schwartz, William T. Hogarth, Michael P. Weinstein

Fig. 1. Fish sampling sites, Cape Fear River and adjacent Atlantic Ocean, 1973- 1980.

Fishes of Cape Fear Estuary 21

and Light Company personnel using meter (505/u mesh) plankton sam- pling efforts conducted throughout the study area.

Voucher specimens are deposited in the curated collection of the Institute of Marine Sciences, University of North Carolina. Specific fish names follow Robins et al. (1980). Exceptions are: Dawson's (1979) sub- specific designation for the opposum pipefish as Oostethus brachyurus lineatus, and Tyler's (1980) definition of Monocanthus and the family Monocanthidae. Deckert's (1973) and Miller's (1976) revisions of Diap- terus are adopted. Contrary to Fahay and Obenchain (1978), we follow McCosker's (1977) placement of the genus Pisodonophis in Ophichthus. We also recognize Ophidion welshi (Nichols and Breder 1922) rather than Rissola marginata (= Ophidion marginatus, Cohen and Nielsen 1978) as the most abundant cusk-eel inhabiting the river.

Rare, common, and abundant fishes are defined as species known from 1, 2 to 10, or more than 10 specimens, respectively. The terms Virginian, Carolinian, and Caribbean refer to geographic provinces along the western Atlantic coast (Briggs 1974). Virginian is that area usually located north of Cape Hatteras; Carolinian is the Continental Shelf area adjacent to the coast between Cape Hatteras and the tip of Florida; and Caribbean is that offshore area seaward of the Carolinian area and located over warm waters of great depths adjacent to the Gulf Stream. Substrate designations are considered sand, mud, clay, muddy- sand or sandy-mud, depending on visual inspection of the amount of mud or sand comprising a Shipek grab at each study station.

RESULTS AND DISCUSSION

Physical and Chemical Parameters. — Water temperatures and salinities usually ranged from 4-26° C and 0-8 ppt in the river near Buoy 42, 3.7-29° C and 3-30 ppt southward of Wilmington to Snows Cut, 5-31.8° C and 10-34 ppt from Snows Cut to Buoy 19 (5.5 km north of Southport), and 5.6-30.7° C and 20-36 ppt in the river south of Buoy 19 and including the nearby Atlantic Ocean (Schwartz 1979, 1979). Waters in the Intracoastal Waterway west of Southport, which are of lower river origin, mirrored water temperatures and salinities usually noted for the river near Southport.

Oxygen content was usually more than 4 ppm. On only one occa- sion was 0 ppm oxygen content noted — for a patch of water that extended from the Intracoastal Waterway side of Caswell Beach, around the island, and along the outer beach southwestward for 2 km.

Fishes.— During 1973-1980, 12,612,022 fishes of 249 species and 85 families were collected throughout the lower Cape Fear estuary (Table 2). Additional fishes recorded in the literature supplement our list by three families and seven species. As expected, several of these

22

Frank J. Schwartz, William T. Hogarth, Michael P. Weinstein

species were freshwater fishes that had been flushed or moved into the river usually following extensive rain and freshwater incursions. Schwartz (1981) provided a list of such fishes. Anadromous fishes ascending the river to spawn in inland freshwaters were: American shad, Alosa sapidissima; alewife, Alosa pseudoharengus; blueback herring, Alosa aestivalis', and hickory shad, Alosa mediocris. The diadromous Americal eel, Anguilla rostrata, was widely distributed within the estu- ary, whereas the conger eel, Conger oceanicus, was usually collected in its higher saline parts. The bulk of the fishes collected were estuarine- dependent forms (Clark et al. 1974), such as spot, Leiostomus xanthu- rus; Atlantic croaker, Micropogonias undulatus; weakfish, Cynoscion regalis\ flounders, Paralichthys spp.; mullets, Mugil spp., and others that usually spawned offshore but used various areas of the estuary as a nursery ground (Weinstein 1979; Weinstein et al. 1980, 1980).

Adults of many species overwintered within the system (Atlantic menhaden, Brevoortia tyrannus; spot; etc.), or passed north and south (cownose ray, Rhinoptera bonasus; cobia, Rachycentron canadum), during spring and fall migrations. The scalloped hammerhead, Sphyrna lewini, and spiny dogfish, Squalus acanthias, also exhibited inshore or offshore seasonal migrations. A few fishes (gobies; blennies; bay anchovy, Anchoa mitchilli; killifishes, Fundulus spp.; and silversides, Menidia spp.) were residents. Several species were sporadic captures that either entered the river in association with sargassum (yellow chub, Kyphosus incisor; Bermuda chub, Kyphosus sectatrix; planehead file-

Table 1 . Species incorrectly reported from the Cape Fear River estuary.

Species

Achirus fascia tus Achirus lineatus Calamus leucosteus Dasyatis violacea Diapterus olisthostomus Etheostoma nigrum

Etropus microstomus Eucinostomus lefroyi Fundulus notti

Gerres cinereus Gobionellus oceanicus Moxostoma pappillosum

Rissola marginata

Probable Species

Trinectes maculatus Trinectes maculatus Calamus sp. * Dasyatis sabina Diapterus auratus Etheostoma olmstedi

Etropus crossotus Eucinostomus argenteus Fundulus lineolatus

Diapterus auratus Gobionellus boleosoma Moxostoma anisurum

Ophidion welshi

Reference

Smith 1907

Louder 1963

Smith 1907

Lawler 1975

Birkhead et al. 1977, 1979

Bayless 1963; Louder

1963; Smith 1907

Lawler 1975

Bayless 1963; Copeland

and Birkhead 1973

Lawler 1975

Louder 1963

Bayless 1963; Smith

1907; Jenkins 1970

Lawler 1975

* Mounted specimen at N. C. State Museum, fell apart 24 September 1975; identification doubtful.

Fishes of Cape Fear Estuary 23

fish, Stephanolepis hispidus; and gray triggerfish, Balistes capriscus) or with intrusion of offshore waters (bearded brotula, Brotula barbata; margined snake eel, Ophichthus cruentifer; and offshore tonguefish, Symphurus civitatus). The O. cruentifer was a larva that, like menhaden larvae (Nelson et al. 1977), apparently had been carried inshore by cur- rents from off the edge of the Continental Shelf, where the species has been reported by Fahay and Obenchain (1978). A limited number of species, such as the pink wormfish, Microdesmus longipinnis, were range extensions from nearby southern habitats (Hammond 1973).

Most common or abundant fishes occurred over wide expanses of the estuary and within wide ranges of water temperatures, dissolved oxygen, and salinity regimes (Table 2). Schwartz (1981) recorded 77 marine fishes that often remained for up to six weeks in 0 ppt salinity intrusion waters. Weinstein et al. (1980) attributed postlarval population oscillations of three fishes — spot, Atlantic croaker, and flounder — to rapid salinity changes in the Buoy 42 area.

As waters warmed during summer, southern warm-water Caroli- nian or Caribbean province species were collected (Spanish mackerel, Scomberomorus eavalla; etc.). Conversely, northern smooth and spiny dogfish, Mustelus canis and Squalus acanthias, of Virginian affiliation were present in the river south of Buoy 18 and especially off Caswell Beach as waters cooled. The only cold-related fish kills occurred during the severe winters of 1976 and 1977 (January-February), when many striped mullet, Mugil cephalus; weakfish; and red drum, Sciaenops ocel- latus, were found dead or dying in the power plant intake canal or on shoals that had iced over.

Gobies, blennies, and the clingfish, Gobiesox strumosus, were usu- ally associated with mud-oyster habitats of higher saline areas of the river. The crested cusk-eel, Ophidion welshi, was most commonly col- lected in dense mud substrates in the dug intake canal, especially during 1973-1976, although it was also collected from the mud-clay main chan- nel and occasionally in sandy habitats. The star drum, Stellifer lanceola- tus, was most abundant in late summer in the mud-clay substrate of the main channel, from Buoy 19 southward. The banded drum, Larimus fasciatus, was a high-saline form associated with sandy substrates at ocean stations and in the river south of Buoy 18. The pink wormfish was found only in the intake canal and at the lower river channel Buoy 18 and sandy shoal 18E stations. The white catfish, Ictalurus catus, rarely strayed southward from freshwater mud habitats near Buoy 42. Most of the remaining species were broadly distributed and exhibited no preference for specific substrates.

The estuarine part of the Cape Fear River possesses a large variety of Virginian, Carolinian, and Caribbean province fishes that are resident,

24

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transient and estuarine-dependent (Schwartz et al. 1979, 1979). It thus serves as an important avenue for seasonal migrations of many species along, into, or away from the area, and is vital as a nursery area for the many fishes that depend on estuaries for growth and survivial for part or all of their lives. Although industrial and domestic pollution enter the river, primarily at Wilmington, and may have an indirect effect on fishes such as herrings (Schwartz 1979), they have not as yet seriously affected its vitality for fish survival.

ACKNOWLEDGMENTS.— We thank the field and laboratory per- sonnel of the University of North Carolina, Institute of Marine Scien- ces; Carolina Power and Light Company; and Lawler, Matusky, and Skelly for their aid with many aspects of this project. Dr. R. Hodson of North Carolina State University provided several larval specimens. The comments of several anonymous reviewers are appreciated. Funding for the project was provided by Carolina Power and Light Company, Raleigh, North Carolina, for which we are grateful. Brenda Bright typed the manuscript. Jane Law and William Link of the Institute of Marine Sciences provided Figure I.

LITERATURE CITED

Bayless, Jack D. 1963. Survey and classification of the Northeast Cape Fear

River and tributaries, North Carolina. N. C. Wildl. Resour. Comm.,

Raleigh. 15 pp. + appdx. 30 pp. Birkhead, William A., C. R. Benett, E. C. Pendleton and B. J. Copeland. 1977.

Nursery utilization of the Dutchman Creek estuary, N. C. 1971-1976.

Carol. Power & Light Co. Rep. 77-2. 1 16 pp. , B. J. Copeland and R. G. Hodson. 1979. Ecological monitoring in

the lower Cape Fear River estuary 1971-1979. Carol. Power & Light Co.

Rep. 79-1. 292 pp. Briggs, John C. 1974. Marine Zoogeography. McGraw-Hill Book Co., N.Y. 475

pp. Brimley, H. H. 1935. Notes on the occurrence of a whale shark (Rhincodon

typus) in the Cape Fear River, near Southport, N. C. J. Elisha Mitchell Sci.

Soc. 57:160-162. Carpenter, James H., and W. L. Yonts. 1979. Dye tracer and current meter

studies, Cape Fear estuary, N. C. 1976, 1977, and 1978. Carol. Power &

Light Co. BSEP Studies, vol. 1. 331 pp. + appdx. 227 pp. Catesby, Mark. 1771. The natural history of Carolina, Florida, and the

Bahama Islands. 2 vol. 3rd ed. London. Clark, John, W. G. Smith, A. W. Kendall, Jr. and M. P. Fahay. 1974. Studies

of estuarine dependence of Atlantic coastal fishes. Data Rep. 2, Southern

Section: New River Inlet, N. C. to Palm Beach, Florida, RV DOLPHIN

Fishes of Cape Fear Estuary 35

cruises 1967-68; zooplankton, volume, surface meter net collections,

temperatures, and salinities. Tech. Rep. Bur. Sport Fish. Wildl. 59. 97 pp. Cohen, Daniel M., and J. G. Nielsen. 1978. Guide to the identification of genera

of the fish order Ophidiiformes with a tentative classification of the order.

NOAA Tech. Rep. NMFS Circ. 417. 72 pp. Copeland, Billy J., and W. S. Birkhead. 1973. Baseline ecology of the lower

Cape Fear River estuary and ocean off Oak Island, N. C. 1971-72. Contr.

33 Pamlico Mar. Lab. N. C. State Univ. 391 pp. , R. G. Hodson and R. J. Monroe. 1979. Larvae and postlarvae in the

Cape Fear River estuary, N. C. during operation of the Brunswick Steam

Electric Plant, 1974-78. Rep. 79-3 N. C. State Univ., Raleigh. 226 p. Davis, James R., and R. B. Cheek. 1967. Distribution, food habits, and growth

of young Clupeids, Cape Fear River system, North Carolina. Proc. 20th

Annu. Conf. Southeast Assoc. Game Fish Comm. 1966:250-258. Dawson, Charles E. 1979. Review of the polytypic Doryrhamphine pipefish

Oostethus brachyurus (Bleeker). Bull. Mar. Sci. 29(4):465-480. Deckert, Gary D. 1973. A systematic review of the genera Diapterus and

Eugerre: with the description of a new genus, Schizopterus (Pisces: Gerrid-

ae). Unpubl. M.S. Thesis, Northern 111. Univ., Dekalb. 74 pp. Fahay, Michael P., and C. L. Obenchain. 1978. Leptocephali of the Ophichthid

genera Ahlia, Myrophis, Ophichthus, Pisodonophis , Callechelys, Lethar-

chus, and Apterichtys on the Atlantic Continental Shelf of the United

States. Bull. Mar. Sci. 25(3):442-486. Fowler, Henry W. 1945. A study of the fishes of the southern Piedmont and

Coastal Plain. Monogr. 7 Acad. Nat. Sci. Phila. 408 pp. Gudger, Eugene W. 1948. The basking shark Cetorhinus maximus on the North

Carolina coast. J. Elisha Mitchell Sci. Soc. 64(\):4\-44. Hammond, Donald D. 1973. A record of Microdesmus longipinnis (Weymouth)

(Pisces: Microdesmidae) from South Carolina waters. J. Elisha Mitchell

Sci. Soc.59(l-2):72-73. Huish, Melvin T., and C. Benedict. 1977. Sonic tracking of dusky sharks in the

Cape Fear River, North Carolina. J. Elisha Mitchell Sci. Soc. 93(l):21-26. Jenkins, Robert E. 1970. Systematic studies of the Catostomid fish tribe Mox-

ostomatini. Unpubl. Ph.D. dissert., Cornell Univ., Ithaca. 799 pp. Lawler, John. 1975. Environmental impact assessment of alterations for the

maintenance of Wilmington harbor, North Carolina, pp. A1-A242, Aquatic

Ecology Studies Cape Fear River Estuary, North Carolina, September 1972

to August 1973, appdx. A. Lawler, Matusky, Skelly Engineers, Tappan, NY. Lawson, John. 1714. The history of Carolina; containing the exact description

and natural history of that country: etc. London. 258 pp. Reprinted 1903

by Observer Printing House, Charlotte, NC. 172 pp. Lindquist, David G., J. R. Shute and P. W. Shute. 1977. Record of bluefin

killifish Lucania goodei in North Carolina. J. Elisha Mitchell Sci. Soc.

95(1): 19-20. , and Recent record of the banded killifish,

Fundulus diaphanus (Lesueur) in southeastern North Carolina. J. Elisha

Mitchell Sci. Soc. 94(2): 113.

36 Frank J. Schwartz, William T. Hogarth, Michael P. Weinstein

Louder, Darrell E. 1963. Survey and classification of the Cape Fear River and

tributaries of North Carolina. N. C. Wildl. Resour. Comm., Raleigh. 20 pp.

+ appdx. 95 pp. Martin, James R., and R. L. Shipp. 1971. Occurrence of juvenile snook, Centro-

pomus undecimalis, in North Carolina waters. Trans. Am. Fish. Soc.

700(1): 131-132. McCosker, John E. 1977. The osteology, classification, and relationships of the

eel family Ophichthidae. Proc. Calif. Acad. Sci. Ser. 4, 41(1): 1-123. Menhinick, Edward F., T. M. Burton and J. R. Bailey. 1974. An annotated

checklist of the freshwater fishes of North Carolina. J. Elisha Mitchell Sci.

Soc. 90(1):24-5O. Miller, Robert R. 1976. Geographical distribution of Central American fresh- water fishes, pp. 125-156 in T. B. Thorson (ed.). Investigation of the Ich-

thyofauna of Nicaraguan lakes. Univ. Nebraska, Lincoln. 663 p. Myers, George S. 1925. Notropis cummingsi, a new minnow from Wilmington,

North Carolina. Am. Mus. Novit. 168. 4 pp. Nelson, William R., M. C. Ingham and W. E. Schaaf. 1977. Larval transport

and year-class strength of Atlantic menhaden, Brevoortia tyrannus. Fish.

Bull. 75(1):23-41. Nichols, John T., and C. M. Breder. 1922. Otophidium welshi, a new cusk eel,

with notes on two others from the Gulf of Mexico. Proc. Biol. Soc. Wash.

55:13-15. Nichols, Paul R., and D. E. Louder. 1970. Upstream passage of anadromous

fish through navigation locks and use of the stream for spawning and

nursery habitat Cape Fear River, N. C, 1962-66. U. S. Fish Wildl. Serv.

Circ. 352. 12 pp. Robins, Charles R., R. M. Bailey, E. E. Bond, J.R. Brooker, E. A. Lachner, R.

N. Lea and W. B. Scott. 1980. A list of common and scientific names of

fishes from the United States and Canada. Am. Fish. Soc. Spec. Publ. 12.

4th ed. 174 pp. Schwartz, Frank J. 1979. Caudal peduncle droop in blueback herring, Alosa

aestivalis, from Cape Fear River, North Carolina. ASB Bull. 26(2):34. Abstract. 1981. Effects of freshwater runoff on fishes occupying the freshwater

and estuarine coastal watersheds of North Carolina, pp. 282-294 in R.

Cross and D. Williams (ed.). National Symposium on Freshwater Inflow to

Estuaries. U. S. Fish Wildl. Serv. Off. Biol. Serv. FWS/OBS-81/04. , and A. F. Chestnut. 1973. Hydrographic atlas of North Carolina

estuaries and sound water, 1972. Univ. N. Carol. Sea Grant Publ. SE 73-12. 132 pp. , and M. P. Dahlberg. 1978. Biology and ecology of the Atlantic sting-

ray, Dasyatis sabina (Pisces: Dasyatidae) in North Carolina and Georgia.

Northeast Gulf Sci. 2(1): 1-23.

, P. Perschbacher, M. McAdams, L. Davidson, K. Sandoy, C.

Simpson, J. Duncan and D. Mason. 1979. An ecological study of fishes and invertebrate macrofauna utilizing the Cape Fear River estuary, Carolina

Fishes of Cape Fear Estuary 37

Beach Inlet, and adjacent Atlantic Ocean. Summary Rep. 1973-1977. Inst. Mar. Sci. Univ. N. Carol., Morehead City. 568 p.

and J. Tate. 1979. An ecological study of fishes... Annual

Report for 1978. Inst. Mar. Sci. Univ. N. Carol., Morehead City. 326 pp.

Smith, Hugh M. 1907. The Fishes of North Carolina. N. C. Geol. Econ. Surv., Raleigh. 453 pp.

Sulak, Kenneth J., P. W. Perschbacher and F. J. Schwartz. 1979. Invasion of the Atlantic by Peprilus burti (Pisces: Stromateidae) and possible implica- tions. Copeia 1979(3):538-541.

Tucker, John W., Jr., and R. G. Hodson. 1976. Early and mid-metamorphic larvae of the tarpon, Megalops atlanticus, from the Cape Fear River estu- ary, North Carolina 1973-74. Chesapeake Sci. 77(2): 123-125.

Tyler, James C. 1980. Osteology, phylogeny, and higher classification of the fishes of the order Plectognathi (Tetraodontiformes). NOAA Tech. Rep. NMFS Circ. 434. 422 pp.

Wade, Richard A. 1962. The biology of the tarpon, Megalops atlanticus, and the ox-eye, Megalops cyprinoides, with emphasis on larval development. Bull. Mar. Sci. Gulf Caribb. 72(4):545-622.

Walburg, Charles H., and P. R. Nichols. 1967. Biology and management of the American shad and status of the fishes, Atlantic coast of the United States 1960. U. S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 550:35-38.

Weinstein, Michael P. 1979. Shallow marsh habitats as primary nurseries for fishes and shellfish, Cape Fear River, N. C. Fish. Bull. 77(2):339-358.

, S. L. Weiss and M. F. Walters. 1980. Multiple determinants of

community structure in shallow marsh habitats, Cape Fear River estuary, North Carolina, USA. Mar. Biol. (Bed.) 55:227-243.

, , R. G. Hodson and L. R. Gerry. 1980. Retention of three

taxa of postlarval fishes in an intensively flushed tidal estuary, Cape Fear North Carolina. Fish. Bull. 78(2)'A 19-436. Welch, Joseph M., and B. B. Parker. 1979. Circulation and hydrodynamics of the lower Cape Fear River, North Carolina. NOAA Tech. Rep. NO580. 108 pp.

Accepted 17 December 1981

Nesting of the Green Turtle, Chelonia mydas (L.), in Florida: Historic Review and Present Trends

C. Kenneth Dodd, Jr.

Office of Endangered Species U.S. Fish and Wildlife Service, Washington, D.C. 20240

ABSTRACT.— Except for accounts in the 1800s, which focused on the Keys and Cape Sable region, there are no reliable literature records for nesting of the green turtle, Chelonia mydas, in Florida. Two nests were reported prior to 1959, but the number recorded has steadily increased and at least 366 nests were confirmed in 1980. The majority were found from Merritt Island south to Key Biscayne, and most were laid on relatively undisturbed beaches. Five reasons can be advanced for this apparent increase in nest reports: better surveillance of nesting beaches; greater awareness of sea turtles and their problems; protective legislation; the success of a head-start program on Hutchinson Island; and the possibility that some turtles are immigrating from populations farther south. Although speculative, it is possible that the nesting pop- ulation of green turtles in Florida is increasing and results from a combination of the latter three reasons. A review of the level of histor- ical green turtle nesting, as well as a year-by-year record of nesting since 1959, are provided.

INTRODUCTION

The green turtle, Chelonia mydas (L.), may be the most familiar of sea turtles to mankind. It has been used as a source of food (meat, eggs, calipee) by indigenous peoples throughout the world (see Nietschmann 1972, 1976 for an example of how important the green turtle is to the Miskito Indians of coastal Nicaragua). The turtle also was the basis for gourmet turtle soup in markets formerly in the United States, and such markets still exist in Europe. While subsistence taking for local use probably has not seriously affected wild stocks, with the possible excep- tion of egg taking in certain areas (see Harrisson 1976), the commercial market has seriously depleted the turtles in many regions (Bacon 1975; Nietschmann 1972). In some parts of the world, green turtles are taken incidentally by trawlers (see Chin 1976). Habitat alteration, particularly of nesting beaches and feeding grounds, and predation by both natural and feral predators, have also combined to seriously deplete popula- tions. Pollution of the oceans, especially by petrochemical products, has a known but as yet unquantified effect on this species (Kleerekoper and Bennett 1976; Witham 1978). The green turtle is perhaps historically the most exploited of all marine turtles (Parsons 1962; King, in press), although the hawksbill, Eretmochelys imbricata (L.), may have that dubious distinction today. Chelonia mydas is considered either threa-

Brimleyana No. 7:39-54. July 1 98 1 . 39

40 C. Kenneth Dodd, Jr.

tened or endangered by most sea turtle biologists, depending on the breeding population involved (World Conference on Sea Turtle Conser- vation 1979).

In the Caribbean, the green turtle has a long history of exploitation (see Parsons 1962 for extensive review), chiefly as a source of meat dur- ing colonial exploration, then for sale primarily to European markets. As a result, the remnant nesting populations, centered at Tortuguero, Costa Rica (Carr et al. 1978) and Aves Island, are believed to be but a fraction of a once huge population. Scattered nesting still occurs on many Caribbean and Gulf of Mexico beaches, but everywhere there is pressure on these populations. Indeed, although protected for many years, concern has been expressed about the survival and viability of the Tortuguero colony (Bjorndal 1980), and Aves Island is threatened by erosion. Because of the nesting site fixity of green turtles, it is likely that each nesting area constitutes a unique gene pool (Smith et al. 1978; William R. Rainey, pers. comm.), thus accentuating the importance of the conservation of each nesting population.

In accordance with Section 7 of the U.S. Endangered Species Act of 1973, as amended, the U.S. Fish and Wildlife Service is responsible for the determination of Critical Habitat for listed Endangered and Threatened species. While I will not discuss the concept in detail, such a determination is a conservation tool that requires Federal agencies to insure that activities they authorize, fund, or carry out are not likely to jeopardize the particular habitat on which the species depends (see Dodd 1978 for a complete discussion of the concept as it applies to marine turtles). Since the Florida population of Chelonia mydas is listed as Endangered on the U.S. List of Endangered and Threatened Wildlife and Plants, the determination of its Critical Habitat might provide an additional important conservation measure. This paper, done in prepa- ration of background material for a proposal of Critical Habitat for the species, reports on the level of green turtle nesting in Florida. It is not an endorsement by the U.S. Fish and Wildlife Service of a proposal under the legal requirements of the Act.

HISTORICAL BACKGROUND

Prior to Carr and Ingle's (1959) first report of definite nesting by two turtles, there appears little in the literature to indicate that green turtles ever nested in significant numbers on Florida beaches. Indeed, there is some question whether any green turtles nested on the mainland at all, since references to them are largely confined to fisheries statistics or secondhand accounts. While Carr and Ingle (1959) provided a brief literature review, it would be well to reexamine these references.

The first reference to green turtles in southeastern U.S. waters was

Green Turtle Nesting in Florida 41

that of Catesby (1731-43). He implied that green turtles nested on the Florida coast when he stated that green turtles do not breed in the Bahamas but come from Cuba and the "Continent. " Bahamian turtlers were said to obtain their turtles in two ways and from two sources: by using a detachable spike mounted on a harpoon to spear turtles in local Bahamian waters, and by traveling to Cuba to turn female turtles as they nested. Catesby stated that Bahamians carried their turtles to the Carolinas (which at that time included all territory between Virginia and Florida) since they were esteemed as a "rarity." Catesby also reported that loggerhead sea turtles, Caretta caretta (L.), were not known from the Continent north of Cape Florida, where they are known to nest in abundance today. As the Florida coast was wild and little known during Catesby's time, it is not surprising that precise information regarding species and nesting areas could be confused, especially by one obtaining much information from secondhand sources. However, it is unlikely that Bahamian turtlers would have hazarded a long and arduous voyage to the remote coasts of Cuba had a large exploitable population of nest- ing green turtles been available so close to home. It seems more likely that if green turtles nested along the Florida coast, they nested in such relatively small numbers that turtling expeditions would have been unprofitable. Why loggerheads would go unnoticed north of Cape Flor- ida remains unknown, although it is possible that many of these beaches were never visited by Bahamian turtlers.

The most reliable early accounts of green turtle nesting are the observations of John James Audubon. His account of "The Turtlers" (1926) clearly stated that the green turtle nested not only in the Tortu- gas, but "resorts either to the shores of the main, between Cape Sable and Cape Florida, or enters Indian, Halifax, and other large rivers or inlets, from which it makes its retreat as speedily as possible, and betakes itself to the open sea." The green turtle was said to approach the shores, enter bays, inlets, and rivers early in April, and deposit three clutches of eggs in May and June. Audubon further described a general- ized nesting sequence, a process he reportedly saw several times, which, while primarily based on loggerheads, may have been based in part on observations of the green turtle. Of particular interest was the notation that green turtles selected the wildest and most secluded nesting locations as opposed to loggerheads; similar observations have been noted by L. Ehrhart (1979, pers. comm.) on Merritt Island.

Green turtles, as noted above, do enter lagoonal or river systems, but it is unclear from Audubon's account whether nesting took place on these shores. J. Fletemeyer (1980, pers. comm.) noted, however, that loggerheads rarely nest in such locations and the use of lagoonal areas

42 C. Kenneth Dodd, Jr.

in eastern Florida as green turtle developmental habitat is also well known (Mendonc^ and Ehrhart 1982). Finally, both Ehrhart and R. Witham (pers. comm.) believe that most Florida green turtles nest two or three times per season. Each of these observations supplies credence to Audubon's notes, although it must be pointed out that he often gen- eralized and his species identifications may have been confused. His delimitation of the area of nesting, Cape Sable to Cape Florida (near Miami), may have been more a reflection of the extent of his travels than a known lack of nesting farther north (see Proby 1974 for a review of Audubon's travels in the Key West area). However, green turtles are not definitely recorded today from beaches of either the Tortugas or Cape Sable, although individuals are not uncommonly observed in off- shore waters. Holbrook's (1842) implication that green turtles nested on Florida beaches as well as the Tortugas may in fact have been based in part on Audubon's observations.

True (1884) reported green turtles nesting in Florida, the Bahamas, and the West Indies, and stated that they may occur from Long Island Sound (uncommonly) south through the Gulf States. He said that the turtles were smaller in the north, indicating "young or dwarfed" indi- viduals, and repeated Holbrook's (1842) nesting description verbatim. Nesting was said to occur from April to July. It is clear in True's account, however, that much of his information came from secondhand sources, and he probably had no direct knowledge of green turtle nest- ing in Florida. The originality of this source is thus open to question.

As Carr and Ingle (1959) noted, William T. Sherman (1889) recorded the importance of green turtles to the economy of eastern Flor- ida, where he was stationed in 1840. Sherman stated: "They were so cheap and common that the soldiers regarded it as an imposition when compelled to eat green turtle steaks instead of poor Florida beef or the usual mess-pork. I do not recall in my whole experience a spot on earth where fish, oysters, and green turtles so abound as at Fort Pierce, Flor- ida." He did not record, however, how the turtles were obtained or their sizes, which would have given an indication of whether nesting might have occurred in the area.

Between 1840 and the mid- 1890s green turtles had clearly suffered a significant decline in the Indian River area of eastern Florida. Turtle fisheries were operated at Sebastian, Ft. Pierce, and Eden, although tur- tles were incidentally taken elsewhere. After reviewing available catch statistics, Wilcox (1896) stated: "There is no doubt that turtle fishing on the Indian River is much less productive than formerly. Mr. Charles Pearke, of Sebastian, who has followed the turtle business during the past ten years, reports a great decrease of turtles as compared with ear-

Green Turtle Nesting in Florida 43

lier years. About 1886 he took 2,500 turtles with eight nets; in 1895 he secured only 60 turtles with six nets. The principal reason assigned for the decrease by Mr. Pearke is that the turtles have been frightened off by the steamboats and launches. The unusual cold of the winter of 1894- 95 is also known to have seriously affected the abundance of turtles. Several hundred turtles were then found floating on the surface in a numbed or frozen condition. On being warmed most of them survived and were soon on their way to the northern markets. Since the cold spell turtles have been much scarcer than ever." This quote has often been incorrectly attributed to Evermann and Bean (1896).

Wilcox also noted that the fishery was a net fishery operating between November and March, and that although occasional turtles weighed 220 pounds, the average weight was 50 pounds in 1891 and 36 pounds in 1895. This would indicate that a juvenile rather than a nesting population was being exploited.

Brice (1896) also noted a decline in green turtle abundance, this time in the Keys region, and said it was probably due to overfishing. He stated: "The fishermen comment upon the fact that for the past few years the green turtles have not been depositing their eggs on Key West and the adjacent keys. It is very probable that this is owing to the exces- sive hunting of this species, and that they now deposit their eggs on the more distant and inaccessible keys." It is clear that Brice was talking about a breeding population, since nesting was mentioned and the tur- tles ranged in size from 125 to 275 pounds, unlike those of the fisheries on Cedar Key and the Indian River. The turtle grounds were principally from Marquesas Key to Alligator Light on the east coast, the Bay of Florida, and the Gulf of Mexico, intervening between the western keys and the mainland. This is roughly the same area with which Audubon (1926) was most familiar. By the time of later turtle statistics (Ingle and Smith 1949; Rebel 1974), Key West and other Florida ports served as outlets for a turtle fishery throughout the Caribbean instead of one solely regionally based; the local green turtle breeding population had been decimated. Until the report of Carr and Ingle (1959), green turtles had not been reported as nesting in Florida since 1896.

NESTING TRENDS SINCE 1959 The numbers of green turtle nests that can be documented since 1959 are provided by site in Table 1. The sources of data for this table are: Carr and Ingle (1959); Routa (1967); Gallagher et al. (1972); Ehrhart (1975, 1979); Worth and Smith (1976); Mann (1977); Ehrhart and Yoder (1978); Fletemeyer (1979, 1980); Stoneburner et al. (1979); Florida Department of Natural Resources (1979, 1980); Schwartz et al.

44 C. Kenneth Dodd, Jr.

				NC. \
				j^i
			sc.
		GA. •
		P2	ATLANTIC OCEAN
	N t		\?
160 km
	1		Va
		FL.	B5
			r6 ■ 7 08	Cape Florida
Cape Sable	10 ?©	£e7s	/ 9

Fig. 1. Location of green turtle nesting sites in the southeastern United States. 1. Camp Lejeune 2. Jekyll Island 3. Merritt Island 4. Sebastian Inlet 5. Hutchinson Island 6. Hobe Sound National Wildlife Refuge 7. Highland Beach 8. Hillsboro Beach 9. John U. Lloyd State Park 10. Loggerhead Key.

(1981); Litwin (1981); and personal communications from C. and M. Barsumian, G. Dalrymple, L. M. Ehrhart, J. Fletemeyer, T. W. Fritts, T. W. Martin, M. Murphy, J. Richardson, D. L. Stoneburner, and R. Witham. The locations of the major nesting areas, originally selected as possible areas of Critical Habitat, as well as two outlier nestings, are shown in Figure 1.

Caution should be exercised when using the figures in Table 1. They indicate a number only, and not necessarily a year-to-year trend, with the exception perhaps of Hutchinson Island, Merritt Island, and Hobe Sound. This is because not every beach received the same inten- sity of survey from one year to the next, and because the same precise area may not have been covered. For instance, Sebastian Inlet North received intensive aerial and ground survey in 1980. However, surveys were not performed in 1981. A single night survey yielding four nests does not necessarily indicate a precipitous decline of nesting on this beach in 1981. Considering these limitations, it is evident that the

Green Turtle Nesting in Florida

45

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46 C. Kenneth Dodd, Jr.

number of green turtle nests reported in Florida is increasing, and that a rudimentary cyclical pattern may be emerging.

Of particular interest is the grouping of nestings from Merritt Island south to Key Biscayne. Early accounts, principally those of Audubon (1926) and Brice (1896), do not record nesting from these beaches, and there are no reliable modern reports of nesting in the Keys and the Cape Sable area. Since green turtle nesting is now confined to the southeast Atlantic coast, this area should receive major concern for sea turtle protection and education projects.

The outlier nestings recorded in Georgia and North Carolina in 1980 are noteworthy in that they constitute the first reliable reports of nestings outside Florida. While green turtles have long been known to occasionally occur in coastal waters as far north as New England, only loggerheads nest occasionally as far north as New Jersey. In the North Carolina report (Schwartz et al. 1981), one turtle is known to have made all the nests; in the Georgia instance, only one nest was reported (Litwin 1981) and it is unknown if this female nested elsewhere in the vicinity. Since there are no beach patrols from Merritt Island north to the Georgia barrier islands, perhaps she deposited additional nests in this area.

In 1979, personnel of the National Park Service (NPS) visited Log- gerhead Key in the Tortugas (Fig. 1). During their stay, one nest of what was believed to be a green turtle was located high up in the beach grass (R. Dawson, pers. comm.). Since the green turtle digs a character- istic body pit quite distinctive from that of the loggerhead, experienced personnel can distinguish the two types of nests. However, no way of verifying this record now exists, so nesting in the Tortugas should at best be considered only a possibility. No green turtle nests could be determined elsewhere during NPS surveys, although many juvenile green turtles were seen in waters off the Tortugas.

HUMAN IMPACT ON FLORIDA NESTING BEACHES

To illustrate potential problems facing green turtle nesting in Flor- ida, human use of selected important beaches is reviewed below.

Merritt Island

Ownership: U.S. Government

Use: Recreational (day only); restricted access to military and National Aeronautics and Space Administration lands; no commercial activities or development.

This area is entirely under the jurisdiction of various Federal agen- cies. The island is largely under the administrative auspices of Merritt Island National Wildlife Refuge, which is administratively superim- posed on the lands of Kennedy Space Center. Public access to the Cen-

Green Turtle Nesting in Florida 47

ter is restricted at all times. The other beaches at Merritt Island are under the jurisdiction of the National Park Service (Canaveral National Seashore). Access to the most popular beach, Playalinda, and all south- ern beaches within the seashore, is restricted to daylight hours. The farthest south section of beach is under the jurisdiction of Cape Canav- eral Air Force Station, and public access is restricted.

Hutchinson Island

Ownership: Private, 15 km; St. Lucie and Martin counties, < 1.6 km

Use: Recreation.

Hutchinson Island is a 36 km barrier island in an area that is expe- riencing rapid development, primarily in the south. It has a dune system which rises to 5 m in some areas. Beaches are rather narrow and sup- ported by sand streaming toward the south. The island is experiencing erosion, which is most severe in the north and gradually tapers off to the south. As of 1979, 37.8 percent (13.6 km) of beachfront had been developed; development is interspersed with undeveloped land in private ownership and county recreation beaches. Sea turtle nesting is lowest in the north in areas of worst erosion and, in the south, most nesting occurs on undeveloped beaches (Gallagher et al., in press).

Hobe Sound

Ownership: U.S. Government, 5.15 km; City (Jupiter Island), 0.16 km; Private, 0.32 km

Use: Recreation, wildlife protection; limited access, day only.

Nearly all beaches in this area are located on Hobe Sound National Wildlife Refuge, where public use after dark is not allowed without spe- cial permission. The principal threats to green turtle nests are raccoon predation and beach erosion, primarily at the northern end of the refuge.

John U. Lloyd State Park

Ownership: State, 4.0 km

Use: Recreation, day only.

Much of the beach here has been renourished by the Corps of Engineers; the sand is derived from 80' - 90' of water from offshore areas. The last such effort occurred about five or six years ago, and extended 1.93 km south of the inlet. John Fletemeyer, Nova University, believes that about 10 years are required for a renourished beach to return to normal characteristics of compaction. Under normal regimes, renourishment is accomplished every 10 years or so. Silt is the big com- pacting problem, not only forming a physical barrier but probably also inhibiting gas exchange and affecting incubation temperatures. As part

48 C. Kenneth Dodd, Jr.

of the renourishment project, the Corps of Engineers has been required by the Florida Department of Natural Resources to relocate nests. For this they have contracted with Fletemeyer to patrol the beach every morning. Since this beach has no night use, nest relocation probably involves no serious detriment to the turtles.

Highland Beach

Ownership: Private, 4.5 km

Use: Recreation, restricted access.

The only potential for public access to this beach is at a motel on the northern section. Like the other areas in private ownership, the resi- dents here are protective of the turtles and have taken measures to insure their continued survival. The measures include funding and par- ticipating in sea turtle beach patrols.

Perhaps the main feature that accounts for green turtle nesting on the beaches indicated in Table 1 is restricted night use. Merritt Island, Hobe Sound, John U. Lloyd State Park, and Key Biscayne allow no night activities; Highland Beach and Hillsboro Beach have restricted public access and owners who are aware of the turtles and the impor- tance of protecting them from disturbance; Sebastian Inlet North, Juno Beach, N. Melbourne Beach, and Ft. Pierce-Sebastian are all presently little disturbed at night. Only at Hutchinson Island is development rather extensive, and here turtles often select what undeveloped beaches remain between high-rise buildings. Many buildings on Hutchinson Island lack bright outdoor lighting facing the ocean, lighting that may inhibit nesting females (R. Witham, pers. comm.). While disturbance may be a primary inhibiting factor to nesting, development of areas behind dunes may lead to increased night use of beaches, regardless of lighting or other building restrictions imposed as mitigation measures. Relatively undisturbed nesting beaches, especially those of Sebastian Inlet North, should remain undeveloped to insure the protection of green turtles. Where development has occurred, planting of trees behind beaches to obscure buildings, such as has been done at John U. Lloyd State Park, may aid in reducing factors disturbing to nesting turtles. Other mitigating measures are provided by Shabica (in press).

The survival of Florida's nesting colony also depends on the survi- val of hatchlings from Florida nests. Since the hatchlings become diso- riented by bright night lights, development of nesting beaches becomes a severe problem, leading to mortality from desiccation and increased exposure to predators (Mann 1977). On developed beaches where green turtles (and loggerheads) are known to nest, it may be necessary to patrol just before sunrise to look for misdirected hatchlings. While this may prove difficult, the alternative is to ignore a potentially serious loss

Green Turtle Nesting in Florida 49

to the population. In the long term, it may be impossible to sustain both viable turtle populations and extensive development, regardless of res- trictive legislation against take or harm. To adequately protect a popu- lation, all habitats used during each life stage must also be protected.

IS THE POPULATION INCREASING?

Despite the spurt of development along much of the southeast Florida coast, there can be little doubt that there are now more green turtles reported nesting than there were 20 years ago. A number of explanations are possible.

Better surveillance. — The first reports of green turtle nesting in Florida (Carr and Ingle 1959; Routa 1967; Gallagher et al. 1971) were based on the observations of a few individuals, made either by chance or while working a limited stretch of beach. Since 1959, there has been a substantial increase in the number of sea turtle beach patrols and the extent of beach covered. In 1979, the Florida Department of Natural Resources listed 20 areas regularly surveyed, and 29 in 1980. As an example, intensive aerial surveys revealed significant nesting in the Seb- astian Inlet North area where only incidental observations had been made in the past. Better surveillance undoubtedly accounts for some of the increase in reported nests, and additional patrols will likely increase future estimates of the level of Florida nesting.

Greater public awareness. — During the last several years, there has been increased publicity concerning sea turtles and the threats facing them. Articles have appeared in numerous newspapers and magazines, and some local governments have even begun to fund sea turtle beach patrols and adopt regulations designed to assist conservation efforts. R. Witham (pers. comm.) believes that greater public awareness may help to decrease poaching, since citizens alert law enforcement agencies of suspected illegal activity. The appearance of a "strange" turtle (as opposed to the numerous loggerheads) is likely to result in reports to appropriate authorities.

Protective legislation. — The State of Florida has had laws protect- ing green turtles, and their eggs and nests, since 1953, although these laws applied only from May through August. In 1974, the Florida Legislature extended the laws to include year-round protection for sea turtles (Shelfer 1978). These laws are strictly enforced by the Florida Marine Patrol of the Department of Natural Resources. In 1978, the green turtle breeding population in Florida was listed as Endangered under provisions of the Federal Endangered Species Act of 1973, as amended (16 U.S. Code 1531-43). The turtles and all parts and pro- ducts, including eggs and shells, are fully protected. The U.S. Fish and Wildlife Service and National Marine Fisheries Service, working in con-

50 C. Kenneth Dodd, Jr.

junction with State and local agencies, have responsibility for en- forcement.

As a result of these laws, most nesting and coastal green turtles are now unmolested, except for incidental take in shrimp trawls and boat related injuries. Vandalism still occurs, however, and only the continued strict enforcement of protective laws, with the imposition of severe penalties, will deter such activity.

Head-start program on Hutchinson Island. — Since 1960, Ross Witham, Florida Department of Natural Resources, has been raising green turtles and releasing them off Hutchinson Island, Florida, in one of the earliest head-start programs in the world. This activity has been supported since 1971 by the Marine Research Laboratory of the Florida DNR. Turtles are raised to a size where natural predation should be reduced (approximately 20 cm), after which they are tagged and released (see Witham and Futch 1977 for details). Head-starting is an experimen- tal technique, and not all sea turtle biologists are enthusiastic about its use as a management tool (see Pritchard 1979 for a discussion). How- ever, turtles released in this program are known to survive in the wild, and are found in the same developmental habitat off Florida's coast where similar-size "wild" turtles occur (Witham and Futch 1977; Witham, pers. comm.). While there is no proof that increased nesting along Florida's east central coast is due to this head-start program, it must be recognized as a possibility, at least in part.

Immigration. — One final possibility is that turtles now nesting in Florida are immigrants from other green turtle populations. Perhaps such individuals represent an expanding population, or they may be less nest site specific than green turtles are supposed to be. Immigrants could follow the western edge of the Gulf Stream during favorable water conditions and nest on beaches adjacent to known developmental habitat. However, Caribbean green turtle stocks are depleted and are not likely to be subject to such population pressure as to force the sur- plus into Florida waters in exponentially increasing numbers. The idea that the Florida population is the result of offspring from a few random founder individuals cannot be easily dismissed. If such were the case, however, it is difficult to understand why nesting is not increasing in the Keys. If most female green turtles return to their natal beach to deposit their eggs, a few successful hatchings on normally unfrequented beaches might lead to range extension, although this would be difficult to determine.

CONCLUSIONS

While earlier authors accepted the idea that Florida beaches once supported a large nesting population of green turtles (Carr and Ingle

Green Turtle Nesting in Florida 51

1959:317), little hard evidence supports this position. What is known is that green turtles have long used coastal waters and lagoons as devel- opmental habitat, and that such populations were appreciably larger than at present. Overharvest, perhaps in conjunction with occasional winter kills (Wilcox 1896; Ehrhart 1977; Ehrhart and Lee 1981), deci- mated these largely juvenile and subadult populations. Overharvest is also implicated in the demise of what may have been a substantial nest- ing population in the Keys and Cape Sable region. There can be little doubt, however, that green turtles are increasingly encountered on east Florida beaches. Due to variances in the amount of time spent in survey and the length of beach covered, it is difficult to estimate a population size; surely many individuals have been unnoticed. Therefore, the fig- ures in Table 1 are minimum numbers and cannot be used compara- tively (except very generally) or to indicate concrete trends. It should be noted that there do appear to be "good" years and "bad" years, a cycle that has been observed in Australian green turtle populations (Limpus 1980) and is very poorly understood. Only systematic, well coordinated, long term research, involving beach patrols, tagging, and correlation with environmental variables, can ultimately shed light on the status of the green turtle in Florida.

The question remains as to whether the Florida nesting population is actually increasing. Better surveillance and greater publicity undoubt- edly account for some of our awareness of green turtle nesting. It is possible that this trend is not as new as Table 1 would make it appear. However, I believe the facts argue that the population is increasing, although I am skeptical that Florida ever had "great nesting assemb- lages" (Carr and Ingle 1959). Instead, I hypothesize that there at one time was a small but stable nesting population, perhaps only a few thousand breeding females, that was decimated by overharvest. Such a population could easily have been exploited, and the dearth of natural- ists in this area could account for the lack of definite records prior to more modern times. Green turtles still frequented offshore waters, but even there they were reduced. By the late 1890s, all nesting may have ceased. However, beginning in the mid-twentieth century, laws were enacted to at least partly protect turtles and their nests. With poaching pressure reduced, early head-starting could have increased the popula- tion somewhat and made it possible for some individuals to exploit a still largely undeveloped coastline. At the same time, immigrant turtles may have provided some new genetic input to the small population. Scientific research, and public awareness of turtles that perhaps resulted from legislation, in turn revealed more turtles nesting. No single explan- tion is deemed adequate, and the long term future of Chelonia mydas in Florida is yet to be determined.

52 C. Kenneth Dodd, Jr.

In the scenerio presented above, many of the nesting beaches avail- able to the green turtle were undeveloped. Turtles were occasionally poached, run over by boats, and faced problems associated with natural beach erosion and predation. But today, regardless of whether the popu- lation is expanding, development of remaining habitat and the increas- ing human presence it entails threatens to permanently end green turtle nesting in Florida. Unless beaches can be preserved, there is no secure future for this remarkable species.

ACKNOWLEDGMENTS.— I would like to thank the following individuals for the generous sharing of data and , in many cases, ideas:

C. and M. Barsumian, A. Carr, G. Dalrymple, R. Dawson, L. Ehrhart, J. Fletemeyer, T. W. Fritts, T. W. Martin, M. Murphy, J. Richardson,

D. L. Stoneburner, and R. Witham. I thank Archie Carr, Thomas Fritts, P. C. H. Pritchard, and Ross Witham for their comments on the manuscript, and Sheila Hulsey for her silent encouragement.

LITERATURE CITED Audubon, John J. 1926. The turtlers. pp. 194-202 in Delineations of American

Scenery and Character. G. A. Baker and Co., New York. 349 pp. Bacon, Peter R. 1975. Review on research, exploitation, and management of the

stocks of sea turtles. FAO Fish. Circ. 334. 19 pp. Bjorndal, Karen A. 1980. Demography of the breeding population of the green

turtle, Chelonia mydas, at Tortuguero, Costa Rica. Copeia 1980(3):525-530. Brice, John J. 1896. The fish and fisheries of the coastal waters of Florida. Rep.

U. S. Comm. Fish Fish. 22:263-342. Carr, Archie F., and R. M. Ingle. 1959. The green turtle (Chelonia mydas) in

Florida. Bull. Mar. Sci. Gulf Caribb. 9:315-320. ,M.H. Carr and A. B. Meylan. 1978. The ecology and migrations of

sea turtles, 7. The west Caribbean green turtle colony. Bull. Am. Mus. Nat.

Hist. 762:1-46. Catesby, Mark. 1731-1743. The Natural History of Carolina, Florida, and the

Bahama Islands. 2 Vols. Author, London. Chin, Lucas. 1976. Notes on marine turtles (Chelonia mydas). Sarawak Mus. J.

25(44):259-265. Dodd, C. Kenneth, Jr. 1978. Terrestrial Critical Habitat and marine turtles.

Bull. Md. Herpetol. Soc. 74:233-240. Ehrhart, Llewellyn M. 1977. Cold water stunning of marine turtles in Florida east

coast lagoons: rescue measures, population characteristics and evidence of

winter dormancy. Am. Soc. Ichthyol. Herpetol. mtg., Gainesville, Fl.

Abstract. 1979. A survey of marine turtle nesting at the Kennedy Space Cen- ter, Cape Canaveral Air Force Station, North Brevard County, Florida.

Rep. to Div. Mar. Resour., Fla. Dep. Nat. Resour. 122 pp. , and R.C. Lee. 1981. Hypothermic stunning of marine turtles in east-

central Florida lagoons in 1981. Soc. Study Amphib. Reptiles and Her- petol. League mtg., Memphis, TN. Abstract.

Green Turtle Nesting in Florida 53

, and R.G. Yoder. 1978. Marine turtles of Merritt Island National

Wildlife Refuge, Kennedy Space Center, Florida. Fla. Mar. Res. Publ. 33:25-30.

Evermann, B.W., and B.A. Bean. 1896. Indian River and its fisheries. Rep. U.S. Comm. Fish Fish. 22:227-248.

Fletemeyer, John. 1979. Sea turtle monitoring project. 1979 Report. Rep. to Broward Co. Environ. Qual. Control Bd., Ft. Lauderdale. 64 pp.

1980. Sea turtle monitoring project. 1980 Report. Rep. to Broward

Co. Environ. Qual. Control Bd., Ft. Lauderdale. 88 pp.

Florida Department of Natural Resources. 1979. Summary of sea turtle activity in Florida, 1979. 20 pp.

1980. Summary of marine turtle activity in Florida, 1980. 39 pp.

Gallagher, Robert M., M.L. Holling, R.M. Ingle and C.R. Futch. 1972. Marine turtle nesting on Hutchinson Island, Florida, in 1971. Fla. Dep. Nat. Re- sour. Res. Lab. Spec. Sci. Rep. No. 37. 1 1 pp.

, J. O'Hara and D.F. Worth. In press. Characteristics of loggerhead

turtles nesting on Hutchinson Island, Florida. Bull. Mar. Sci. 33(1).

Harrisson, D.S.L.J.T. 1976. Green turtles in Borneo. Brunei Mus. J. 5:196-198.

Holbrook, John E. 1842. North American herpetology; or a description of the reptiles inhabiting the United States. 2nd ed., Vol. 1. J. Dobson, Philadel- phia, xv + 152 pp.

Ingle, Robert M., and F.G.W. Smith. 1949. Sea turtles and the turtle industry. Spec. Publ. Univ. Miami, Coral Gables. 107 pp.

King, F. Wayne. In press. Historical review of the status of the green turtle, Chelonia my das, and hawksbill, Eretmochelys imbricata. In K.A. Bjorndal (ed.). Proc. World Conf. Sea Turtle Conserv., Smithson. Inst. Press, Washington.

Kleerekoper, H., and J. Bennett. 1976. Some effects of the water soluble pollu- tion fraction of Louisiana crude on the locomotor behavior of juvenile green turtle {Chelonia my das) and sea catfish (Arius felis). Preliminary results. Rep. Contract 206-76 Am. Petrol. Inst. 7 pp.

Limpus, Colin J. 1980. The green turtle, Chelonia mydas (L) in eastern Aus- tralia, pp. 5-22 in Management of Turtle Resources. James Cook Univ. North Queensland Res. Monogr. No. 1. 72 pp.

Litwin, Salome C. 1981. Chelonia mydas mydas (Green turtle). Nesting. Her- petol. Rev. 72(3):81.

Mann, Thomas M. 1977. Impact of developed coastline on nesting and hatch- ling sea turtles in southeastern Florida. Unpubl. M.S. thesis, Fla. Atlantic Univ., Boca Raton. 100 pp.

Mendonc,a, Mary T., and L.M. Ehrhart. 1982. Activity, population size and structure of immature Chelonia mydas and Caretta caretta in Mosquito Lagoon, Florida. Copeia 1982(1): 161-167.

Nietschmann, Bernard. 1972. Hunting and fishing focus among the Miskito Indians, eastern Nicaragua. Hum. Ecol. 7:41-67.

1976. Memorias de arrecife tortuga. Ser. Geograf. Natur. No. 2,

Banco de America, Managua, Nicaragua. 258 pp.

54 C. Kenneth Dodd, Jr.

Parson, James J. 1962. The green turtle and man. Univ. Fla. Press, Gainesville.

126 pp. Pritchard, Peter C.H. 1979. 'Head-starting' and other conservation techniques

for marine turtles. Cheloniidae and Dermochelyidae. Int. Zoo Yearb.

79:38-42. Proby, K.H. 1974. Audubon in Florida. Univ. Miami Press, Coral Gables. 384

pp. Rebel, Thomas P. 1974. Sea turtles and the turtle industry of the West Indies,

Florida, and the Gulf of Mexico. Rev. ed. Univ. Miami Press, Coral

Gables. 250 pp. Routa, Robert A. 1967. Sea turtle nest survey of Hutchinson Island, Florida. J.

Fla. Acad. Sci. 50:287-294. Schwartz, Frank J., C. Peterson, H. Passingham, J. Fridell and J. Wooten.

1981. First successful nesting of the green turtle, Chelonia my das, in North

Carolina and north of Georgia. ASB Bull. 2#(2):96. Abstract. Shabica, Stephan V. In press. Planning for protection of sea turtle habitat. In

K. Bjorndal (ed.). Proc. World Conf. Sea Turtle Conserv. Smithson. Inst.

Press, Washington. Shelfer, L. 1978. Florida's enforcement of marine turtle conservation laws. Fla.

Mar. Res. Publ. 55:65. Sherman, William T. 1889. Memoirs. Vol. I. D. Appleton and Co., New York. Smith, Michael H., H.O. Hillestad, M.N. Manlove, D.O. Straney and J.M.

Dean. 1978. Management implications of genetic variability in loggerhead

and green sea turtles. XHIth Congr. Game Biol.:302-312. Stoneburner, D.L., D. Gilmore, J. Hinesley, D. Gross, D. Hall and J.L.

Richardson. 1979. Observations on Chelonia my das: a northerly extension

of known nesting range. Herpetol. Rev. 10(3): 103-104. True, Frederick W. 1884. The useful aquatic reptiles and batrachians of the

United States. Part II, Sect. I: The Fisheries and Fishery Industries of the

United States. U.S. Comm. Fish Fish., Washington, pp. 141-162. Wilcox, William A. 1896. Commercial fisheries of Indian River, Florida. Rep.

U.S. Comm. Fish Fish. 22:249-262. Witham, Ross. 1978. Does a problem exist relative to small sea turtles and oil

spills? Proceedings Conference on Assessment of Ecological Impacts of Oil

Spills. AIBS. pp. 630-632. , and C.R. Futch. 1977. Early growth and oceanic survival of pen- reared sea turtles. Herpetologica 55:404-409. World Conference on Sea Turtle Conservation. 1979. Sea Turtle Conservation

Strategy. IUCN, Gland, Switzerland. 38 pp. Worth, Dewey F., and J.B. Smith. 1976. Marine turtle nesting on Hutchinson

Island, Florida, in 1973. Fla. Dep. Nat. Resour. Mar. Res. Publ. 18:1-17.

Accepted 29 March 1982

Thermal Preferenda and Diel Activity Patterns

of Fishes from Lake Waccamaw

W. W. Reynolds and M. E. Casterlin

Biothermal Research Institute R. D. 3, Box 10, Wyoming, Pennsylvania 18644

and

D. G. LlNDQUIST '

Department of Biology University of North Carolina, Wilmington, North Carolina 28406

ABSTRACT.— We tested seven fish species from Lake Waccamaw in Ichthyotron electronic shuttleboxes to determine their preferred temper- atures and diel activity patterns. All preferred temperatures between 25 ° and 31 ° C, which corresponded to water temperatures measured in the lake in August 198 1 . All species avoided temperatures below 20 ° or above 36° C. One species was diurnal, two were nocturnal, and four were crepuscular in diel activity pattern under the LD 14:10 August photoperiod.

INTRODUCTION

Lake Waccamaw, North Carolina, is one of the shallow "bay lakes" of the southeastern coastal plain of North America. It is unusual among these lakes in having significant calcareous buffering capacity, and thus a pH normally at or above neutrality. Another unusual feature of Lake Waccamaw is that it harbors three endemic fish species: Fundulus wac- camensis Hubbs & Raney, the Waccamaw killifish (Cyprinodontidae); Menidia extensa Hubbs & Raney, the Waccamaw silverside (Atherini- dae); and Ethe o stoma per longum (Hubbs & Raney), the Waccamaw dar- ter (Percidae, Etheostomatinae). An undescribed ictalurid, Noturus sp., the broadtail madtom, also occurs in the lake.

In connection with a survey to assess the conservation status of the endemic species, we measured the thermoregulatory behavior (preferred and avoided temperatures) and diel activity patterns of several fish spe- cies. They included the three endemic and one undescribed species named above, and three others that had never been so tested — Lepomis margina- tus (Holbrook), the dollar sunfish (Centrarchidae); Enneacanthus chae- todon (Baird), the blackbanded sunfish (Centrarchidae); and Noturus gyrinus (Mitchell), the tadpole madtom (Ictaluridae).

1 Direct reprint requests to DGL

Brimleyana No. 7:55-60. July 198 1 . 55

56 W. W. Reynolds, M. E. Casterlin, D. G. Lindquist

MATERIALS AND METHODS

The fishes (38-60 mm standard length) were captured by seine or hand net, either in the lake or in backwaters of the Big Creek tributary, and tested individually in Ichthyotron electronic shuttleboxes as described by Reynolds (1977). These devices permit a fish, by passing through an electronic gate (light beam), to control water temperatures by normal swimming movements, which are monitored automatically via photo- transistors. The fish were tested during the first half of August 1981, under a natural photoperiod of 14 h light: 10 h dark (LD 14:10). Final thermal preferenda, which are independent of thermal acclimation (Rey- nolds & Casterlin 1979, 1980), were measured, and hourly activity as registered by the phototransistors (mean light beam interruptions per hour) were used to characterize diel activity patterns (Table 1). In gen- eral, more precise behavioral thermoregulators will show greater activity. The relative amount of activity (light beam interruptions) during day, night, and crepuscular periods is used to determine the diel patterns.

RESULTS

All species exhibited mean final thermal preferenda between 25 ° and 30° C (Table 1). Various measures of central tendency failed to coincide for each species because of skewness in the distributions of occupied temperatures. Of these species, Lepomis marginatus was the most precise behavioral thermoregulator, exhibiting the smallest range and standard deviation, while Menidia extensa was the least precise. Thus, the former species also exhibited the highest hourly activity rate, while the latter was the least active. Noturus gyrinus preferred the warmest temperatures, and Enne acanthus chaetodon the coolest, in terms of mean value.

Fundulus waccamensis was the only diurnal species as determined by maximal activity, while Etheostoma perlongum and Noturus sp. were nocturnally most active (Table 1). The other species were crepuscular, being most active around dawn and/ or dusk. Here there was a marked behavioral difference between the two Noturus species, which often share the same microhabitats and nesting sites (Lindquist, unpubl.).

During the first half of August 198 1 , normally a time of warmest water, the temperatures in the lake ranged from 27.4 ° to 3 1 .0 ° C over all areas and depths. Warmest temperatures occurred near the mouth of Big Creek, the major tributary. In the creek and associated canals, tempera- tures ranged from 24.8 ° to 30.9 ° C during the same period.

DISCUSSION

Lake Waccamaw does not normally freeze completely over in winter, nor does it stratify thermally to any significant extent in summer, as its maximum depth is only about 3 m. The temperatures preferred by the

Behavior of Lake Waccamaw Fishes

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58 W. W. Reynolds, M. E. Casterlin, D. G. Lindquist

fishes tested correspond fairly well to normal maximum summer temper- atures in the lake. This is a typical result, in that freshwater and marine fishes often prefer temperatures (final preferenda) that are normal sea- sonal maxima in their habitats and geographic ranges (Reynolds & Thomson 1974; Reynolds et al. 1977; Casterlin & Reynolds 1979a, 1982).

Field observations of fish distributions and movements appeared lar- gely consistent with our laboratory measurements. For example, as inshore lake temperatures over a two-week period warmed from 27.4 ° to 29.4 °C at one location, fewer Fundulus and Menidia could be seen or captured by seine inshore during the day, although they moved inshore at night as some cooling occurred. Enne acanthus chaetodon was captured in the cooler depths of backwater canals, which had lower temperatures than occurred in the lake at that time. We did not see this species in the lake, where temperatures were above its mean final preferendum.

A more dramatic field example of apparent behavioral thermoregula- tion occurred during the unusually dry and hot summer of 1980. Diurnal temperatures in the lake shallows reached 40 ° C, and all fish then vacated these lethally hot regions, at least during daylight hours (Lindquist and Yarbrough, unpubl.).

We often found males of both Noturus species guarding nests under the same spawning cover, and their thermal preferenda (Table 1) appar- ently differ little. The striking difference in their diel activity patterns (one being nocturnal, the other crepuscular and mainly active at dawn) sug- gests a temporal niche partitioning between these otherwise closely sim- ilar species.

The very high activity level of Lepomis marginatus (Table 1) is note- worthy. It apparently reflects territorial or reproductive aggression, which was occurring at that time, and may not be typical of other sea- sons. For example, intense agonistic interactions, with resultant body damage, was seen among individuals in holding tanks, although this could not occur during the testing of single individuals in the Ichthyo- tron. It is also interesting that L. marginatus preferred lower tempera- tures than Lepomis macrochirus Rafinesque, the bluegill sunfish, which prefers about 31 °C (Reynolds & Casterlin 1979) but ranges much farther north. Enneacanthus chaetodon prefers temperatures similar to those preferred by Enneacanthus gloriosus (Holbrook), the bluespotted sunfish (Reynolds & Casterlin 1980; Casterlin & Reynolds 1979b).

Etheostoma perlongum is only the third darter species to have been successfully tested for thermoregulatory behavior in the laboratory. Hill & Matthews (1980) reported thermal preference data for Etheostoma spectabile (Agassiz) and Etheostoma radiosum (Hubbs & Black) from the Blue River in Oklahoma, and found a correlation between thermoregula- tory precision and the thermal stability of microhabitats occupied by

Behavior of Lake Waccamaw Fishes 59

these species in nature. In our tests, E. perlongum exhibited a thermoreg- ulatory precision that was intermediate among the species tested. Lind- quist et al. (1981) observed seasonal onshore-offshore spawning migra- tions of E. perlongum, which might be cued by changes in water temperature. Although darters generally appear to be diurnal (see Adam- son & Wissing 1977; Cordes & Page 1980; Mathur 1973), our data sug- gest that E. perlongum is nocturnal. However, our results may not accu- rately represent the fish's activity pattern in nature, and further tests are necessary to resolve this apparent discrepancy.

Enneacanthus gloriosus exhibits a crepuscular activity pattern (Caster- lin & Reynolds 1980), as does E. chaetodon (Table 1). Centrarchids generally tend to be diurnal or crepuscular (Reynolds & Casterlin 1976; L. marginatus in Table 1). In contrast, ictalurids such as Noturus (Table 1) tend to be nocturnal or crepuscular, as are the Ictalurus species (Rey- nolds & Casterlin 1977, 1978). Ictalurids and centrarchids are considered typical "warm-water" fish species (Reynolds 1979), in contrast to "cold- water" species such as salmonids that prefer temperatures below 20 °C. Percids are more variable, with various species preferring widely different temperatures (Reynolds 1979).

ACKNOWLEDGMENTS.— We thank John McNeill for generously allowing us the use of his cabin at Lake Waccamaw, and John R. and Peggy W. Shute for help in capturing several of the species. This study was made possible by grant-in-aid funds provided under section 6 of the Endangered Species Act of 1973 (PL 93-205) to David G. Lindquist. These funds were administered by the North Carolina Wildlife Resources Commission.

LITERATURE CITED Adamson, Scott W., and T. E. Wissing. 1977. Food habits and feeding periodicity

of the rainbow, fantail, and banded darters in Four Mile Creek. Ohio J. Sci.

77:164-169. Casterlin, Martha E., and W. W. Reynolds. 1979a. Shark thermoregulation.

Comp. Biochem. Physiol. 64 A AS 1-453. ,and 1979b. Thermoregulatory behavior of the bluespotted

sunfish, Enneacanthus gloriosus. Hydrobiologia 64:3-4. , and 1980. Diel activity of the bluespotted sunfish, Ennea-

canthus gloriosus. Copeia 1980 (2):344-345. , and 1982. Thermoregulatory behavior and diel activity of

yearling winter flounder, Pseudopleuronectes americanus (Walbaum). Environ. Biol. Fishes 7: 177- 180. Cordes, Lynn E., and L. M. Page. 1980. Feeding chronology and diet composi- tion of two darters (Percidae) in the Iroquois River System, Illinois. Am. Midi. Nat. 104:202-206.

60 W. W. Reynolds, M. E. Casterlin, D. G. Lindquist

Hill, Loren G., and W. J. Matthews. 1980. Temperature selection by the darters

Etheostoma spectabile and Etheostoma radiosum (Pisces: Percidae). Am.

Midi. Nat. 704:412-415. Lindquist, David G., J. R. Shute and P. W. Shute. 1981. Spawning and nesting

behavior of the waccamaw darter, Etheostoma perlongum. Environ. Biol.

Fishes 6:177-191. Mathur, Dilip. 1973. Food habits and feeding chronology of the blackbanded

darter, Percina nigrofasciata (Agassiz), in Halawakee Creek, Alabama.

Trans. Am. Fish. Soc. 702:48-55. Reynolds, William W. 1977. Fish orientation behavior: an electronic device for

studying simultaneous responses to two variables. J. Fish. Res. Board Can.

54:300-304. (ed.). 1979. Symposium on thermoregulation in ectotherms. Am.

Zool. 79:191-384. , and M. E. Casterlin. 1976. Locomotor activity rhythms in the bluegill

sunfish, Lepomis macrochirus. Am. Midi. Nat. 9(5:221-225. , and 1977. Diel activity in the yellow bullhead. Prog. Fish- Cult. 59:132-133. , and 1978. Ontogenetic change in preferred temperature and

diel activity of the yellow bullhead, Ictalurus natalis. Comp. Biochem. Physiol. 54/1:409-411.

, and 1979. Behavioral thermoregulation and the 'final pref-

erendum' paradigm. Am. Zool. 79:21 1-224. , and 1980. The role of temperature in the environmental

physiology of fishes, pp. 497-518 in M. A. Ali (ed.). Environmental Physi- ology of Fishes. Plenum Press, New York. 723 pp. , and D. A. Thomson. 1974. Responses of young Gulf grunion, Leu-

resthes sardina, to gradients of temperature, light, turbulence and oxygen. Copeia 1974(3):747-758.

, and and M. E. Casterlin. 1977. Responses of young Califor- nia grunion, Leuresthes tenuis, to gradients of temperature and light. Copeia 1977(1):144-149.

Accepted 19 May 1982

Helminths of Some Seabirds from North Carolina

Ronald W. Mobley and Grover C. Miller

Department of Zoology, North Carolina State University, Raleigh, North Carolina 27650

ABSTRACT. — Eighty-five birds of the orders Procellariiformes, Pelecaniformes, and Charadriiformes were examined for helminths between May 1977 and September 1979. At least nine species of helminths were recovered — one species of Digenea, at least six of Cestoda, one Nematoda, and one Acanthocephala. The following new host records were obtained: Opisthovarium elongation and Falsifilicollis altmani in the common tern; Tetrabothrius laccocephalus in the north- ern fulmar and Audubon's shearwater; T. minor in Cory's shearwater; T sp. in the red-billed tropicbird; and T. filiformis , Choanotaenia sp., and Contracaecum sp. in Wilson's storm petrel. Feeding habits of the hosts were directly reflected by their helminth fauna. Procellariiform and pelecaniform hosts showed a predominance of helminths that probably use fish or squid as intermediate hosts, whereas charadrii- form hosts contained helminths whose intermediate hosts are either snails or terrestrial crabs.

INTRODUCTION

Much literature is available on the helminths of sea birds. Most early studies were concerned primarily with morphology and taxonomy. The best pre- 1900 descriptions of these helminths are found in the works of Diesing (1850), Fuhrmann (1899), and Loennberg (1899) as reported by Baer (1954). The twentieth century marked the beginning of polar exploration and consequently provided increased access to sea birds for parasitological study. Earliest reports were by Linstow (1905) on the Russian Polar Expedition, 1900-1903; Raillet and Henry (1912) on the cestodes from the French Antarctic Expedition; and Leiper and Atkinson (1914, 1915) on helminths recovered from the British Antarc- tic (Terra Nova) Expedition, 1910-1913. Later reports included those of Fuhrmann (1921) on the German South Polar Expedition, 1901-1903; Johnston (1937) and Mawson (1953) on the Australian National Antarctic Research Expeditions; and Johnston and Mawson (1945) and Prudhoe (1967) on parasitic helminths collected by the British, Austral- ian and New Zealand Antarctic Research Expedition, 1929-1931.

Cestodes have probably been the most studied helminths of sea birds and literature is abundant. Spatlich (1909), Ransom (1909), Nybelin (1916), and Szpotanska (1925, 1929) all made important descriptive con- tributions. Linton (1927) in America, Tseng (1932, 1933) in China, Yamaguti (1935) in Japan, and Joyeux and Baer (1935, 1939) in France described specimens from their respective countries. Later studies cen-

Brimleyana No. 7:6 1 -68. July 1 98 1 . 61

62 Ronald W. Mobley and Grover C. Miller

tered more on taxonomic revisions and included works by Johnston (1935), Lopex-Neyra (1943, 1952), Wardle et al. (1952, 1974), and Baer (1954). The trematodes of sea birds were extensively studied by Linton (1928), Yamaguti (1958, 1971) and Cable et al. (1960). Van Cleave (1918, 1934, 1939, 1947), Perry (1942), Reish (1950), and Petrotchenko (1958) provided excellent information of the taxonomy and biology of the acanthocephalans of these birds. Recent contributions to sea bird helminthology included those of Deblock (1966), Jones and Williams (1967, 1968, 1969), Threlfall (1971), Ellis and Williams (1973), Williams and Ellis (1974), Riley and Owen (1975), and Bourgeois and Threlfall (1979).

A variety of literature on the identification and ecology of the bird hosts is available in publications by Bent (1921, 1922), Murphy (1936), Roberts (1940), Palmer (1941), Robertson (1964), and Watson (1966).

The objectives of our study were to (1) determine the gastrointesti- nal helminth fauna of certain sea birds, and their prevalence and inten- sity ranges; and (2) correlate differences in helminth fauna with aspects of host biology such as feeding habits and geographical range.

METHODS AND MATERIALS

Eighty-five birds were collected between May 1977 and August 1979, including seven species of Procellariformes, one species of Peleca- niformes, and two species of Charadriiformes. All were collected from the Atlantic Ocean east of and adjacent to Oregon Inlet, North Caro- lina. Twelve- or sixteen-gauge shotguns and either No. 2 or No. 4 high velocity shot were used to collect the hosts.

Wilson's storm petrels, Oceanites oceanicus, were attracted into range using "chum" consisting of poultry offal. Fulmars Fulmarus gla- cialis; shearwaters, Puffinus spp.; and gadfly petrels, Pterodroma spp., were generally unresponsive to chum slicks but were occasionally encountered in close proximity to commercial fishing vessels. Red-billed tropicbirds, Phaethon aethereus, were collected singly near the collect- ing vessel. Common terns, Sterna hirundo, and bridled terns Sterna anaethetus, were collected either on the wing or as they rested on flot- sam. After collection each bird was labeled with respect to time and date, placed in an individual plastic bag, and put on ice. Hosts were hard forzen within eight hours of collection and examined as soon as possible, usually within 48 hours. Bird identifications were based on Watson (1966).

Each bird was incised midventrally from cloaca to sternum. The alimentary tract, from gizzard to rectum, was removed intact to a glass dish containing saline, where the tract was then longitudinally dissected. The gut contents were agitated, rinsed, and allowed to settle. The super-

North Carolina Seabird Helminths 63

natant was decanted into another dish for examination with a stereo- scopic microscope. All original sediment was washed repeatedly and decanted as above. Gross examination of intestinal tissue was made in an attempt to find attached helminths. Trematodes, cestodes, and acan- thocephalans were fixed and stored in AFA (Alcohol-Formalin-Acetic Acid), stained with Semichon's acetocarmine, cleared in methyl salicy- late, and mounted in Kleermount. Nematodes were fixed in 70 percent alcohol, stored in glycerin-alcohol, cleared in glycerin, and mounted in glycerin jelly.

RESULTS AND DISCUSSION

Nine species of parasites were recovered (Table 1). One species of digenetic trematode and one species of acanthocephalan were recovered from the common tern; five species of cestodes were found in various species of procellariiforms, and one species in the red-billed tropicbird; and one species of nematode was found in two Wilson's storm petrels. No helminths were recovered from the one black-capped petrel, Ptero- droma hasitata, or the seventeen bridled terns, Sterna anaethetus, examined.

Sea birds vary greatly in the degree to which they are associated with the open sea. Truly pelagic species may encounter land only during breeding periods, while others, such as some species of gulls and terns, may live and feed inshore except during migrations. Resulting differen- ces in habits will affect exposure to the helminth fauna.

Although fish probably represent a major food item for most spe- cies examined, the coast-inhabiting terns also secondarily eat a variety of crustaceans, insects, worms, and molluscs (Watson 1966). The acan- thocephalans found in common terns probably used crustacean inter- mediate hosts (Reish 1950; McDonald 1969). In 2 individuals the number of acanthocephalans recovered exceeded 200, thus indicating extensive feeding on the intermediate host.

The procellariiform and pelecaniform hosts are all pelagic and primarily consume fish and squid (Watson 1966, 1975). The tetrabo- thriid cestodes and the heterocheilid nematode, Contracaecum sp., were probably transmitted through fish or squid intermediate hosts (Baer 1954; Ellis and Williams 1973; Yamaguti 1959). However, the cestode, Choanotaenia sp., recovered from Wilson's storm petrel, is from a genus known to use insects as intermediate hosts (Yamaguti 1959). Infections may have occurred during breeding, perhaps by ingestion of infected insects associated with the nesting sites.

A high prevalence of infection may best be explained by birds defe- cating in an area where suitable intermediate hosts are present and their ingestion is likely. A highly prevalent helminth might tend to be less

64

Ronald W. Mobley and Grover C. Miller

Table 1 . Prevalence and intensity range of gastrointestinal helminths of sea birds from North Carolina.

Taxon (No. examined)

Prevalence of infection (%)

Intensity range

Order Procellariiformes northern fulmar (1) Cestoda: Tetrabothrius laccocephalus ' Tetrabothrius procerus

Cory's shearwater (7) Cestoda: Tetrabothrius minor ' Tetrabothrius laccocephalus

greater shearwater (3) Cestoda: Tetrabothrius laccocephalus

Audobon's shearwater (5) Cestoda: Tetrabothrius laccocephalus '

South Trinidad petrel (2) Cestoda: Tetrabothrius sp.

Wilson;s storm petrel (37) Cestoda: Tetrabothrius filiform is ' Choanotaenia sp. ' Unidentified fragments

Nematoda: Contraceacum sp. '

Order Pelecaniformes red-billed tropicbird (2) Cestoda: Tetrabothrius sp. '

Order Charadriiformes common tern (11) Digenea:

Opisthovarium elongatum ' Acanthocephala: Falsifilicollis altmani '

100	>10
100	>10
29	1-5
15	10

100

40

100

6

8

11

1 ->10

1-2

1 1-4

50 2

9 3 pairs in copuh

45 2 - >200

1 New host record

North Carolina Seabird Helminths 65

host specific and could thus be transmitted through a variety of hosts. Conversely, if the prevalence of a particular helminth is low, host speci- ficity would tend to be higher, definitive or intermediate host popula- tions may be sparse, or definitive-intermediate host interactions might be poorly developed.

The prevalence of Falsifilicollis altmani in the common tern is rela- tively high. Correspondingly, the definitive host population is fairly dense in the coastal areas of North Carolina. While the intermediate host is unknown, a variety of crustacean species occurs in the area. The parasite demonstrates only limited specificity; it has been found in at least four other bird genera (Ehrhardt 1966, Yamaguti 1963).

An example of low infection is that of Tetrabothrius filiformis in Wilson's storm petrel. Host population density is relatively low except on the breeding grounds, and even there the birds are far-ranging in feeding habits (Bent 1922). Again, the intermediate hosts are unknown, but populations of fish and squid vary from sparse in warm open waters to dense in the cold waters surrounding the breeding grounds (Watson 1975). Like other tetrabothriids, T. filiformis is probably transmitted through fish or squid, both of which seem to be secondary to fish oil slicks and krill in the diet of Wilson's storm petrel (Watson 1975). Krill may be an intermediate/ paratenic host.

The levels of infection in the northern fulmar and the shearwaters reported here and elsewhere (Bourgeois and Threlfall 1979) at first seem high for birds of the open sea. The tetrabothriid species (T. laccocepha- lus, T. minor, and T. procerus) recovered from these hosts, however, are found in a variety of related hosts and over a wide geographical range (Baer 1954). This range would seem to indicate that the helminths can use a large variety of intermediate host species or an intermediate host with a wide range. These factors, combined with the hosts' habits of feeding almost exclusively on squid and fish (Watson 1966, 1975), seem to support the relatively high rates of parasitism in these birds.

Most of the cestodes recovered constitute new locality records, and several cestode species were found in hosts coming from both north and south polar regions. In conclusion, it appears that the offshore waters of North Carolina provide an area of intergradation for northerly/ south- erly seabirds and their parasites. We are doing additional work on these hosts and their parasites.

ACKNOWLEDGMENTS.— Appreciation is extended to David S. Lee, Steven P. Platania, and other members of the North Carolina State Museum of Natural History staff for providing hosts and host identifi- cations. Acknowledgment is due Micou M. Browne for his assistance and suggestions in the preparation of this material.

66 Ronald W. Mobley and Grover C. Miller

This is Paper No. 7089 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, North Carolina.

LITERATURE CITED

Baer, Jean G. 1954. Revision taxonomique et etude biologique des Cestodes de la famille des Tetrabothriidae, parasites d Oiseaux de haute mer et de mammiferes marins. Mem. Univ. Neuchatel Ser. inquarto 1:1-21.

Bent, Arthur C. 1921. Life histories of North American gulls and terns; order Longipennes. U. S. Natl. Mus. Bull. 107. 345 pp.

1922. Life histories of North American petrels and pelicans and their

allies: order Tubinares and order Steganopodes. U. S. Natl. Mus. Bull. 121. 343 pp.

Bourgeois, C. E., and W. Threlfall. 1979. Parasites of the greater Shearwater (Puffinus gravis) from Newfoundland, Canada. Can. J. Zool. 57(7): 1355-1357.

Cable, Raymond M., R. S. Connor and J. W. Balling. 1960. Digenetic trema- todes of Puerto Rican shore birds. Scientific survey of Puerto Rico and the Virgin Island 77(2): 187-239.

Deblock, Stephanie. 1966. Six cestodes d oiseaux de mer ou de rivage de 1 hemisphere austral (ile Europa). Description de Tetrabothrius mozambi- quus n. sp. et de Baerbonaia baeribonae n. gen., n. sp. Mem. Mus. Nat. Hist. Nat. Paris. 41(\): 103-1 23.

Diesing, C. M. 1850, Systema Helminthum I. Vinobonae. Weinheim, Germany. 588 pp.

Ehrhardt, Warren R. 1966. Some helminths from two species of gulls from North Carolina. Unpubl. M.S. Thesis, N. C. State Univ., Raleigh. 225 pp.

Ellis, Catherine, and I. C. Williams. 1973. The longevity of some species of helminth parasites in naturally acquired infections of the lesser blackbacked gu\\,Larus fuscus L., in Britain. J. Helminthol. 47(3):329-338.

Furhmann, O. 1899. Mitteilungen uber Vogeltaenien. Centralbl. Bakt. Parasitol. 2(5:622-627.

1921. Die Cestoden der Deutschen Sudpolar-Expedition 1901-1903.

Deutsche Sudpolar Exped. 1901-1903, 16 Zoologie 5:469-524.

Johnston, Harvey. 1935. Remarks on the Cestode genus Porotaenia. Trans. R. Soc. S. Aust. 59:164-167.

1937. Cestoda. Sci. Rep. Aust. Antarctic. Exped. 191 1-1914 70(4):74 pp.

, and P. M. Mawson. 1945. B.A.N.Z. Antarctic Research Expedition

1929-1931. Vol. V, Part 2. Parasitic Nematodes. Adelaide, pp. 73-160.

Jones, N. V., and I. C. Williams. 1967. The cestode parasites of the Sheat-bill, Chionis alba (Gmelin), from Signy Island, South Orkney Islands. J. Hel- minthol. 47:151-160.

, and 1968. The Trematode Parasites of the sheathbill, On- onis alba (Gmelin), from Signy Island, South Orkney Islands. J. Helminthol. ¥2:65-80.

, and 1969. The nematode and acanthocephalan parasites of

the sheathbill, Chionis alba (Gmelin), at Signey Island, South Orkney

Islands, and a summary of host parasite relationships in the sheathbill. J. Helminthol. 43:59-61.

North Carolina Seabird Helminths 67

Joyeux, C. H., and J. G. Baer. 1934. Sur quelques Cestodes de France. Arch.

Mus. Nat. Hist. Nat. (Paris) 77:165-170. , and 1939. Sur quelques cestodes de Charadriiformes. Bull.

Soc. Zool. Fr. 64:171-187. Leiper, R. T., and E. L. Atkinson. 1914. Helminths of the British Antarctic

Expedition 1910-1913. Proc. Zool. Soc. London, pp. 222-226. , and 1915. Parasitic worms with a note on a free-living

nematode. British Antarctic (Terra Nova) Expedition, 1910. Natural His- tory Report, Zoology 2: 19-60. Linstow, Otto von. 1905. Helminthen der Russichen Polar-Expedition 1900-

1903. Mem. Acad. Imp. Sci. St. Peterbs. Phys. Math. 75:1-17. Linton, E. 1927. Notes on Cestode parasites of birds. Proc. U. S. Natl. Mus. 70.

73 pp. 1928. Notes on trematode parasites of birds. Proc. U. S. Natl. Mus.

73. 1-11. Loennberg, E. 1899. Ueber einige cestoden aus dem Museum zur Bergen. Ber-

gens Mus. Arb. 4: 1-23. Lopez-Neyra, Carlos R. 1942. Division del genero Hymenolepis Weinland (S.I.)

en ostros mas naturales. Rev. Iber. Parasitol. 2:46-93, 1 13-256. 1952. Byrocoelia albaredai n. sp. relaciones con Tetrabothriidae y

Dilepididae. Rev. Iber. Parasitol. 72:319-344. McDonald, Malcolm E. 1969. Catalogue of helminths of waterfowl (Anatidae).

U. S. Fish Wildl. Serv. Spec. Sci. Rep. Wildl. 726. 692 pp. Mawson, Patricia M. 1953. Parasitic Nematoda collected by the Australian

National Antarctic Research Expedition: Heard Island and Macquarie

Island, 1948-1951. Parasitol. 45:219-297. Murphy, Robert C. 1936. Oceanic Birds of South America. Amer. Mus. Nat.

Hist., New York. 2 vols. Nybelin, Orvar. 1916. Neue Tetrabothriiden aus Vogel. Zool. Anz. 47:297-301. Palmer, R. S. 1941. A behavior study of the Common Tern {Sterna hirundo

hirundo L.). Proc. Boston Soc. Nat. Hist. 42:1-119. Perry, M. L. 1942. A new species of the acanthocephalan Filicollis. J. Parasitol.

25:385-388. Petrotchenko, Vladamir I. 1958. Acanchocephala of domestic and wild animals.

Rabot. Gel Mintol. 80 Let. Skrjabin, Akad. Sel-Skokhoziaistv. Nauk.

7:10458. Prudhoe, Stephan. 1969. Cestodes from fish, birds and whales. Report of the

B.A.N.Z. Antarctic Research Expedition, 1929-31 9:17:193. Railliet, A., and A. Henry. 1912. Helminthes recueillis par 1 expedition antarc-

tique francaise du Pourquoi Pas I. Cestodes d Oiseaux. Bull. Mus. Nat.

Hist. Nat.:35-39. Ransom, H. B. 1909. The teanioid cestodes of North American Birds. U. S.

Natl. Mus. Bull. 69. 129 pp. Reish, Donald. 1950. Preliminary note on the life cycle of acanthocephalan,

Polymorphus kenti Van Cleave, 1947. J. Parasitol. 56:496. Riley, John, and R. W. Owen. 1975. Competition between two closely related

Tetrabothrius cestodes of the fulmar (Fulmarus glacia/is L.) Z. Parasitenk. D.

46(3):221-228.

68 Ronald W. Mobley and Grover C. Miller

Roberts, B. 1940. The life cycle of Wilson's Petrel. British Graham Land

Exped., Sci. Rep. 7:141-194. Robertson, W. B., Jr. 1964. The terns of the Dry Tortugas. Bull. Fla. State

Mus. Biol. Sci. 5:1-14. Spatlich, Walter. 1909. Untersuchungen uber Tetrabothrien. Ein Beitrag zur

Kenntnis des Cestoden-korpers. Zool. Jahrb. Anat. 25:539-594. Szpotanska, I. 1925. Etude sur les Tetrabothriides des Procellariiformes. Bull.

Acad. Pol. Sci. Lt. Ser. B.:673-727. 1929. Recherches sur quelques Tetrabothriides d Oiseaux. Bull.

Acad. Pol. Sci. Lt. Ser. B.: 129-152. Threlfall, William. 1971. Helminth parasites of alcids in the northwestern North

Atlantic. Can. J. Zool. 49(4):46 1-466. Tseng, S. 1932. Studies on avian cestodes from China. Part I. Cestodes from

Charadriiform birds. Parasitol. 24:87-106. 1933. Studies on avian cestodes from China. Part II. Cestodes from

Charadriiform birds. Parasitol. 24:500-511. Van Cleave, Harley J. 1918. The Acanthocephala of North American birds.

Trans. Am. Microsc. Soc. 37:19-47. 1934. Observations on the status of certain genera of Acanthoce- phala, chiefly from birds. J. Parasitol. 20:324. 1939. A new species of acanthocephalan genus Polymorphus and

notes on the status of the name Profilicollis. J. Parasitol. 25:121-131. 1947. Analysis of distinction between the acanthocephalan genera

Fillicollis and Polymorphus with descriptions of a new species of Polymor- phus. Trans. Am. Microsc. Soc. 6(5:302-313.

Wardle, R. A., and J. A. McLeod. 1952. The Zoology of Tapeworms. Univ. Minn. Press, Minneapolis. 780 pp.

Wardle, R. A., J. A. McLeod and S. Radinovsky. 1974. Advances in the Zool- ogy of Tapeworms, 1950-1970. Univ. Minn. Press, Minneapolis. 274 pp.

Watson, George E. 1966. Seabirds of the Tropical Atlantic Ocean. Smithsonian Identification Manual. Smithsonian Press, Washington. 120 pp.

1975. Birds of the Antarctic and Sub-Antarctic. Am. Geophy.

Union, Washington. 350 pp.

Williams, I. C, and C. Ellis. 1974. The helminth parasites of the sheathbill, Chionis alba (Gmelin), and the diving petrels, Pelecanoides georgicus (Murphy and Harper) and P. urinatrix (Gmelin), at Bird Island, South Georgia. J. Helminthol. 48(3): 195-197.

Yamaguti, Satyu. 1935. Studies on the helminth fauna of Japan. Part VI. Ces- todes of birds I. Jpn. J. Zool. 6:184-232.

1959. Systema Helminthum. Vol. II. Cestodes of Vertebrates.

Interscience Publ., New York. 860 pp.

1961. Systema Helminthum. Vol. III. The Nematodes of Vertebrates.

Interscience Publ., New York. 1261 pp.

1963. Systema Helminthum. Vol. V. Acanthocephala. Interscience

Publ., New York. 423 pp.

1971. Synopsis of Digenetic Trematodes of Vertebrates. Keigaku

Publ. Co., Tokyo. 1074 pp.

Accepted 29 January 1982

Life History of a Coastal Plain Population of the Mottled Sculpin, Coitus bairdi (Osteichthyes: Cottidae),

in Delaware

Fred C. Rohde

Charles T. Main, Inc., Prudential Center, Boston, Massachusetts 02199

AND

Rudolf G. Arndt

Faculty of Natural Sciences and Mathematics, Stockton State College, Pomona, New Jersey 08240

ABSTRACT. — Aspects of the biology of a Coastal Plain population of Cottus bairdi in a lowland forest stream in Delaware were examined during 1973-81. Cottus bairdi occurs primarily over a gravel substrate; naturally-occurring rocks are absent. Individuals are shorter-lived and attain sexual maturity at a smaller size than in other studied popula- tions. Two age groups, and a presumed third, were found. Only 33% of age group 0 survives to spawn in age group I. Spawning occurs in February and March at a water temperature of 6.5° to 12.5° C, and mean ovum diameter is 2.15 mm. There is a strong relationship between ova number and standard length (Y = -157.11 + 6.80 SL). Fecundity is lower than that reported for most populations of C. bairdi. Most eggs are deposited in single-spawn clusters, with a mean egg number of 134.2 per cluster. Condition factor indicates that males are generally more robust than females. Most growth in length occurs in age group 0, males are longer than females, and there is a highly significant difference in sex ratio of mature specimens taken.

Dominant food is trichopteran and dipteran larvae, and plecopte- ran nymphs; crustaceans, ephemeropteran nymphs, and coleopteran larvae are of minor importance. There is seasonal variation in food items. Diet differs only slightly as a function of sculpin size.

INTRODUCTION

The mottled sculpin, Cottus bairdi, is widely distributed in the United States and Canada (Lee 1980) and there are numerous studies of aspects of its life history (Gage 1878; Smith 1922; Hann 1927; Koster 1936, 1937; Bailey 1952; Daiber 1956; Ludwig and Norden 1969; Patten 1971; Nagel 1980). There are, however, only four known Atlantic Coastal Plain populations, all located in the Nanticoke River system, Chesa- peake Bay drainage. We studied one of these, in Butler Mill Branch and

Brimleyana No. 7:69-94. July 1 98 1 . 69

70 Fred C. Rohde and Rudolf G. Arndt

its tributaries, near Seaford, Sussex County, Delaware, during 1973-75 and 1978-81 and report here on habitat, reproduction, age and growth, food habits, and abundance. Franz and Lee (1976) reported on a popu- lation in Caroline County on the Eastern Shore of Maryland, 4.0 km north of Federalsburg and 15.2 km northwest of Butler Mill Branch, and briefly considered habitat, food habits, and size. In spring 1981 we discovered a third population, in upper Sullivan Branch and the lower parts of its tributaries, Wolfpit and Raccoon branches, 6.4 km north of Federalsburg, Caroline County, Maryland, and a fourth in Skinners Run, at a point just upstream of Route 307 and 4.0 km southwest of Federalsburg, Dorchester County, Maryland. All data herein refer to the Butler Mill Branch population, unless otherwise specified.

MATERIALS AND METHODS

Specimens of C. bairdi (N = 1009) were collected with a 3.0 m X 1.2 m, 3.2 mm mesh nylon flat seine. All habitats were sampled. Collections were made on 28 October and 4 November 1973; 20 January, 17 Febru- ary, 17 March, 20 July, 8 September and 30 November 1974; 2 and 23 March 1975; 7 September and 26 November 1978; 16 March and 25 July 1979; 9, 15, 22 and 30 March, 13 April, 4 and 20 May, 8 and 28 June, 12 and 25 July, and 9 August 1980; and 22 and 27 February, 7, 14, 22, 26 and 29 March, 5, 11, 18 and 25 April, and 19 May 1981, between 1050 and 2000 hours. Fish were preserved in 10% formalin and stored in 40% isopropyl alcohol until examined. For conservation pur- poses, the last specimens taken in each collection, regardless of sex, size, and reproductive condition, were usually released. Through 1979, only relatively small samples were taken, because of the presumed small size of the population. Sampling was intensified in 1980 and 1981 to pin- point spawning time, locate eggs, and collect larvae. In 1981, fish were primarily observed in the field, and most of those collected were released alive.

An additional 41 adult C. bairdi were taken by seine in spring 1981 at the new Caroline County locality, 6 adults at the Dorchester County locality, and 1 adult at the locality given by Franz and Lee (1976); these were preserved.

Air and water temperature, pH (Hach Kit), and dissolved oxygen (Yellow Springs meter) were recorded at time of capture. To help char- acterize the habitat in areas particularly frequented by C. bairdi, semi- quantitative substrate samples of about 8 liters were taken from Butler Mill Branch and about 1.2 liters from an upper tributary. The samples were dried and sieved, and the percentage by weight of several gravel/ sand particle size categories determined.

Coitus Life History in Delaware 71

Gonadal development was determined by calculating the gonoso- matic ratio (gonad weight as a percentage of total weight) of 235 females and 121 males taken during 1973-80. This sample included only sculpins that could be sexed and with gonads of 0.001 g or heavier (young-of-the-year from September, October, and November collec- tions, and all adults). Gonads were weighed to the nearest 0.001 g. Fecundity of 67 pre-spawning females collected in January (20) and February (3) 1973, March (10) 1979, and March (34) 1980 was deter- mined. Ten ova from each of 57 mature females were measured to the nearest 0.1 mm using an ocular micrometer, and the ova growth rate determined. Deviation from a 1:1 ratio of ova number between right and left ovary in 22 fish was tested with chi-square, as was deviation from a 1:1 sex ratio. Standard length (SL) and total length (TL) were measured to the nearest mm and total body weight to the nearest 0.01 g.

Eggs were collected where adults were most common. Pieces of concrete block, bricks, and water-logged wood suitable as egg attach- ment sites were provided on 9 March 1980 and 22 February 1981. Color was recorded on live eggs; other data are from preserved eggs. Attempts to collect larvae with a fine-mesh dip net and a 0.5 mm plankton net were made in the same area.

Age was determined by the length-frequency method. Although otoliths have been used to determine age in sculpins (Koster 1936: Lud- wig and Norden 1969; Patten 1971; Petrosky and Waters 1975), our results using otoliths from both fresh and preserved C. bairdi were inconclusive. The relationship between growth in weight and in length for specimens that could be sexed was determined by fitting a regression line on the logarithms of mean weights and SL for 3 mm intervals. Fitness was determined with the coefficient of condition (K) using the formula K - W X 1Q5, where W is weight (g) and L is SL (mm). L3

Based on examination of 539 stomachs, we determined the number of food items in five fish SL groups (< 20 mm, 20-29 mm, 30-39 mm, 40-49 mm, > 49 mm), and the percent occurrence of food by month.

Voucher specimens from all localities were deposited in the Academy of Natural Sciences of Philadelphia (ANSP 145647, 145648, and others), and specimens from the Butler Mill Branch locality were also deposited at Iowa State University (ISU 1995, 1996), and the Uni- versity of Florida (UF 30136, 30137).

STUDY AREA

The Butler Mill Branch system lies between 8 km northwest and 4 km southwest of Seaford, Sussex County, Delaware. It consists of Horse

72 Fred C. Rohde and Rudolf G. Arndt

Pen, Green Briar, and Butler Mill branches, and several unnamed tribu- taries. Its maximum length is 10.5 km (9.0 km straight-line) and its area 23.8 km2. The creeks are bordered by mature lowland forest of red maple, Acer rubrum; American holly, Ilex opaca; green ash, Fraxinus pennsylvanica; tulip tree, Liriodendron tulipifera; sweetgum, Liquidamber styraciflua; black tupelo, Nyssa sylvatica; and sweetbay, Magnolia virgin- iana; some upper parts, however, pass through agricultural fields.

In the upper system the dominant creek substrate is mud and mud- sand, and typical stream width and depth in spring are 1 m or less, and some 20 cm, respectively. Gravel patches appear downstream and become more common lower in the system. Greatest stream width is 7 m, at a point below an impoundment (Craigs Pond) on lower Butler Mill Branch. The uppermost waterways sometimes dry completely, and areas just downstream of these form pools. Current and flow in the lower system are strong all year. Storms often cause local creekside flooding. Aquatic vegetation, consisting of several vascular species and filamentous green algae, is local and often luxuriant. Rocks, usually prominent in C. bairdi habitat, are absent.

Coitus bairdi is found only in areas of permanent and pronounced flow (Fig. 1). In its upstream distribution, C. bairdi is found only over gravel patches, where concomitantly there is a stronger current. Creek width here is often only 1 m, with depths to 5 cm. Patches are typically about 1 m long and 0.3 m wide, comprised of fine gravel and sand, and located several dozen meters apart. Cottus bairdi occurs but sparingly on these patches. Gravel patches and C. bairdi are both progressively more common farther downstream in Horse Pen, Green Briar, and Butler Mill branches. Cottus bairdi is not found below or just above Craigs Pond, although prior to impoundment it likely did occur in both areas.

All preserved specimens and related data, unless otherwise specified, were collected in a sampling area located in the downstream part of the study area (Fig. 1), over and near two large and easily accessible gravel riffles (of 4.6 m X 6.5 m and 4.6 m X 28.0 m). The strongly meandering stream here is 4 to 5 m wide, with a substrate of alternating gravel riffles, mud-bottom pools, and sand. The riffles are several cm deep, the pools to 1.2 m, and sand occurs at intermediate depths. The banks are steep and frequently undercut, with masses of exposed fine tree roots. The south bank abuts lawns behind homes, and the north side is mostly forested. Concrete block riprap has been placed along the developed shore to prevent erosion. Dominant aquatic vegetation is pondweed, Potamo- geton sp.; water starwort, Callitriche palustris; water purslane, Ludwigia palustris\ bur reed, Sparganium sp.; and unidentified filamentous green algae. Vegetation occurs primarily in cleared, sunlit areas near the homes and highway. Cottus bairdi is abundant throughout. The water, usually

Cottus Life History in Delaware

73

i

N

0 .5 1.0

Kl LOMETERS

...... indicates distri- bution of Cottus bairdi in Butler Mill Branch and tributaries • •• — indicates sampling

SEAFORD

-no

7T\

SAMPLING

Fig. 1. The Butler Mill Branch system near Seaford, Sussex County, Delaware, showing Cottus bairdi distribution and sampling area.

clear, is turbid after rain. Temperature in the sampling area ranged from 5.5° C (February) to 20.0° C (July), pH from 5.7 to 7.0 (typically 6.5), and dissolved oxygen concentrations from 9.1 to 12.8 ppm.

Stream flow data were taken in the sampling area on 22 dates between 30 June 1955 and 16 April 1969 by the U.S. D.I. Geological Survey, Water Resources Division (Robert H. Simmons, pers. comm.) at low-flow conditions (after periods of no rain for four or five days). Aver- age stream cross-section area was 0.76 m2 (range 0.20-2.17), mean flow 0.16 m3/sec (0.04-0.46), and mean water velocity 0.21 m/ sec (0.10-0.29). The lowest values were from summer, the highest from spring and late fall.

Fifteen fishes were taken with C. bairdi (number of times in paren- theses) in a total of 26 collections made in the sampling area during 1973-80: Lampetra aepyptera (20), Anguilla rostrata (18), Umbra pyg- maea (11), Esox americanus (1), Esox niger (6), Notemigonus crysoleucas (4), Erimyzon oblongus (17), Ictalurus natalis (2), Ictalurus nebulosus (1),

74 Fred C. Rohde and Rudolf G. Arndt

Aphredoderus sayanus (10), Acantharchus pomotis (1), Enneacanthus gloriosus (3), Lepomis gibbosus (5), Lepomis macrochirus (6), and Etheostoma olmstedi (26).

Cottus bairdi habitat in Caroline and Dorchester counties, Mary- land, including physicochemical characteristics, is as described for Butler Mill Branch. Maryland fish species-associates include some of those listed previously, plus Etheostoma fusiforme and Percaflavescens.

RESULTS AND DISCUSSION

Reproduction

Spawning occurs in February and March. Two females taken 17 March 1974 each contained only two ova, indicating the end of spawning. On 2 March 1975, 1 of 3 1 females was spent and, although other females contained ova, there were fewer than in 1974, 1979, and 1980 pre- spawning samples, indicating spawning was under way. Spawning was completed by 23 March; all ovaries were empty except for some ova being resorbed. Spawning in 1979 was later, and females collected on 16 March still possessed a full ova complement. In 1980 females spawned primarily between 9 and 22 March: on 9 March none had spawned, on 15 March 2 of 22 were spent, by 22 March all but 1 were spent, and on 30 March all taken were spent. Examination of specimens in the field in 1981 yielded the following ratios of females heavy with eggs to spent females: 22 February, 13:1; 27 February, 10:2; and 7 March, 7:3. On subsequent dates all taken had completed spawning.

February and March spawning is generally earlier than that reported for C. bairdi. Other reports based on direct (deposited eggs) or indirect (mature ova or specimens in breeding condition) evidence are: Nagel (1980), Tennessee, early April; Ludwig and Norden (1969), Wisconsin, 1 April to 3 May; Ricker (1934), Ontario, middle of May; Bailey (1952), Montana, most of June; Gage (1878), New York, April; Koster (1936), New York, April to June; Smith (1922), Michigan, April and May; Rob- ins (1954), Appalachian region, late March to early May. Pflieger (1975) found presumed C. bairdi eggs in Ozark streams from early November to late February. William L. Pflieger (pers. comm.) has confirmed records of eyed eggs from 2 February 1971 and of males guarding egg masses with embryos far enough advanced to show movement on 23 and 29 February 1971, which indicates spawning occurred at least as early as January. This early spawning, however, may not be attributable to C. bairdi, as most Ozark populations are considered to represent an undescribed species.

Ovary weight as a measure of gonad development during 1973-75 was low in October and November 1973, increased rapidly in January, and peaked in February 1974 (Fig. 2, Table 1). The gonosomatic ratio

Cottus Life History in Delaware

75

SONDJ FMAMJ JASONDJ FMAMJ JA 1973 1974 1975

197B 1979 1980

MON THS

Fig. 2. Relationship between gonad weight and body weight in male and female Cottus bairdi from Butler Mill Branch, Sussex County, Delaware, October 1973- March 1975 (solid lines) and September 1978-August 1980 (broken lines).

dropped markedly in March 1974, at spawning, was low through summer, and increased again in November. The maximum value was reached early in March 1975, and was followed rapidly in mid-March by another sharp post-spawning decrease. The gonosomatic ratio pattern for males was similar to that for females, but monthly differences were not as pronounced (Fig. 2, Table 1). Maximum development in 1974 was in January, followed by a summer decline and a fall increase, and values

76 Fred C. Rohde and Rudolf G. Arndt

Table 1. Average combined monthly gonosomatic ratios (expressed as %) for Cottus bairdi from Butler Mill Branch, Sussex County, Delaware, 1973- 1980. Number of specimens in parentheses.

	1973-75	1978-80
Month	Male	Female	Male	Female
January	1.52 (4)	7.57 (20)
February	1.40 (2)	19.51 (3)
March	1.05(15)	20.73 (33)	0.85 (35)	16.51 (71)
May			0.09 (3)	0.65(14)
June			0.09 (5)	0.74(17)
July	0.21 (5)	0.57(14)	0.16 (6)	0.56(13)
August			0.20 (1)	0.53 (1)
September	0.53 (3)	0.71 (14)	0.16(14)	0.49 (5)
October	1.21 (7)	1.46 (4)
November	1.26(17)	2.18(25)	1.94 (4)	2.68 (1)

remained high from November through March. Both sexes in 1978-80 showed patterns similar to those of 1973-75. The March 1979 value for females, however, was markedly higher than that of other years, and reflects a later spawning in 1979.

Fecundity, a general term that refers to number of ova produced, is restricted in this study to the number of mature ova present in the ovaries from January to mid-March. Mean ova number per female (N = 67) is 93.7 (range 37-156), and mean female SL is 36.9 mm. The regression equation for the relationship between SL and ova number (Y) is: Y = -157.1 1 + 6.80 SL (Fig. 3). The correlation coefficient, r = 0.90, indicates a strong positive relationship between ova number and SL (P< 0.001).

Fecundity of the Butler Mill Branch sculpins is low when compared with data for other species of Cottus and for other populations of C. bairdi in Williams (1968), Ludwig and Norden (1969), Patten (1971), Foltz (1976), and Nagel (1980). Exceptions are the diminutive C. pyg- maeus, with a range of 30 to 43 ova (Williams 1968), and C. bairdi from northeastern Tennessee with means of 55.5 to 67.7 ova (Nagel 1980). The low fecundity observed in our specimens and in C. pygmaeus is correlated with their small size, while in the Tennessee population it appears to be correlated with a greater ova size. Females taken in 1978-80 were more fecund (Y = -209.87 + 8.13 SL) than in 1973-75 (Y = -142.24 + 6.46 SL). This difference is significant (0.05 level, testing the slope), and may be a function of size. However, although females in 1978-80 were larger (x SL = 38.2 mm) than in 1973-75 (xSL= 34.5 mm), the difference of the means is not significant. Nagel (1980) noted a similar difference in fecundity

Cottus Life History in Delaware

77

160-

140

120-

« 100

LU

CO

s

D

Z 80-

< > O

601-

40-

20-

10 20 30 40

STANDARD LENGTH (mm)

50

Fig. 3. Relationship between number of mature ova and standard length of Cottus bairdi from Butler Mill Branch, Sussex County, Delaware.

between two sampling periods, in which there also was no significant difference in size of adults.

Hann (1927) generalized that fecundity in C. bairdi is approximately proportional to the cube of the female's length. This is in agreement with relationship, Log F = -2.738 + 2.962 Log SL, observed by Ludwig and Norden (1969) in Wisconsin. Their fecundity-length relationship is higher than that observed by us for all Butler Mill Branch females examined: LogF= -2.129 + 2.608 Log SL.

The regression equation for the relationship of ova number (Y) to

78 Fred C. Rohde and Rudolf G. Arndt

body weight (W) is: Y = 49.19 + 31.43 W. The correlation coefficient of 0.77 is significant (P< 0.001), and indicates a positive relationship between ova number and gravid female weight.

Mature ova (large and orange) were not observed in 235 females until January. In 1974, mean diameter of ova from 14 preserved females increased from 1.18 mm in January to 2.07 mm just prior to spawning in March. The largest ovum measured 2.4 mm (early March). Values in 1975 (N - 23 females) fell within these ranges. In March 1980 (N = 20 females) the range was 1.8 to 2.5 mm (x = 2.15 mm). There was no difference in mean ovum diameter between ovaries (N = 22 females). Ludwig and Norden (1969) gave ova diameters of 1.50 to 2.06 mm (x = 1.88 mm) in Wisconsin, and Hann (1927) of 2 to 2.5 mm in Michigan. Nagel (1980) observed a larger average ovum size (3.32 mm) in Tennessee C. bairdi and suggested that it correlated with the smaller number of ova produced.

The relationship between diameter of mature ova and SL is signifi- cant (r = 0.55; P<0.01). This is in agreement with Hoar (1957) who stated that mature ova size depends both on size of parent and on nutri- tion during the pre-spawning period.

In seven females, the right ovary was usually longer (x = 9.5 mm) and narrower (x - 4.2 mm) than the left (x = 8.7 and 5.0 mm, respectively). The right contained a mean of 41.2 ova, the left 39.9. There is no signifi- cant chi-square difference from the expected 1:1 ratio (P>0.05).

A total of 2 egg clusters was found in 1980, and 37 in 1981. The larger number was due to additional egg attachment sites provided in February 1981, and to more extensive searching. In 1980, the first cluster was found on 15 March and the second on 22 March (deposited in the interim). In 198 1 , the first seven clusters were found on 27 February, five others on 7 March, and another six on 14 March. Additional clusters found on subsequent dates resulted (at least primarily) from searching in areas not examined earlier, rather than from later egg deposition. All clusters were found in and near two gravel riffles in the sampling area (Fig. 1), and to about 30 m upstream. All were in areas of fast current, usually near the stream center, and over gravel or gravel-sand substrate. Water depth by the eggs (on several dates in early 1980 and 1981) ranged from 3 to 50 cm (x = 20.0 cm, N = 40 measurements).

With one exception, every cluster was attached to the underside of a submerged object. The exception was in a hollow on the upper surface of a submerged log. One cluster was under a beer can and another under a bottle, and three others were under waterlogged wood: a branch, a piece of board, and a piece of cut log. Favored egg-attachment sites were pieces of concrete block and bricks, and 23 (58.9%) of the clusters were found on such objects. Ten clusters (25.6%) were on ironstone and silicified

Cottus Life History in Delaware 79

sandstone (both usually flat and concave beneath). Both occur naturally in this part of the Delmarva Peninsula, but the large local concentration was probably dumped there for old construction. Only two clusters (5.1%) were found on naturally-occurring material (log and branch). The general absence of rock or other suitable spawning sites in Delmarva Peninsula streams may limit C. bairdi distribution.

Four substrate samples from these two gravel riffles yielded particles that ranged in size from greater than 2.5 cm to smaller than 0.05 cm; about half the particles (by weight) were in the size category 0.8 to 2.5 cm (Table 2). By contrast, about half the particles (by weight) in three samples from three gravel patches in Horse Pen Branch were 0.05 cm or smaller (Table 2). The largest pieces of gravel collected in the sampling area riffles measured (maximum length and width, cm) 7.5 X 3.9, 6.1 X 3.5, and 5.9 X 4.2, while the largest from Horse Pen Branch were 2.2 X 1 .7, 2.3 X 1 .4, and 2.0 X 1 .4. Cottus bairdi was much more common over the riffles of larger gravel, perhaps because under natural conditions these facilitate spawning.

We consider 32 of the 39 clusters to represent the spawn of 1 female, 3 the spawn of 2 females, and 4 the spawn of 3 females. Ludwig and Norden (1969) noted a mean of 3.3 spawns per cluster. Koster (1936) observed up to six clusters in a nest, but noted that most nests contained from one to four egg masses. Clusters resulting from multiple spawnings can be identified by their larger size, varied egg mass colors, and irregular configuration. There were four instances of two single spawns found under one piece of substrate (each cluster was counted as the spawn of one female), and several cases of single- and multiple-spawn clusters on substrate with multiple-spawn clusters. Thus, the result of a total of 50 spawnings was found, deposited on a total of 31 pieces of substrate.

Color of live eggs was usually pink-orange, but the range in color of clusters was from orange to yellow to tan, and one cluster was almost white. About half of the clusters contained one to several white eggs, presumed to be unfertilized. Most clusters were approximately circular in outline, with eggs arranged in irregular rows, usually to four or five layers deep. They adhered strongly to the substrate and to each other. Egg cluster mean length, width, and height, respectively, and the range of each (in mm), for the total sample sizes given previously were: single spawn, 24.8 (16-37), 20.6 (13-29), 8.1 (6-12); two spawns, 40.0 (28-45), 24.3 (22-26), 8.6 (7-10); and three spawns, 48.2 (40-56), 30.0 (28-32), 1 1.7 (8-19).

Egg number in 1 1 single-spawn clusters ranged from 64 to 201, with an egg diameter range (measured to nearest 0.1 mm) of 2.2 to 3.5 mm, and a mean egg diameter per cluster range of 2.32 to 3.09 mm; egg number for one triple-spawn cluster was 623, with a range in egg diameter

80

Fred C. Rohde and Rudolf G. Arndt

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Cottus Life History in Delaware

81

of 2.5 to 3.4 mm, and a mean egg diameter of 2.87 mm (Table 3). Range in egg number was higher than that determined by examination of ova in females, as was the mean of 134.2 eggs (measured only for single-spawn clusters) versus 93.7 ova per female. These differences are not readily explained. Butler Mill Branch sculpin eggs are smaller in diameter than those found by Nagel (1980) in Tennessee (x - 3.73 mm). There was no relationship between egg diameter and date collected, or between egg diameter and cluster size.

Eggs preserved on the following dates were in the indicated stages of development: 27 February 1981, morula or earlier; 14 March 1981, morula and blastoderm; 15 March 1980, morula or earlier; 22 March 1981, morula or earlier ( 1 cluster), and with large, well-developed, curled embryos with large, black-pigmented eyes (3); 26 March 1981, with less well-developed young than the latter 22 March eggs, with large, gray eyes (1), and with well-developed young (1); and 29 March 1981, with well- developed young. Eggs in many clusters in the field on 22 March 1981 contained well-developed, active young. The number of eggs and clusters found with such advanced embryos increased on subsequent dates.

Eggs hatching in the field were first noted on 29 March 1981, when several empty egg shells were found in each of four clusters. Embryos in the remaining eggs were advanced and active. One cluster missing on this date may have totally hatched (rather than being destroyed by predators or high water). By 5 April 1981, all eggs in nine clusters had hatched, by

Table 3. Data on Cottus bairdi egg clusters collected in 1980 and 1981 at Butler Mill Branch, Sussex County, Delaware.

Date collected and preserved

N spawns/ N eggs/ cluster cluster

Range

egg diameter

(mm)

x egg

diameter

(mm)

N eggs measured

27 February 1981	170	2.6	- 3.4	2.86	20
27 February 1981	201	2.5	- 2.9	2.66	20
14 March 1981	117	2.2	- 2.5	2.32	20
15 March 1980	133	2.2	- 2.5	2.33	30
22 March 1980	64	2.3	- 2.4	2.35	20
22 March 1981	119	2.6	- 2.9	2.77	20
22 March 1981	170	2.8	- 3.1	2.94	20
22 March 1981	126	2.7	- 3.1	2.85	20
22 March 1981 ;	5 623	2.5	- 3.4	2.87	50
26 March 1981	126	2.3	- 2.7	2.51	20
26 March 1981	181	2.7	- 3.5	3.09	20
29 March 1981	70	2.4	- 2.7	2.56	20

82 Fred C. Rohde and Rudolf G. Arndt

1 1 April almost all clusters had totally hatched, and by 18 April all had hatched.

Relatively detailed data on hatching are available for two single- spawn clusters. On 7 March 1981, a pair of C. bairdi and eight eggs were found under a brick and we assumed the fish were spawning. No eggs were found here on 27 February. On 14 March, what we assumed to be the remainder (and much larger portion) of that cluster was found nearby on the brick. (Lifting of the brick on 7 March presumably interrupted spawning and caused its resumption at another point.) On 22 March, eggs of the group of eight contained well-developed embryos. On 26 March, well-developed young were still visible, with earlier-stage young in the larger portion. On 29 March, the eight eggs were missing and we presume they hatched. The incubation period was thus some 22 days. The remainder had not hatched by 5 April. The brick with eggs and guarding male were then placed in a plastic gallon jar with netting at each end to allow passage of a current of water, and replaced in the stream. On 1 1 April, most eggs and hatchlings had decayed, but three eggs were near hatching. The development period of these eggs was thus about 35 days. About half the eggs of another cluster found on 7 March 1981 (not present on 27 February) was hatched by 5 April, and the remainder contained well-developed embryos. On 1 1 April only 10 eggs remained, and on 18 April there was none. Time to hatching was some 29 to 42 days.

As in the field, eggs of a cluster also hatch over a period of days in the laboratory. A field-collected cluster maintained at approximately field water temperatures (10° to 12° C) hatched as follows: 7-8 April, 4; late 8 April, 59; 9 April, 14; evening of 9 April to mid-afternoon on 12 April, several; by mid-afternoon on 13 April, 4. Three eggs from another cluster hatched on 8 April 1981.

A male guards each cluster, and a male guarding a multiple-spawn cluster was presumably successful in spawning with up to three females. Koster (1936) stated that a successful male spawns with one or more females, and Scott and Crossman (1973) noted that it is usual for more than one female to deposit eggs in a male's nest. The same male stays with a given cluster until the last eggs hatch. He is usually hidden under the egg-deposition site, but sometimes lies with the head or body exposed. Rarely a guarding male was found with a second male, and once two males were hidden with a female heavy with eggs. These latter occurren- ces, however, may have been sampling artifacts.

Early in the 1981 reproductive season, from 22 February through 7 March, each male examined was in dark reproductive coloration. The dorsum and sides were olive-drab to blackish, the head and fins blackish, and the first dorsal fin edged with dull yellow and orange. The posterior

Cottus Life History in Delaware 83

venter was smoky-gray with fine black stippling, and the breast and abdomen dull whitish. The normally prominent black saddles were indis- tinct. On 14 March, seven of eight males were so colored, while one had the more typical tan background color with prominent chocolate-brown saddles. On each subsequent date, through 29 March 1981, some males had the dark coloration described above, but the number of such males then found per date never exceeded those with the brown body. Even some males guarding eggs on and after 14 March 1981 were brown.

Water temperature during the spawning period ranged from 7.5° to 12.5° C in March 1975, and from 6.5° to 11.0° C in March 1980. Temperatures from other years were within these ranges. Koster (1937) reported spawning in upper New York State at 10° C, Bailey (1952) in Montana at 7.8° to 12.8° C, Ludwig and Norden (1969) in Wisconsin at 8.9° to 13.9° C, and Nagel (1980) in Tennessee at 12° C and 14° C. Water temperature fluctuates considerably during egg development. Tempera- tures (°C) at Butler Mill Branch in 1981, from the date before the first eggs were found (22 February) through the date after the last were found (18 April), were: 10.0, 9.0, 7.5, 8.0, 7.0, 11.0, 11.5, 13.5, 12.0, and 16.0. Eggs in 1980 (two dates) were found at 8.0° C and 6.5° C.

The difference between number of males (69) and females (156) collected during the reproductive season was highly significant (chi- square, P<0.001, expected ratio 1:1). The observed values may be a sampling artifact. However, as a male is known to spawn with more than one female, it is also possible that the observed values reflect a real difference. Ludwig and Norden (1969) noted a reduction in number of males at commencement of the spawning season, and suggested it resulted from competition for nesting sites and some segregation of sexes. Koster (1936) found no significant difference in the proportion of males to females.

Age and Growth

The Butler Mill Branch sculpins are more short-lived than reported for any other Cottus species. This may be related to early onset of sexual maturity. Further, only few Butler Mill Branch C. bairdi that reach age group I survive to spawn twice. Length-frequency distributions based on the total sample through 1980 indicate the existence of three age groups: O, I, and a presumed II (Figs. 4,5). In 1974, group I fish were present from January to September, but none was collected in November. Of later samples, some group I fish were still present in November 1978. A few may have survived the winter, as indicated by the large specimens Q> 50 mm SL) collected in March and July 1979, which would then have been in age group II. Fish corresponding to this age group were also present in the 1980 samples. Of the 403 sculpins collected in 1980, 262 (65.0%) were

84

Fred C. Rohde and Rudolf G. Arndt

10

20

15-

10-

5-

UJ 10

CO

i 5

20

15

10

5-

T

ikOa..

20

*

30 40

STANDARD LENGTH (mm)

2,23 MAR. 1975 N 65

30 NOV. 1974 N 122

8 SEPT. 1974 N 101

20 JULY 1974 N 94

17 MAR. 1974 N 5

17 FEB. 1974 N 5

20JAN. 1974 N 26

28 OCT., 4 NOV. 1973 N 17

Fig. 4. Length frequencies of Cottus bairdi from Butler Mill Branch, Sussex County, Delaware, October 1973-March 1975. Solid bars = males; open bars - females; hatched bars = unsexed young.

Coitus Life History in Delaware

85

15- 10-

10- i 5-

d EL.

tHbi--

cLAm. ..

30 40 50

STANDARD LENGTH (mm)

9 AUG. 1980 N 55

25 JULY 1980 N 68

12 JULY 1980 N 77

28 JUNE 1980 N 61

8 JUNE 1980 N 35

4,20 MAY 1980 N 17

9,15,22,30 MAR. 1980 N 90

25 JULY 1979 N 67

16 MAR. 1979

N 17 26 NOV. 1978

N 6

7 SEPT. 1978 N 46

Fig. 5.

County

females:

Length frequencies of Cottus bairdi from Butler Mill Branch, Sussex Delaware, September 1978-August 1980. Solid bars = males; open bars = hatched bars = unsexed young.

86 Fred C. Rohde and Rudolf G. Arndt

in age group O at time of capture, 133 (33.0%) in group I, and 8 (2.0%) in presumed group II. Although the data are limited, and assuming that each age group was collected proportional to its actual number in the population, 33.0% of group 0 survived to spawn as group I fish, and 2.0% to spawn a second time. Survival rates have not been calculated for other populations of any species of Cottus.

Three just-hatched young (measured to nearest 0.1 mm) from one cluster averaged 6.56 mm SL, and 24 from a second cluster averaged 6.05 mm. Hann (1927) observed a mean size at hatching of 6.4 mm SL. Newly hatched young in other studies ranged from 5.6 mm SL to 9.8 mm TL (see Nagel 1980). The body of a newly hatched young is slightly opaque, with no evidence of the dark saddles, and with a prominent yolk sac. Five siblings of 98 hours of the second cluster averaged 6.54 mm SL. The yolk sac of these is greatly reduced and only slightly protrudes from the body, and melanophores form weakly-defined saddles. Five siblings of 195 hours averaged 6.96 mm SL. Their yolk sac appears fully resorbed, melanophores are conspicuous on the body, and the saddles are better defined. Three additional siblings of 288 hours averaged 6.63 mm SL.

Two young-of-year taken on 19 May 1981 were 10 mm SL and 12 mm SL, and resembled the adult in morphology and coloration. No young were taken until 8 June in 1980 (Table 4, Fig. 5), at which time they averaged 13.5 mm SL (range 1 1-16 mm), an assumed post-hatching increase of 7.5 mm. Subsequent samples taken approximately biweekly revealed a decrease in growth rate, from a 4.9 mm increase over 20 days to 28 June, to but a 2.4 mm increase in the 15 days to 9 August.

Most growth occurred in the first calendar year of life. Young taken in July 1974 ranged from 16 to 26 mm SL (x= 21.1 mm). From hatching at 6 mm in early to mid-April, growth in the 1974 year class was rapid: to 21.1 mm in July, 24.2 mm in September, and 30.0 mm in November

Table 4. Monthly mean standard lengths, by sex, for 1978, 1979, and 1980 year classes of Cottus bairdi from Butler Mill Branch, Sussex County, Delaware. Number of specimens in parentheses.

1978 Year Class 1979 Year Class 1980 Year Class

Date Unsexed Male Female Unsexed Male Female Unsexed

37.8(10)

44.0 (8) 23.1 (56)

59 (1) 59.5 (2)

7 Sept. 1978	30.9(19)	-
16 Mar. 1979	-	49.9 (7)
25 Jul. 1979	-	43.3 (3)
9, 15,22,30,
Mar. 1980	-	59.4 (4)
4, 20 May 1980	-	-
8 Jun. 1980	-	61 (1)
28 Jun. 1980	-	-
12 Jul. 1980	-	-
25 Jul. 1980	-	-
9 Aug. 1980	-	-

45.0 (24)	38.2(62)	-
47.3 (3)	41.4(13)	-
47.0 (3)	42.8 (8)	13.5(23)
45 (1)	41.3 (7)	18.4(51)
47.0 (4)	41.5 (4)	22.2 (69)
50 (1)	43 (1)	24.5 (66)
51 (1)	45 (1)	26.9(53)

Cottus Life History in Delaware 87

Table 5. Monthly mean standard lengths, by sex, for 1973 and 1974 year classes of Cottus bairdi from Butler Mill Branch, Sussex County, Delaware. Number of specimens in parentheses.

Date	1973 Year Class	1974 Year Class
Male	Female	Unsexed	Male	Female
28 Oct.,
4 Nov. 1973	32.5 (6)	27.7(10)
20 Jan. 1974	36.6 (5)	33.8(21)
17 Feb. 1974	41.5(2)	38.7 (3)
17 Mar. 1974	41.0(3)	38.0 (2)
20 Jul. 1974	46.7 (3)	41.5 (8)	21.1 (83)	-	-
8 Sep. 1974	43.5 (6)	40.5(13)	24.2 (82)	-	-
30 Nov. 1974	-	41 (1)	30.0(121)	-	-
2 Mar. 1975	-	-	-	30.7(18)	29.8 (37)
23 Mar. 1975	-	-	-	35.0 (3)	30.4 (7)

(Table 5). By next March these fish, now in age group I, were mature and had spawned. Males (N = 21) now averaged 31.3 mm SL and females (N = 44) 29.9 mm.

Growth in the 1973 year class was at first similar, with a combined October-November sample mean length of 29.5 mm SL. However, growth did not decrease pronouncedly or cease in winter as noted in the 1974 year class and as reported for a population by Bailey (1952). Rather, sculpins of the 1973 year class apparently continued to grow from November to March. For combined February-March 1974 samples, both sexes (5 males, 5 females) averaged some 10 mm longer than those taken in early March 1975. The reason for this difference is not known; it may be an artifact of small sample size. Nagel (1980) also noted such between- year variation in growth rate. The 1979 year class (age group I) exhibited little growth, if any, from June through August (Table 4). A few fish presumed of the 1978 year class (age group II) were also present in the 1980 samples. These were 10 to 15 mm longer than those collected the previous spring, interpreted as growth continuing during the second year of life.

Cottus bairdi in Montana exhibited a growth rate comparable to Butler Mill Branch specimens during age group 0, but none matured until in age group II and when usually longer than 74 mm TL (Bailey 1952). Ludwig and Norden (1969) observed growth similar to Sussex County Cottus in age groups 0 to II, but reported earliest maturity in Wisconsin sculpins in age group II (x SL = 54.5 mm, range 37-70). Nagel (1980) noted reproduction in Tennessee at the end of the second year of life, at

88 Fred C. Rohde and Rudolf G. Arndt

50-81 mm TL. Hann (1927) stated that sexual maturity in Michigan is attained by two years of age, at 45-70 mm SL. Koster (1936) found that first spawning in a stream-dwelling population near Ithaca, New York, occurred at the end of the second year of life (age group II), while some specimens of a lake-dwelling population from nearby Cayuga Lake apparently matured at the end of the first year of life (age group I). This early maturity may result from a genetic or an environmental difference between these populations.

Butler Mill Branch sculpins reached sexual maturity at a smaller size than any other population of C. bairdi (x SL = 37.4 mm, range 22-64, N = 225). Mean length of 33 Butler Mill Branch adults taken in 1981 was 44.7 mm SL (range 35-65 mm). Adults from the new localities in Caroline and Dorchester counties, Maryland, were of similar size (x SL = 38.7 mm, range 29-62, N = 48), and Franz and Lee (1976) noted small size in the other Caroline County population.

Mature males in the March and May 1978-80 samples were larger (x SL = 47.6 mm, range 37-64, N = 38) than mature females (x SL = 38.9 mm, range 32-59, N = 86), a highly significant difference (P<0.001). However, there was no significant difference at the 0.05 level in the January, February, and March 1973-75 samples (mature males, x SL = 33.7 mm, range 25-47, N = 31; mature females, x SL = 31.7 mm, range 22-40, N = 70). The reason for the size difference of 1978-80 is not appar- ent. Such sexual dimorphism was also observed by Hann (1927), Bailey (1952), Ludwig and Norden (1969), and Nagel (1980), but not in New York sculpins by Koster (1937).

Regressions for the weight-length relationship derived for males and females from both sampling periods are: (1973-75) males, Log W = - 10.8985 + 3.0585 Log L, r = 0.99; females, Log W = -10.3999 + 2.8814 Log L, r = 0.99; (1978-80) males, Log W = -1 1.8313 + 3.3681 Log L, r = 0.99; and females, Log W = -1 1.0505 + 3.1412 Log L, r= 0.99. Analysis of covariance reveals significant differences (P<0.05) in slope between sexes and between years. A slope greater than 3.00, indicating that relative weight increased faster than length (Ricker 1971), is noted in three of four regressions. Only females in 1973-75 showed a slightly faster increase in length when compared to weight.

There is a strong, and expected, positive correlation between SL and TL: SL= 1.21 + 0.76 TL, r= 0.93.

Condition factor (K) for both sexes is lowest during the coldwater months preceding spawning (Table 6). Highest K values were recorded during the warmwater period subsequent to spawning. With the excep- tions of October and November, males were more robust than females.

Cottus Life History in Delaware

89

Table 6. Combined monthly averages of coefficient of condition (K) for male and female Cottus bairdi from Butler Mill Branch, Sussex County, Delaware, 1973-1980. Male N = 145, female N = 278.

Month

	Male
Gonads	Gonads
in situ	excised
2.06	2.06
2.10	2.08
2.88	2.84
3.07	3.07
2.88	2.88
2.53	2.70
3.33	3.33
2.36	2.26
2.19	2.17
2.34	2.32

Female Gonads Gonads

in situ excised

Jan.

Feb.

Mar.

May

Jun.

Jul.

Aug.

Sep.

Oct.

Nov.

2.06	1.93
2.44	1.96
2.58	2.12
2.63	2.62
2.67	2.65
2.27	2.26
2.68	2.66
2.14	2.06
2.20	2.17
2.39	2.34

Food Habits

Based on stomach contents of 539 fish ( 1 2-64 mm SL), food is prim- arily larval trichopterans (in 55.7% of stomachs) and dipterans (42.3%), and plecopteran nymphs (19.7%); less important are crustaceans (9.2%), ephemeropteran nymphs (8.0%), and coleopteran larvae (2.4%) (Table 7). Nineteen taxa were identified. One hundred and twenty-eight (23.7%) stomachs were empty. Franz and Lee (1976) found 5 taxa in 10 speci- mens. Ricker (1934), Koster (1936, 1937), Dineen (1951), Bailey (1952), and Daiber (1956) also found that the primary food was benthic insect larvae.

Seasonal variation in diet is evident (Table 7). Trichopterans, mainly Hydropsychidae, were eaten in all months except February, and were dominant in March, July, August, and September. Dipterans, primarily Chironomidae, were the most common food in January, June, October, and November. Plecopterans were common from January through May.

Prey taxa differed only slightly by fish size group (Table 8). Chiro- nomids were relatively more important to sculpins smaller than 19 mm SL than to longer fish. Larger specimens (>49 mm SL) ate fewer chiro- nomids but more larger simulids. A 43 mm SL sculpin contained one 25 mm SL Etheostoma olmstedi, and a 53 mm SL specimen had eaten an egg of Lampetra aepyptera.

90

Fred C. Rohde and Rudolf G. Arndt

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Cottus Life History in Delaware

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92 Fred C. Rohde and Rudolf G. Arndt

Table 8. Mean number of food items, by length groups, in stomachs of Cottus bairdi from Butler Mill Branch, Sussex County, Delaware, 1973-1980.

Food

Cottus SL (mm) <20 20-29 30-39 40-49 >49

	.01	.04
	.01	.01
.15	.04	.16
.01
.01	.36	.53
	.03	.04
.13	.15	.06
.04	.10	.06
.02	.02
.02	.02	.01

Nematoda

Oligochaeta

Crustacea

Amphipoda .22 .15 .04 .16 .08

Isopoda Insecta Plecoptera

Perlodidae

Chloroperlidae

Unidentified Ephemeroptera

Heptageniidae .06 .04 .06

Leptophlebiidae

Unidentified Odonata

Anisoptera .01

Zygoptera .01

Trichoptera

Hydropsychidae 1.17

Phryganeidae

Unidentified Coleoptera

Elmidae

Gyrinidae

Unidentified Diptera

Chironomidae 7.89

Simuliidae

Tipulidae

Unidentified .01

Osteichthyes

Lampetra aepyptera .08

Etheostoma olmstedi .0 1

Number of stomachs with countable food items 18 122 152 68 13

Abundance

Throughout most of this study we assumed that the Butler Mill Branch sculpin population was small. Work in early 1980, however, revealed the species to be more widely distributed here than previously

.89	.94	1.16	1.23
.01
.09	.19	.04
.01	.06	.01
.01	.01
1.61	2.51	.28	.23
.10	.05 .03	.19	.46

Cottus Life History in Delaware 93

known, and in March both frequency of sampling and sample size were increased. Surprisingly, no effect of increased collecting on the popula- tion was noted. Although on each collection date the sampling area was almost cleared of specimens, and no additional sculpin could be found, on each subsequent visit the area again was productive. Thus, it appears that many fish escaped capture, or that there was extensive and rapid movement into the sampling area from nearby. Probably both were important. Certainly larval C. bairdi are secretive, which probably explains our failure to catch them at Butler Mill Branch. Hatchlings in the laboratory are benthic to at least 288 hours. When disturbed they are strong swimmers, but then settle rapidly and remain quietly in gravel. This larva-gravel association may explain why C. bairdi is restricted to areas of gravel.

Further, lower Butler Mill Branch, including all of the sampling area, was poisoned with rotenone in May 1976 by Delaware Division of Fish and Wildlife personnel to foster game fish in Craigs Pond (Roy W. Miller, pers. comm.). Sampling for sculpins in 1978 revealed no effect of the rotenoning. We conclude that the population of C. bairdi here is healthy, and the adjacent Maryland populations also appear to be strong.

ACKNOWLEDGMENTS.— We thank the following individuals for their help. Johnson C.S. Wang, Nancy J. Bieber, Ruth A. Hermansen, Judith K. Heyman, Michael J. Hozik, Joan A. Pienta, and several Stock- ton State College students assisted in collecting. J. C.S. Wang also exam- ined sculpin eggs, and Michael J. Hozik provided information on geol- ogy. Robert H. Simmons, U.S. D.I. Geological Survey, Water Resources Division, Dover, Delaware, provided stream flow data. Roy W. Miller, Supervisor of Finfisheries, Delaware Division of Fish and Game, Dover, supplied information on rotenoning in Butler Mill Branch. William L. Pflieger, Missouri Department of Conservation, Columbia, provided information on the Ozark populations of C. bairdi. Robbin N. Rogers searched samples for fish eggs and larvae. Bruce W. Menzel and Steve W. Ross made numerous helpful comments during manuscript preparation. Nancy J. Bieber and Deborah K. Wozniak typed drafts of the manus- cript. John E. Cooper and an anonymous reviewer provided helpful comments during manuscript review.

LITERATURE CITED Bailey, Jack E. 1952. Life history and ecology of the sculpin Cottus bairdi

punctulatus in southwestern Montana. Copeia 1952(4):243-255. Daiber, Franklin C. 1956. A comparative analysis of the winter feeding habits of

two benthic stream fishes. Copeia 1956 (3): 141-151. Dineen, Clarence F. 1951. A comparative study of the food habits of Cottus

bairdi and associated species of Salmonidae. Am. Midi. Nat. 46(3): 640-645.

94 Fred C. Rohde and Rudolf G. Arndt

Foltz, Jeffrey W. 1976. Fecundity of the slimy sculpin, Cottus cognatus, in Lake Michigan. Copeia 1976(^:802-804.

Franz, Richard, and D.S. Lee. 1976. A relict population of the mottled sculpin, Cottus bairdi, from the Maryland Coastal Plain. Chesapeake Sci. /7(4):301-302.

Gage, Simon H. 1878. Notes on the Cayuga Lake star gazer. Cornell Rev. 6(2):91-94.

Hann, Harry W. 1927. The life history of the germ cells of Cottus bairdi Girard. J. Morphol. Physiol. 43(2):427-497.

Hoar, W.S. 1957. The gonads and reproduction, pp. 287-317 in M.E. Brown (ed.). The Physiology of Fishes, Vol. I. Academic Press, Inc., New York. 465 pp.

Koster, William J. 1936. The life-history and ecology of the sculpins (Cottidae) of central New York. Unpubl. Ph.D. dissert., Cornell Univ., Ithaca. 87 pp. 4- Appendix and figures.

. 1937. The food of sculpins (Cottidae) in central New York. Trans.

Am. Fish. Soc. (56:374-382.

Lee, David S. 1980. Cottus bairdi Girard. Mottled sculpin. p. 805 in D.S. Lee et al. Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. Hist., Raleigh, x + 867 pp.

Ludwig, Gerald M., and C.R. Norden. 1969. Age, growth and reproduction of the northern mottled sculpin (Cottus b. bairdi) in Mt. Vernon Creek, Wis- consin. Milw. Public Mus. Occas. Pap. Nat. Hist. 2:1-67.

Nagel, Jerry W. 1980. Life history of the mottled sculpin, Cottus bairdi, in north- eastern Tennessee (Osteichthyes: Cottidae). Brimleyana 4:1 15-121.

Patten, Benjamin G. 1971. Spawning and fecundity of seven species of north- west American Cottus. Am. Midi. Nat. #5(2):493-506.

Petrosky, Charles E., and T.F. Waters. 1975. Annual production by the slimy sculpin population in a small Minnesota trout stream. Trans. Am. Fish. Soc. 104(2):237-244.

Pflieger, William L. 1975. The Fishes of Missouri. Mo. Dep. Conserv., Jefferson City. 343 pp.

Ricker, William E. 1934. An ecological classification of certain Ontario streams. Univ. Toronto Stud. Biol. Ser. No. 37. 1 14 pp.

. 1971. Methods for assessment of fish production in fresh waters. 2nd

ed. IBP Handbook No. 3. Blackwell Sci. Publ., Oxford. 348 pp.

Robins, C. Richard. 1954. A taxonomic revision of the Cottus bairdi and Cottus carolinae species groups in eastern North America (Pisces, Cotti- dae). Unpubl. Ph.D. dissert., Cornell Univ., Ithaca. 248 pp.

Scott, W.B., and E.J. Crossman. 1973. Freshwater Fishes of Canada. Fish. Res. Board Can. Bull. 184. 966 pp.

Smith, Bertram G. 1922. Notes on the nesting habits of Cottus. Pap. Mich. Acad. Sci. Arts Lett. 2:222-224.

Williams, James D. 1968. A new species of sculpin, Cottus pygmaeus, from a spring in the Alabama River Basin. Copeia 1968(2):334-342.

Accepted 28 December 1981

Distribution and Ecology of the Seepage Salamander

Desmognathus aeneus Brown and Bishop (Amphibia: Plethodontidae),

in Tennessee

R. L. Jones '

Graduate Program in Ecology, University of Tennessee, Knoxville, Tennessee 37916

ABSTRACT. — The seepage salamander, Desmognathus aeneus, is found at elevations from 280 to 1000 m in the Unicoi Mountains of Polk and Monroe counties, southeastern Tennessee. It inhabits leaf litter along small streams and seepage areas. Oviposition probabh occurs in late April and early May, and hatching from mid-June through mid-July. Clutch sizes range from 8 to 15 (x - 12.2) eggs. The major prey items for Tennessee D. aeneus appear to be mites ana collembolans.

INTRODUCTION

Two dwarf species of salamanders in the genus Desmognathus occur in Tennessee. The pygmy salamander, Desmognathus wrighti King, is found primarily at higher elevations along the main ridge of the Unaka Mountains (Great Smokies, Bald Mountains, and Roan Moun- tain) on the Tennessee-North Carolina border. The seepage salamander, Desmognathus aeneus Brown and Bishop, has been reported from one locality in the extreme southeastern corner of the state (Harrison 1967). I here present additional information on the distribution, status, and ecology of D. aeneus in Tennessee. Most of my data were obtained during a 1976 study of habitat use among plethodontid salamanders in the Unicoi Mountains (Jones 1977). Supplemental information was obtained during later collecting trips and from specimens in the Verte- brate Zoology Collection, University of Tennessee, Knoxville.

RESULTS

Distribution

Harrison (1967) reported D. aeneus from Turtletown, Polk County. Additional localities are shown in Figure 1. The species seems to occur in isolated populations throughout the Unicoi Mountains in both Polk and Monroe counties. All localities are within the Blue Ridge Physiographic Province and range from 280 to 1000 m elevation. Harri-

1 Present address: Mississippi Museum of Nature Science, Jackson, MS 39202 Brimleyana No. 7:95-100. July 1981. 95

96

R. L. Jones

Georg i a

Fig. 1. Distribution of D. aeneus in Tennessee. Solid symbols represent localities from which specimens were examined, open symbol represents a literature record, dashed line represents approximate western boundary of Blue Ridge Physiographic Province.

son (1967) felt that the Little Tennessee River acted as a barrier to the dispersal of this species. My efforts to locate populations in the area immediately north of this river in Tennessee have been unsuccessful.

Habitat

Desmognathus aeneus has been characterized as an inhabitant of seepages and of the leaf litter near small streams in wooded areas (Fol- kerts 1968). I found that the mean distance from nearest open water was 7.8 m (range 0.3-30.1 m) for 55 D. aeneus from the Citico Creek watershed, Monroe County, during summer 1976. Sixty percent of these were found within 5 m of streams and seepage areas (Table 1). How- ever, specimens also can be found a considerable distance into the sur-

Desmognathus aeneus in Tennessee 97

Table 1. Values of 3 habitat variables for 55 D. aeneus in the Citico Creek watershed, Monroe Co., Tennessee. Values in parentheses represent percentages of salamanders found in each habitat category.

Cover	type	Substrate type		Distance from water (meters)
Leaf litter	22(40.0)	Leaf-soil interface	22(40.0)	0-5 33(60.0)
Moss	16(29.1)	Rock-soil interface	2( 3.6)	6-10 10(18.2)
Log	11(20.0)	Leaf litter	6(10.9)	11-20 8(14.5)
Rock	6(10.9)	Log Soil Rock	9(16.4) 10(18.2) 6(10.9)	>20 4( 7.3)

rounding woodland, since almost 22 percent occurred farther than 10 m from open water. Hairston (1973) also found D. aeneus in relatively terrestrial situations in the Nantahala Mountains of North Carolina. Both mosses and small logs were frequently used as cover objects, but I found most salamanders beneath leaf litter at the leaf-soil interface (Table 1). Canopy cover at the point of capture for the 55 specimens ranged from 60 to 90 percent (x = 78.4%). In the Citico Creek watershed, the species seems to especially favor areas with heavy growths of Rho- dodendron maxium. The habitat requirements for D. aeneus in this area are most similar to those of Eurycea bislineata (unpubl. data), although D. aeneus also shares its habitat to some extent with Desmognathus ochrophaeus Cope, D. fuscus (Rafinesque), D. monticola Dunn, and Plethodon glutinosus (Green).

Diet

Donovan and Folkerts (1972) reported that the stomach contents of D. aeneus from Alabama and Georgia consisted primarily of arthro- pods. Insects (collembolan and dipteran larvae) and arachnids (primar- ily mites) were the two most frequently ingested prey types. I also found arthropods to be the most common prey items in the stomachs of 47 D. aeneus from the Citico Creek watershed, Monroe County (Table 2). This group comprised slightly over 95 percent of the prey items ingested and was found in almost 96 percent of the stomachs. The most fre- quently ingested arthropods were mites, representing about 58 percent of the total prey items and found in almost 79 percent of the stomachs. Insects, primarily collembolans, were found in almost 62 percent of the stomachs but made up only 30 percent of all prey items ingested. The composition of prey found in the stomachs suggests that D. aeneus does

98

R. L. Jones

Table 2. Stomach contents of 47 D. aeneus from the Citico Creek watershed, Monroe Co., Tennessee.

Category	N prey	% freq.	N stomachs	% freq.
Nematoda	2	0.8	2	4.2
Mollusca	10	4.1	8	17.0
Arthropoda	233	95.1	45	95.7
Crustacea (Isopoda)	1	0.4	1	2.1
Insecta	71	30.0	29	61.7
Collembola	40	16.3	17	36.2
Diptera	7	2.9	6	12.8
Coleoptera	5	2.0	5	10.6
Hymemoptera	4	1.6	4	8.5
Lepidoptera	2	0.8	2	4.2
Hemiptera	2	0.8	2	4.2
Insect larvae	11	4.5	9	19.1
Chilopoda	1	0.4	1	2.1
Diplopoda	3	1.2	2	4.2
Arachnida	154	62.9	38	80.8
Araneae	3	1.2	3	6.4
Pseudoscorpionida	8	3.3	6	12.8
Acarina	143	58.4	37	78.7
Unidentified arthropods	3	1.2	3	6.4

most of its foraging within the leaf litter rather than at the surface. This idea is further supported by approximately 72 hours of monitoring noc- turnal activity of salamanders during both wet and dry nights in the Citico Creek watershed in 1976; no D. aeneus were observed active on the surface in either seepage areas of leaf litter along small streams, although they were known to be abundant in the immediate area. I have found the species crossing roads during rainstorms, but it appears to rarely leave the protection of surface cover in its habitat, an observation first noted by Folkerts (1968) for D. aeneus in Alabama.

Reproduction

Folkerts (1968) found evidence for both spring and fall periods of oviposition in Alabama D. aeneus. He suggested that the species may deposit eggs at any time of year except mid-winter. Harrison (1967) reported egg deposition from late April to early May, but suggested that considerable variation probably occurred in the time of oviposition

Desmognathus aeneus in Tennessee 99

from year to year. Folkerts (1968) found that clutch size varied from 5 to 17 (x = 8.8) and that hatching in the laboratory occurred in 43 to 54 days. Harrison (1967) observed hatching from late May to early August, and reported that hatching in the laboratory occurred in 34 to 45 days. He found that clutch size varied from 6 to 18 (x = 10.7). Eleven D. aeneus clutches have been found in the Citico Creek watershed. The earliest date for a clutch containing uncleaved eggs was 10 May. Clutches that were ready to hatch or hatching were observed on 23 June and 26 June, respectively. This indicates that, in Tennessee, oviposition probably occurs from late April through early May and hatching from mid-June through mid-July. I found no fall clutches. Clutch size ranged from 8 to 15 (x = 12.2). Most of the 11 clutches were beneath moss at ground level near streams or seepage areas. However, a female attend- ing a clutch was found beneath a small log, 1.6 m from a small stream; a second female and clutch were beneath moss on a rotting tree stump, 28 cm above ground level and 7.2 m from a stream.

DISCUSSION

The range of D. aeneus in Tennessee is apparently restricted to the Unicoi Mountains of Polk and Monroe counties. Tennessee populations are probably continuous with those in western North Carolina and northern Georgia, since D. aeneus has been collected in counties border- ing Tennessee in both states (Martof, et al. 1980; Harrison 1967).

Within the Unicoi Mountains, D. aeneus is an apparently wides- pread but relatively uncommon species. Its range in Tennessee lies primarily within the Cherokee National Forest, which should provide some measure of protection since much of this area remains relatively undisturbed. However, populations of D. aeneus are sensitive to drying conditions caused by clear cutting (Folkerts 1968), a common method of timber harvest in the Cherokee National Forest. Disruption by wild boar, Sus scrofa, of seepage area habitats is also a relatively common phenomenon within the range of D. aeneus in Tennessee. The apparent susceptibility of its populations to habitat destruction, coupled with its restricted distribution in the state, indicate that D. aeneus should be monitored for any sudden population changes in order to insure its con- tinued survival in Tennessee.

ACKNOWLEDGMENTS.— A. C. Echternacht, University of Ten- nessee, and W. H. Redmond, Jr., Tennessee Valley Authority, provided many useful comments on earlier versions of this manuscript. Two anon- ymous reviewers also provided comments.

100 R. L. Jones

LITERATURE CITED

Donovan, Lois A., and G. W. Folkerts. 1972. Foods of the seepage salamander Desmognathus aeneus Brown and Bishop. Herpetologica 2#(l):35-37.

Folkerts, George W. 1968. The genus Desmognathus Baird (Amphibia: Ple- thodontidae) in Alabama. Unpubl. Ph.D. dissert., Auburn Univ., Auburn. 129 pp.

Hairston, Nelson G. 1973. Ecology, selection and systematics. Breviora 414:1-21.

Harrison, Julian R. 1967. Observations on the life history, ecology and distri- bution of Desmognathus aeneus aeneus Brown and Bishop. Am. Midi. Nat. 77(2):356-370.

Jones, R. L. 1977. Resource partitioning among six species of the salamander family Plethodontidae. Unpubl. MS thesis, Univ. Tennessee, Knoxville. 59 pp.

Martof, Bernard S., W. M. Palmer, J. R. Bailey and J. R. Harrison III. 1980. Amphibians and Reptiles of the Carolinas and Virginia. Univ. North Caro- lina Press, Chapel Hill. 264 pp.

Accepted 11 January 1982

Dental and Cranial Anomalies in the River Otter (Carnivora: Mustelidae)

Thomas D. Beaver, George A. Feldhamer and Joseph A. Chapman

Appalachian Environmental Laboratory,

Gunter Hall, Frostburg State College Campus,

Frostburg, Maryland 21532

ABSTRACT.— Dental or cranial anomalies were noted in 65 (32.2%) of 202 skulls of river otters, Lutra canadensis, collected from the Ches- apeake Bay region of Maryland and Virginia from 1974 to 1979. The most frequent anomaly was alveolar thinning. Anomalies probably did not adversely affect individuals or the population structure.

INTRODUCTION

The dentition and cranial structure of many mammalian species have been well studied because of their importance in systematics and taxonomy. Also, the condition of teeth affects nutritional status, which in turn directly affects behavior, reproduction and longevity (Robinson 1979). As a result, anomalies have been described for several orders and numerous mammalian species (Choate 1968; Colyer 1936; Hershkovitz 1970; LaVelle and Moore 1972; Manville 1963; Pavlinov 1975; Sheppe 1964; Shultz 1923). However, relatively few studies have dealt with the Mustelidae (Hall 1940; Heran 1970; Parmalee and Bogan 1977). Here we describe dental and cranial anomalies noted in skulls of river otters, Lutra canadensis, from the Chesapeake Bay region of Maryland and Virginia and assess their probable impact upon individuals and the population.

MATERIALS AND METHODS

A total of 202 skulls was collected from trappers in Dorchester County, Maryland, during the trapping seasons of 1974 through 1979. The material was considered a random sample representative of the river otter population as no skull was obtained specifically because it had an anomaly. All skulls were individually numbered and are housed in the museum collection of the Appalachian Environmental Labora- tory (AEL). Each skull was examined for the following: 1) plagio- cephaly — asymmetrical cranial growth due to premature closure of one frontal-parietal suture; 2) bregmatic bones — extra bones derived from accessory ossification in any of the fontanelles; 3) heterotopic bones — small accessory bones; 4) caries — decay of dental tissue; 5) alveolar thinning — exposure of the buccal tooth roots; 6) supernumerary teeth — those in excess of the normal dental pattern; 7) congenital agenesis

Brimleyana No. 7:101-109. July 1981. 101

102 Thomas D. Beaver, George A. Feldhamer, Joseph A. Chapman

— reduced dental complement due to teeth that failed to develop; and 8) irregular placement — teeth in positions other than the normal pat- tern. Skulls were X-rayed to provide confirmation of suspected agenesis or supernumeration.

RESULTS AND DISCUSSION

A total of 137 skulls (67.8%) exhibited no cranial abnormalities or deviation from the normal dental pattern of 3/3, 1/1,4/3, 1/2 = 36 in L. canadensis. Anomalies were found in 65 skulls (32.2%). This is similar to the results of Colyer (1936) who noted "marked irregularities" in 75 of 161 (46.6%) skulls of Lutra sp. — a greater percentage of occurrence of anomalies than in any other genus of Mustelidae he examined.

Cranial anomalies. — Cranial anomalies occurred in only 2 (1.0%) of the skulls examined. Plagiocephaly was found in an adult female over 3 years of age, and there was a small hole in the frontal bone (Fig. 1). Mowbray et al. (1979) indicated that 1 1% of the 296 river otter carcasses that they examined from this study area showed evidence of gunshot wounds, although few animals (1.7%) were judged to have died from such wounds. Dougherty and Hall (1955) and van Soest et al. (1972) noted that characteristics similar to those seen in Figure 1 may result from metastrongylid nematode (Skrjabingylus sp.) involvement. Thus, the plagiocephaly may have resulted from an extrinsic agent, either a nonlethal gunshot wound or a parasite, rather than genetic factors. Heterotopic bones occurred on the premaxilla of an adult male over 3 years of age (Fig. 2); there were no indications of previous injury. Whether this condition was genetic in origin or resulted from previous trauma could not be determined. No instances were found of bregmatic bones.

Dental anomalies. — Dental anomalies were much more prevalent than cranial anomalies. With the exception of caries, an example of each dental anomaly was represented in the sample. The lack of caries in wild animals is consistent with previous studies. Hall (1940) found caries in only 8 of 3,761 specimens (0.2%) of North American carniv- ores, and all cases occurred within the Ursidae. Colyer (1936) reported only 4 cases of caries in 7,635 specimens (0.05%) of North American carnivores. The reason for lack of caries in the river otter, as well as most other carnivores, is unknown, although it may be associated with absence of carbohydrates in the diet.

Irregular tooth placement was noted in two individuals: a juvenile female had teeth that were abnormally far apart (Fig. 3), and in an adult female they were exceedingly close together (Fig. 4).

Alveolar thinning (Fig. 5) was the most common anomaly, occurr ing in 47 skulls (23.3%). Of these, 39 (82.9%) involved the last upper

River Otter Skull Anomalies

103

Fig. 1. Plagiocephaly in adult female river otter (AEL-302); also note small hole in frontal bone.

Fig. 2. Heterotopic bones in adult male river otter (AEL-303).

104 Thomas D. Beaver, George A. Feldhamer, Joseph A. Chapman

molar, and 7 others involved both upper and lower premolars in which thinning was associated with the posterior premolar in the tooth row. Smith et al. (1977) also found alveolar thinning associated "almost exclusively" with the last upper molars in three species of platyrrhine monkeys. They attributed the condition to internal pressures associated with mastication. Such thinning may predispose underlaying tissues to periodontal disease, and both local and systemic factors have been implicated in this process (Clark et al. 1970).

Congenital agenesis occurred in 8 (3.9%) of the skulls. Three addi- tional skulls appeared to possess this condition, however X-ray roent- genograms revealed a non-erupted tooth. Agenesis was bilateral in two individuals; one case involved the second lower premolars and the other the second upper premolars (Fig. 6). The other six cases, all unilateral, involved two different locations in the maxillary tooth row: five instances occurred at the first premolar and one at the second premolar.

Agenesis can result from delayed tooth formation and eruption, or a genetically induced reduction associated with phylogenetic shortening of the tooth row (LaVelle and Moore 1972). Delayed eruption was noted in three instances; however, this was not the case for the nine individuals in which no unerupted tooth or alveolus was present. In all cases congenital agenesis involved the premolars. Hall (1940:1 18) stated, "when a premolar is missing the place most often is at the anterior end of the premolar series." This was the situation in only 5 of 9 individuals (55.6%) in our sample; the remainder involved the second premolar.

Supernumerary teeth were found in 8 (3.9%) of our specimens. Seven cases occurred near the first upper premolar; one was posterior to the last upper molar. X-rays revealed distinct alveoli for three of the supernumerary premolars and the molar. In the other four cases, two teeth appeared to protrude from the same alveolus, although this was difficult to verify. Previous explanations for supernumerary dentition include: 1) splitting of a permanent or milk tooth bud; 2) failure to shed deciduous teeth; 3) atavism; and 4) "arising of tooth as a new crea- ture (genetic conditioning possible)" (Pavlinov 1975:516). We regard the latter as the least probable. Splitting of the tooth bud is a reasonable explanation in the four examples of two teeth, virtually identical in size, protruding from the same alveolus (Fig. 7).

To attribute supernumerary upper premolars in the river otter to atavism is unreasonable. The resulting five premolars exceeds the primi- tive number of four — already present in the normal upper premolar complement of the genus. The supernumerary premolars with distinct alveoli, therefore, are more likely due to failure of the deciduous teeth to shed. However, atavism is a likely explanation regarding the super- numerary molar. Because teeth tend to be lost at the end of a row (Hall

River Otter Skull Anomalies

105

Fig. 3. Irregular spacing of dentition in immature female river otter (AEL-304).

Fig. 4. Irregular tooth placement resulting in rotation from the toothrow (AEL-305).

106 Thomas D. Beaver, George A. Feldhamer, Joseph A. Chapman

'if

Fig. 5. Extreme alveolar thinning associated with upper molars (AEL-306).

•>

I'.

V

Fig. 6. Bilateral congenital agenesis of second upper premolars (AEL-307).

River Otter Skull Anomalies

107

	i ■

Fig. 7. Two teeth protruding from same alveolus. Apparent difference in length due to one tooth partially forced from alveolus (AEL-308).

Fig. 8. Supernumerary molar assumed to be atavistic (AEL-309).

108 Thomas D. Beaver, George A. Feldhamer, Joseph A. Chapman

1940), placement of the extra molar at the posterior end of the series (Fig. 8) would be expected if the condition were an atavistic trait.

Based solely on the age of certain anomalous skulls, dental and cranial anomalies in the river otter did not appear to be detrimental to the overall condition or survival of individuals. As noted, anomalous animals more than three years of age were found. The carcasses were not available to assess body fat or other indices of physical condition, however, so this conclusion must remain conjectural. We suspect, though, that in contrast to the possible detrimental effects of anomalies on individual mantled howler monkeys, Alouatta palliata, and subse- quent effects postulated for dominance relationships and population dynamics (Smith et al. 1977), anomalies probably are of little impor- tance in the population dynamics of the river otter.

ACKNOWLEDGMENTS.— We thank Dr. William Newman, Pam Askins and Denise Andrews, Radiology Department, Sacred Heart Hospital, Cumberland, Maryland, for providing X-rays of the study material. Dr. J. Edward Gates, Appalachian Environmental Labora- tory, critically reviewed the manuscript, and an anonymous reviewer provided helpful comments. This is Contribution No. 1284-AEL, Appala- chian Environmental Laboratory, Center for Environmental andEstuar- ine Studies, University of Maryland. Specimens were collected as part of a study funded by Maryland Federal Aid in Wildlife Restoration W-49-R.

LITERATURE CITED Choate, Jerry R. 1968. Dental abnormalities in the short-tailed shrew, Blarina

brevicauda. J. Mammal. 49{2):25 1-258. Clark, James W., E. Cheraskin and W. M. Ringsdorf, Jr. 1970. Diet and the

periodontal patient. Charles C. Thomas, Springfield, IL. 370 pp. Colyer, F. 1936. Variations and diseases of the teeth of animals. John Bale,

Sons and Danielsson, Ltd., London. 750 pp. Dougherty, Ellsworth C, and E. R. Hall. 1955. Biological relationships between

American weasel (genus Mustela) and nematodes of the genus Skrjabingy-

lus Petrov 1927 (Nematoda: Metastrongylidae), the causative organism of

certain lesions in weasel skulls. Rev. Iber. Parasitol., Tomo Extraordinario:

531-576. Hall, E. Raymond. 1940. Supernumerary and missing teeth in wild mammals of

the orders Insectivora and Carnivora, with some notes on disease. J. Dent.

Res. 79:103-143. Heran, Ivan. 1970. Anomalies in the position of lower teeth in the fisher, Martes

pennanti Erxl. (Mammalia; Mustelidae). Can. J. Zool. 48:1465. Hershkovitz, Phillip. 1970. Dental and periodontal diseases and abnormalities

in wild-caught marmosets (Primates — Callithricidae). Am. J. Phys.

Anthropol. 52:377-394.

River Otter Skull Anomalies 109

LaVelle, C. L. B., and W. J. Moore. 1972. The incidence of agenesis and poly- genesis in the Primate dentition. Am. J. Phys. Anthropol. 55:671-680.

Manville, Richard H. 1963. Dental anomalies in North American lynx. Z. Saeugeterkd. 28(3): 166-169.

Mowbray, Elmer E., D. Pursley and J. A. Chapman. 1979. The status, popula- tion characteristics and harvest of the river otter in Maryland. Md. Wildl. Admin. Publ. Wildl. Ecol. No. 2:1-16.

Pavlinov, I. Y. 1975. Tooth anomalies in some Canidae. Acta Theriol. 20(33): 507-519

Parmalee, Paul W., and A. E. Bogan. 1977. An unusual dental anomaly in a mink. J. Tenn. Acad. Sci. 52(3): 1 15-1 16.

Robinson, P. T. 1979. A literature review of dental pathology and aging by dental means in nondomestic animals — Parts I and II. J. Zoo Anim. Med. 70:57-65,81-91.

Sheppe, Walter. 1964. Supernumerary teeth in the deer mouse, Peromyscus. Z. Saeugeterkd. 29( l):33-36.

Shultz, Adolph H. 1923. Bregmatic fontanelle bones in mammals. J. Mammal. 4(2):65-77.

Smith, James D., H. H. Genoways and J. K. Jones, Jr. 1977. Cranial and dental anomalies in three species of Platyrrhine monkeys from Nicaragua. Folia Primatol. 25:1-42

van Soest, R. W. M., J. van der Land and P. G. H. van Bree. 1972. Skrjabingy- lus nasicola (Nematoda) in skulls of Mustela erminea and Mustela nivalis (Mammalia) from the Netherlands. Beaufortia 20:85-97

Accepted 30 November 1981

Life History and Ecology of

Chauliodes rastricornis Rambur and C. pectinicornis (Linnaeus)

(Megaloptera: Corydalidae) in Greenbottom Swamp,

Cabell County, West Virginia

Pamela S. Dolin and Donald C. Tarter

Department of Biological Sciences,

Marshall University, Huntington, West Virginia 25701

ABSTRACT. — Life history and ecology of two congeneric fishflies, Chauliodes rastricornis Rambur and C. pectinicornis (Linnaeus), were investigated from March 1978 to September 1980 at Greenbottom Swamp, Cabell County, West Virginia. Frequency histograms of head width indicate both species are univoltine. Larval C. rastricornis showed greatest growth (66%) between June and October, while C pectinicornis had two growth periods, from July to October (106%) and April to June (17%). Larvae of both species ingested organic detritus, copepods, and algae during the winter months, and ostracods, rotifers, cladocerans, and immature insects during the warmer seasons. Chauliodes rastricornis pupated from April to May and emerged at dusk from May to June; peak emergence was on May 9. Chauliodes pectinicornis pupated in June and emerged at night during July. The chi-square test showed a significant difference (0.05 level) from the 1:1 sex ratio in adults of C. rastricornis. Total counted eggs ranged from 922 to 1612 (x = 1 166) and 967 to 1500 (x = 1250) eggs per female in C. rastricornis and C pectinicornis, respectively.

INTRODUCTION

Several authors, including Moody (1877), Brimley (1908), Cuyler (1956, 1958), Hazard (1960), Baker and Neunzig (1968), Watkins et al. (1975), Tarter and Watkins (1974), Tarter et al. (1976) and Tarter et al. (1977), have reported ecological and taxonomical studies of members of the genus Chauliodes.

Chauliodes rastricornis Rambur is known from 24 eastern states and Ontario, Manitoba, and Nova Scotia (Davis 1903; Hazard 1960; Parfin 1952; Tarter et al. 1976), and Chauliodes pectinicornis (Linnaeus) from 26 eastern states and Quebec (Davis 1903; Hazard 1960; Tarter et al. 1976). These species are often sympatric. Chauliodes larvae inhabit a variety of lentic waters including marshes, swamps, swamp channels, ponds, small lakes, woodland pools, and sinkholes (Cuyler 1956). The objective of this investigation was to compare the life history and ecol- ogy of C. rastricornis and C. pectinicornis in Greenbottom Swamp, Cabell County, West Virginia.

MATERIALS AND METHODS

Greenbottom Swamp is located in the Union district of northwest Cabell County, West Virginia (Krebs and Teets 1913). The swamp is

Brimleyana No. 7:111-120. July 1981. Ill

112 Pamela S. Dolin and Donald C. Tarter

approximately 8050 m east of Homestead, West Virginia, on State Road n2. The study area is located 550 m from the Ohio River's south edge. The swamp proper is 1450 m long and has a contiguous marsh on the north end that continues another 1200 m, forming about 14 ha of swamp forest. The mean water depth varies from 0.5 m during dry peri- ods to 1.5 m during high waters. Elevation is 168 m above sea level. In the permanently inundated areas, the only tree species is black willow, Salix nigra Marsh. Button-bush saplings, Cephalanthus occidentalis L., have the greatest density of any woody plant seedling in the swamp.

The study began in March 1978 and ended in September 1980 (Dolin 1980). Larval collections of both species were made on a monthly basis, by searching under moss, duckweed, and loose bark of small stems and decaying logs.

Monthly water chemistry was assessed with a Hach chemical kit, Model AL-36B. The measurements taken included: hydrogen ion con- centration (pH) determined colorimetrically; dissolved oxygen and car- bon dioxide (mg/ 1); and total hardness and alkalinity (mg/ 1 of CaCO^). Water temperature ranges were measured with a Taylor maximum- minimum thermometer secured 0.3 m below the surface.

Head width was measured to obtain growth records. Total width was taken at the widest section of the head capsule, usually at the eyes. Using a compound microscope and a caliper, the measurements were taken to the nearest 0.1 mm. Monthly differences in head widths of 246 C. rastricornis and 1 12 C pectinicornis larvae were determined to show the mean, range, standard deviation, and two standard errors of the mean in a Hubbs and Perlmutter (1942) population range diagram. Per- cent growth was calculated monthly by comparing the mean head width values. Size classes were assessed by frequency histograms arranged in 0.5 mm head width categories.

Generally, foreguts were examined from five larvae of each species per month. After the head was removed with microdissecting scissors, the abdomen was cut and the foregut extracted. The gut was then slit ventrally and the contents placed on a clean slide for identification. The number of foreguts containing the various taxa was recorded, and the mean and percent frequency of occurrence were calculated.

Chauliodes pupae were obtained by digging into the grooves of logs that had started to desiccate due to receding spring waters. For both species, a few larvae in the last instars were reared to the pupal and adult stages. Rearing was accomplished by leaving the larvae in place and quickly transporting the log section to aerated pint jars containing water from the swamp. Larvae were placed one to a jar to help insure pupation by reducing competition for foodstuffs, to prevent cannibal- ism, and to determine when the individuals pupated. Pupae were placed

Corydalid Life Histories 113

in a rearing cage, and the range and mean pupation dates were recorded.

Adults were collected at night by means of a black light trap. For C. rastricornis, the trap was set out between 9 and 10 PM twice a week during the emergency period to obtain a quantitative number of emer- gents per hour. Various times wee sampled in hopes of finding the peak period of emergence for C. pectinicornis. The range and peak emergence times were registered.

To determine fecundity, the thorax and head of each adult fishfly was removed with microdissecting scissors. The abdomen was then cut ventrally and the ovaries with the eggs teased out. All eggs were counted and the length (mm) of the female's body was recorded. The mean and range were computed for each species.

RESULTS AND DISCUSSION

Swamp environment . — The pH ranged from 6.0 in February to 7.3 in November; x = 6.8. Dissolved oxygen values ranged from 2.0 to 11 mg/ 1 in August and March, respectively; x = 6.0 mg/ 1. In July, a car- bon dioxide value of 85 mg/ 1 was recorded versus 15 mg/ 1 in the win- ter and spring. Carbonate alkalinity values ranged from 51.3 mg/ 1 in December and January to 102.6 mg/ 1 in July and August; x = 73.4 mg/ 1. Total hardness ranged from 68.4 to 171.0 mg/ 1 in June and May, respectively; x = 99 mg/ 1.

Larval development .— Frequency histograms of head width indi- cated that the life cycles of both species are univoltine (Figs. 1, 2). In C. rastricornis, hatching probably begins in early June, but due to small size and concealment larvae were not collected until the end of June (Fig. 1). April was the last month larvae were found before they pupated. Hatching of C. pectinicornis occurred in July, making this month unique in that two different size classes of larvae as well as pupae were found (Fig. 2). Apparently, two cohorts were present.

Population range diagrams of larval head capsule widths show the monthly growth of both fishflies (Figs. 3, 4). Chauliodes rastricornis showed the greatest growth (66%) between June and October, while C. pectinicornis had two growth periods, from July to October (106%) and April to June (17%). Growth of both was retarded when the monthly water temperatures averaged below 15°C (November through March). The second growth period of C. pectinicornis corresponds with the absence of C. rastricornis larvae from the swamp (Fig. 5).

In North Carolina, Cuyler (1956) found prepupal C. rastricornis and larval C. pectinicornis from March 28 to April 17. He collected C. pectinicornis larvae on land in April even though no adults were found until the second week in May. This leaves five weeks for the fishfly

114

Pamela S. Dolin and Donald C. Tarter

A

d

40% -i

30% 207,, 10% 0%

50% -. 40% - 30% 20% H 10% 0%

A

Head Width (mm)

Fig. 1. Head-width frequencies at monthly intervals of Chauliodes rastricornis larvae from Greenbottom Swamp, Cabell County, West Virginia. Number in parentheses after month is sample size.

larvae to pupate in the drier environment before emerging as adults.

Larval food habits. — The following taxa were found in the fore- guts of Chauliodes larvae (Dolin 1980): ostracods, cladocerans, copep- ods, amphipods, odonates, rotifers, dipterans, and the alga Tribonema. Detritus and unidentifiable materials were also found.

Based on a yearly mean, organic detritus ranked first (96%) in fre- quency of occurrence in C. rastricornis, and ostracods (26%) were second. Chironomids were found with a frequency of 16 percent. Based on a yearly mean, organic detritus ranked first (82%) in frequency of occurrence in C. pectinicornis, and ostracods (14.5%) and cladocerans (14.5%) ranked second.

Copepods were found only in winter in both fishflies. The unidenti-

Corydalid Life Histories

115

40% 30%

20% 10%

0%

Aug. (10)

=-"J ,

50% g 40% = 30%

f 20% -

0%

20% 10% 0%

Sept. (101

60% -i Nov. (10)

50% -

Ak

Head Width (mm)

Fig. 2. Head-width frequencies at monthly intervals of Chauliodes pectinicornis larvae from Greenbottom Swamp, Cabell County, West Virginia. Number in parentheses after month is sample size.

fied materials, amphipods, and chironomids occurred sporadically throughout the year in both species. Empty foreguts were found 20 per- cent of the time in C. rastricornis and 30 percent in C. pectinicornis. These insects apparently indiscriminately ingested whatever food source was available during the season, with detritus being the only constant category in their diet.

Under laboratory conditions, Weed (1889) and Lintner (1892) fed fishfly larvae the backswimmer Notonecta undulata, dipteran larvae, and spiders. Davis (1903) reported the larvae as carnivores that ate smaller members of their own species, larvae of Sialis, caddisflies, dipte- rans, and other accessible soft-bodied insects.

116

Pamela S. Dolin and Donald C. Tarter

Head Width (mm)

Water Temp.

(C)

Fig. 3. Monthly variations in larval head widths of Chauliodes rastricornis from Greenbottom Swamp, Cabell County, West Virginia. Horizontal lines = means; vertical lines = ranges; open rectangles = standard deviation; closed rectangles = two standard errors of the mean; numbers above vertical lines = sample size; and solid line = mean water temperatures.

Pupal stage. — Pupae have been collected under bark and in decay- ing logs and stumps by various authors: Walsh (1863), Weed (1889), Needham and Betten (1901), and Brimley (1908). Parfin (1952) and Davis (1903) reported Chauliodes pupating under rocks, and Davis also observed them in rotten wood and earthen cells. Cuyler (1956) reported finding C. rastricornis in pine and oak stumps as far as 25 feet from water and observed eight C. pectinicornis pupae under the bark of one large log on the swamp bank.

In May, eight C. rastricornis pupae were found deep in rotten logs that were drying because of receding waters. Pupae of C. pectinicornis were found under bark of drying logs, and in earthen cells under logs left on the bank of the swamp after water had receded.

Seven C. rastricornis and eight C. pectinicornis were reared from larvae to adults. Chauliodes rastricornis larvae transformed to pupae from April 23 to May 1, with the pupal stage lasting a range of 6 to 18

Corydalid Life Histories

117

days; x = 12. Chauliodes pectinicornis larvae transformed to pupae between June 26 and July 28, with the pupal stage lasting from 8 to 17 days; x = 10.

Adult stage. — Seventy-seven C. rastricornis adults were collected from 9 May 1979 to 2 June 1980. The peak emergence was on 9 May, with 28 adults trapped in the night light per hour. Only five C. pectini- cornis males were collected between 14 and 30 July by light-trapping, even though trapping between 9 PM and 3 AM occurred from May to the end of August. This could be due to the low occurrence of C. pecti- nicornis in the swamp.

Davis (1903) reported collecting adult Chauliodes between 14 and 16 June, noting they lived only a few days in captivity and would not eat. Smith (1922) reported collecting adult Chauliodes in June, and Chaplain and Kirk (1926) trapped C pectinicornis from 11 July to 6 August in a fermented syrup of molasses and water. Tarter et al. (1977) recorded C pectinicornis emerging from 28 February in Louisiana to 1 1 November in Maryland, with the West Virginia emergence from 22 May to 9 September. Chauliodes rastricornis emerged from 10 January to 28

Head Width (mm)

1

July

Water Temp.

(C)

Fig. 4. Monthly variations in larval head widths of Chauliodes pectinicornis from Greenbottom Swamp, Cabell County, West Virginia. Legend symbols are the same as in Figure 3.

118

Pamela S. Dolin and Donald C. Tarter

^^H

Pupae

Larval Head Width (mm)

5 - Larval 4 - 3 - 2 _ 1 -

Width (mm)

Adults

. Pupae

January

I I

O November

Fig. 5. Summary graph of life cycles of Chauliodes rastricornis (bottom), and C. pectinicornis (top), from Greenbottom Swamp, Cabell County, West Virginia. Larval head width means are plotted against months of the year. Straight lines = emergence periods of pupae and adults.

December in Florida, and specimens of C. rastricornis in West Virginia were only found on 2 June (Tarter et al. 1977).

The C. rastricornis pupae reared in our laboratory transformed into adults between 29 April and 13 May, and the C. pectinicornis emerged between 10 July and 27 August. This separation of emergence times helps keep these two species from interbreeding. No mating, ovip- ositing, eating, or drinking was observed.

Brimley (1908) reported rearing larval Chauliodes caught on 4 April 1906 to adults, which first emerged 21 April and continued trans- forming until 20 May 1906. Parfin (1952) recorded rearing C. rastricor- nis, which first emerged on 5 June and lived approximately six days on honey and water.

The sex ratio of 77 C. rastricornis captured by the light traps was 5 males: 1 female, and a chi-square test indicated a significant difference (0.05 level). Since all five C. pectinicornis trapped were males, a chi- square value was not calculated.

Adult C. rastricornis, reared from pupae, showed a fecundity range of 922 to 1612 eggs (x = 1 166) per female, but light-trapped females of the same species contained between 8 and 637 eggs (x = 388). Chauliodes

Corydalid Life Histories 119

pectinicornis adults light-trapped by other collectors contained either all or none of their eggs, indicating only one oviposition. The fecundity ranged from 967 to 1500 eggs (x = 1250). No egg masses were found in the field.

Adult C. rastricornis laid their eggs a short distance away from the water (Needham and Betten 1901). The grayish eggs were layed in V- shape rows attached to flat surfaces. A pair of C. pectinicornis was observed and photographed while copulating on a fern. After the male left, the female, who had already laid most of her eggs, continued ovip- ositing using the outline of the pattern she had already begun. Around 900 eggs were laid, all of which hatched 17 days later. Davis (1903) reported 30 to 40 egg masses, of 1000 to 2000 eggs each, on the under- side of a wooden boat-landing in June and July. Other masses were as high as 10 to 15 feet above water on stones and leaves. Hatching was reported to occur five to six days later. On 16 June, Smith (1922) observed C. pectinicornis depositing 1000 to 2000 eggs in winding paral- lel rows one layer deep. The eggs hatched eight days later in a 45-minute period at night. A complete description of the egg masses and eggs of C. pectinicornis was given by Baker and Neunzig (1968).

ACKNOWLEDGMENTS.— Special thanks are extended to Miss Vickie Crager for typing the manuscript. Also, we express our appreciation to the following persons for help in the field work: Kerry Bledsoe, Bill Cremeans, Steve Lawton, Tom Rone, Mary Beth Roush, and Jan Taylor.

LITERATURE CITED Baker, J. R., and H. H. Neunzig. 1968. The egg masses, eggs, and first instar

larvae of eastern North America Corydalidae. Ann. Entomol. Soc. Am.

6/(5): 118 1-1 187. Brimley, C. S. 1908. Notes on some Neuropteroids from Raleigh, N.C. Entomol.

News 79:133-134. Chaplain, A. B., and H. B. Kirk. 1926. Bait pan insects. Entomol. News

57(9):288-291. Cuyler, R. D. 1956. Taxonomy and ecology of the larvae of sialoid Megaloptera

of east-central North Carolina, with a key to and descriptions of the larvae

of genera known to occur in the U. S. Unpubl. MS Thesis. N. C. State

Coll., Raleigh. 150 pp. 1958 The larvae of Chauliodes Latreille (Megaloptera: Corydalidae).

Ann. Entomol. Soc. Am. 57:582-586. Davis, K. C. 1903. Sialididae of North and South America. In Aquatic Insects

in New York State. N.Y. State Mus. Bull. 65:442-487. Dolin, Pamela S. 1980. The life history and ecology of Chauliodes rastricornis

Rambur and C. pectinicornis (Linnaeus) (Megaloptera: Corydalidae) in

Greenbottom Swamp, Cabell County, West Virginia. Unpubl. MS Thesis.

Marshall Univ., Huntington. 58 pp.

120 Pamela S. Dolin and Donald C. Tarter

Hazard, Edwin I. 1960. A revision of the genera Chauliodes and Nigronia

(Megaloptera: Corydalidae). Ann. Entomol. Soc. Am. 57:582-586. Hubbs, Carl L., and A. Perlmutter. 1942. Biometric comparison of several

samples, with particular reference to racial investigations. Am. Nat.

76:582-592. Krebs, C. E., and D. D. Teets. 1913. West Virginia Geological Survey County

Reports, Cabell, Wayne and Lincoln Counties. Wheeling News Litho Co.,

Wheeling. 483 pp. Lintner, J. A. 1892. Chauliodes pec Unicornis, rastricornis, and serricornis. N.Y.

State Entomol. 8th Annu. Rep.: 155-1 59. Moody, Henry L. 1877. Habits and transformations of Chauliodes pectinicor-

nis. Psyche 2:52-53. Needham, J. C, and C. Betten. 1901. Aquatic Insects in the Adirondacks. N.Y.

State Mus. Bull. 47:383-612. Parfin, Sophy L. 1952. The Megaloptera and Neuroptera of Minnesota. Am.

Midi. Nat. 47(2):42 1-434. Smith, J. B. 1922. Hatching in three species of Neuroptera. Ann. Entomol. Soc.

Am. 75:169-176. Tarter, Donald C, and W. D. Watkins. 1974. Distribution of the fishfly genera

Chauliodes Latreille and Nigronia Banks in West Virginia. Proc. W. Va.

Acad. Soc. 46(2): 146-150. , , M. Little and J. T. Goodwin. 1976. New state records of

fishflies (Megaloptera: Corydalidae). Entomol. News <S7(7-8):223-228. , , and D. L. Ashley. 1977. Seasonal emergence

patterns of fishflies east of the Rocky Mountains (Megaloptera: Corydali- dae). Entomol. News 88(3-4):69-16.

Walsh, B. D. 1863. Notes on Neuroptera. Proc. Entomol. Soc. Phil. 75:182-272.

Watkins, William D., D. D. Tarter, M. L. Little and S. D. Hopkins. 1975. New records of fishflies for West Virginia (Megaloptera: Corydalidae). Proc. W. Va. Acad. Sci. 47(1): 1-5.

Weed, C. M. 1889. Studies in pond life. Ohio Agric. Exp. Stn. Tech. Bull. 7:4-17.

Accepted 29 April 1982

Annotated Checklist of the Mammals of Georgia

Joshua Laerm, Lloyd E. Logan, M. Elizabeth McGhee

Museum of Natural History and Department of Zoology, University of Georgia, Athens, Georgia 30602

AND

Hans N. Neuhauser

The Georgia Conservancy, Savannah, Georgia 31405

ABSTRACT. — Previous accounts listed 79 species of mammals occurring in Georgia. Taxonomic changes and improved distributional records now allow listing of 90 species — 74 non-marine — with 20 of them represented by more than one subspecies. Consequently, Georgia has 1 18 taxonomically distinct mammalian species and subspecies.

INTRODUCTION

Since publication of Mammals of Georgia (Golley 1962) there has been a considerable increase in knowledge concerning this group of ver- tebrates in the state. There have been, for example, several new state records (e.g., Wharton and White 1967), a number of taxonomic and/ or nomenclatural changes (e.g., Williams and Genoways 1980), and some significant range extensions (Laerm et al. 1980). A review of Georgia's cetaceans (Neuhauser and Ruckdeschel 1978) provided a much needed update of the known ocurrences of marine mammals in Georgia's coas- tal waters. Recent regional studies (Wharton 1973; Neuhauser and Baker 1974; Laerm et al. 1980) provided useful information regarding the distribution and abundance of a number of species.

In general, the mammals of Georgia are poorly known. Many spe- cies are extremely rare, some are known only from very old historical data, and there are a number of taxonomic problems with others. Although we are preparing a book on the mammals of Georgia, its completion will require several more years of work. We provide here an updated and comprehensive annotated checklist of mammal species and subspecies known to occur in Georgia and its coastal waters. For each species and subspecies, we include distributional data and brief com- ments regarding their taxonomic and conservation status.

MATERIALS AND METHODS

Scientific and common names are those used by Jones et al. (1979). Distributional ranges are based on voucher specimens in a number of museums, published accounts, our observations, and University of

Brimleyana No. 7:121-135. July 1981. 121

122 Joshua Laerm, et al.

Georgia Museum of Natural History records. Subspecific designations are those of Hall (1981), unless otherwise noted. Data were obtained from mammal collections in the following institutions: Academy of Nat- ural Sciences of Philadelphia; American Museum of Natural History; Auburn University; Carnegie Museum of Natural History; Charleston Museum of Natural History; Cornell University; Delaware Museum of Natural History; Field Museum of Natural History; Florida State Museum; Museum of Comparative Zoology, Harvard University; Museum of Texas Tech University; National Museum of Natural His- tory; University of Georgia Museum of Natural History; University of Illinois Museum of Natural History; University of Kansas Museum of Natural History; and University of Michigan Museum of Zoology.

SPECIES LIST

Didelphis virginiana (Linnaeus), Virginia Opossum. Statewide; common in nearly all habitats, including most urban areas. Two sub- species are recognized (Gardner 1973; Hall 1981): Didelphis virginiana virginiana Kerr occurs from the Upper Coastal Plain southward, and is replaced northward by D. v. pigra Bangs.

Sorex cinereus cinereus Kerr, Masked Shrew. A rare species that reaches the southern limit of its range in northern Georgia. Within the state it is known from only three specimens, taken in mesic forest habi- tats of Towns County (Wharton 1968a).

Sorex longirostris longirostris Bachman, Southeastern Shrew. Statewide but uncommon. The species generally inhabits mesic wood- land and open field habitats (Golley 1962).

Sorex fumeus fumeus Miller, Smoky Shrew. An uncommon shrew that reaches the southeastern limit of its range in northeast Georgia, where it is known from Fannin, Murray, Rabun, Towns, and Union counties. It is most commonly found in deciduous forest leaf litter (Hamilton 1940).

Microsorex hoyi winnemana Preble, Pygmy Shrew. This rare shrew, the smallest North American mammal, reaches the southern limit of its range in northeast Georgia (Long 1974; Wharton 1968a), where it occurs in mesic woodlands of Towns County. Long (1972, 1974) identi- fied the species occurring in Georgia as M. thompsoni, but we follow van Zyll de Jong (1976) and Hall (1981).

Blarina brevicauda churchi Bole and Moulthrop, Short-tailed Shrew. This common shrew occurs in mesic habitats in the northern half of the state, exclusive of the extreme northwest, south to the Fall Line (French 1982).

Blarina carolinensis carolinensis (Bachman), Southern Short-tailed Shrew. A common inhabitant of mesic habitats in the Coastal Plain,

Georgia Mammals 123

south of the Fall Line. This shrew also occurs in the extreme northwest- ern part of the state (French 1982).

Cryptotis parva Say, Least Shrew. An uncommon shrew of field habitats throughout the state, which is most abundant on the Coastal Plain (Golley 1962). Two subspecies are recognized in Georgia (Hall 1981): Cryptotis parva parva (Say) is statewide except in the extreme south, and C. p. floridana (Merriam) is known only from Thomas, Grady, Camden and Mcintosh counties (Golley 1962; Neuhauser 1972).

Scalopus aquaticus (Linnaeus), Eastern Mole. A common fossorial species inhabiting a variety of habitats statewide. Three subspecies are recognized in Georgia (Yates and Schmidly 1978; Hall 1981): Scalopus aquaticus aquaticus (Linnaeus) is known only from the extreme north- east; S. a. australis (Chapman) occurs in the southeastern third of the state; and S. a. howelli (Jackson) occurs throughout the northwestern two-thirds of the state.

Condylura cristata parva Paradiso, Star-nosed Mole. A very rare, largely aquatic mole that reaches the southern limit of its range in Georgia. It is known only from Charlton, Chatham, Clinch, Effingham, Jackson, and Union counties (Peterson and Yates 1980).

Myotis lucifugus lucifugus (LeConte), Little Brown Myotis. This bat is known only from Bartow, Dade, Polk, Towns, and Walker coun- ties. Other previously published localities (Golley 1962; Hall and Kelson 1959; Hall 1981) were based on misidentifications (Davis and Rippy 1968). The type locality, "Georgia, probably the LeConte Plantation, near Riceboro, Liberty County . . ." (Miller and Allen 1928) is ques- tionable because there are no specimens with data to suggest that M. lucifugus occurs in the Coastal Plain (Davis and Rippy 1968). This uncommon species inhabits buildings and caves.

Myotis austroriparius (Rhoads), Southeastern Myotis. Statewide, but not known from the southeastern tier of counties adjoining South Carolina. The species is common in the southern part of the state and is often confused with M. lucifugus (Davis and Rippy 1968). It inhabits trees, caves, and buildings (Hall and Kelson 1959).

Myotis grisescens Howell, Gray Myotis. Known in Georgia from only two localities, in Polk and Clarke counties (Baker 1965). These specimens were probably transients, since the primary range of this cave dweller is northwest of Georgia. The species is considered endangered under State and Federal regulations (Odom et al. 1977).

Myotis keenii septentrionalis (Trouessart), Keen's Myotis. This sol- itary bat is rare in Georgia, where most records are from the northwest- ern mountains. There it inhabits caves (Fitch and Schump 1979). Van Zyll de Jong (1979) suggested that the nominal subspecies septentriona- lis is a separate species from M. keenii. We follow Hall (1981) and Jones

124 Joshua Laerm, et al.

et al. (1979) in recognizing M. k. septentrionalis.

Myotis sodalis Miller and Allen, Indiana Myotis. This species, con- sidered endangered under State and Federal regulations (Odom et al. 1977), is known in Georgia from only a single limestone cave in Dade County.

Myotis leibii (Audubon and Bachman), Small-footed Myotis. This rare, solitary bat reaches the southeastern limit of its range in north Georgia. Only three specimens from two localities in Dade and Union counties are known (Baker 1965). It inhabits caves, trees, and buildings.

Lasionycteris noctivagans (LeConte), Silver-haired Bat. This migra- tory tree dweller is fairly common statewide except in the lower Coastal Plain.

Pipistrellus subflavus (Cuvier), Eastern Pipistrelle. The pipistrelle is common in caves and trees statewide. Two subspecies are recognized in Georgia (Hall 1981): Pipistrellus subflavus subflavus (Cuvier) is state- wide except in the extreme southeastern Coastal Plain, where it is replaced by P. s. floridanus (Davis).

Eptesicus fuscus fuscus (Palisot de Beauvois), Big Brown Bat. Common statewide in attics of rural and urban homes as well as in outbuildings. The extreme southeastern counties may represent a region of intergradation between E. f fuscus and E. f osceola Rhoads, which occurs in Florida.

Lasiurus borealis borealis (M tiller), Red Bat. A common migratory bat that roosts in trees statewide.

Lasiurus seminolus (Rhodes), Seminole Bat. Statewide, except for the mountain regions, and the most common bat in the Coastal Plain. It occurs most commonly in trees.

Lasiurus cinereus cinereus (Palisot de Beauvois), Hoary Bat. A large, uncommon tree dweller that is statewide during migration.

Lasiurus intermedius floridanus (Miller), Northern Yellow Bat. In Georgia, this rare tree dweller is known from less than a dozen speci- mens, all from the Coastal Plain.

Nycticeius humeralis humeralis (Rafinesque), Evening Bat. This common colonial bat, statewide in occurrence, is frequently found in attics of rural homes (Watkins 1972).

Plecotus rafinesquii Lesson, Rafinesque's Big-eared Bat. Although probably statewide in distribution, this bat is an uncommon inhabitant of man-made shelters or hollow trees (Jones 1977). Golley (1962), Jones (1977), and Hall (1981) indicated its range as statewide, but there are no collection records from the entire Piedmont Plateau and only one from the upper Coastal Plain (Grady County). It is apparently declining in numbers in south Georgia, perhaps due to habitat manipulation (Laerm et al. 1980). Two subspecies are recognized in Georgia (Jones 1977; Hall

Georgia Mammals 125

1981): Plecotus rafinesquii rafinesquii Lesson occurs in the extreme northern tier of counties, and P. r. macrotis LeConte in the Coastal Plain.

Tadarida brasiliensis cynocephala (LeConte), Brazilian Free-tailed Bat. This uncommon bat occurs in the Piedmont and Coastal Plain of Georgia. It is found most commonly in buildings and under bridges, and colonies may be fairly large.

Dasypus novemcinctus mexicanus Peters, Nine-banded Armadillo. This species was first recorded in Georgia in the early 1950s (Fitch et al. 1952) and is now common in a variety of habitats in most of the lower Coastal Plain Sand Hills (Humphrey 1974). There are isolated records from the upper Coastal Plain as far north as Stewart and Bibb counties, just south of the Fall Line.

Sylvilagus palustris palustris (Bachman), Marsh Rabbit. A com- mon inhabitant of lowlands, marshes, and flood plain habitats in the Coastal Plain.

Sylvilagus floridanus mallurus (Thomas). Eastern Cottontail. State- wide, and the most abundant rabbit in Georgia, where it occurs in a variety of habitats.

Sylvilagus transitionalis (Bangs), New England Cottontail. Known only from a few specimens collected in the Blue Ridge Province in 1908 and 1909. There are no recent records of its occurrence in the state, so it may be extirpated.

Sylvilagus aquaticus aquaticus (Bachman), Swamp Rabbit. Occurs north of the Fall Line in Georgia, but extends south into the upper Coastal Plain in the western counties. It is a common inhabitant of flood plains and creeks throughout its range.

Tamias striatus striatus (Linnaeus), Eastern Chipmunk. Found in the mountains, northern sections of the Piedmont, and scattered locali- ties in western sections of the upper Coastal Plain. Common in open woodlands and urban areas.

Marmota monax monax (Linnaeus), Woodchuck. This species reaches the southern limit of its range in north Georgia, where it is known from few and generally scattered localities in the mountains, south to Cherokee and Barrow counties. It occurs in woodlands, uncul- tivated fields, and roadside habitats.

Sciurus carolinensis carolinensis (Gmelin), Gray Squirrel. This is the most common squirrel in Georgia, where it occurs statewide in a variety of habitats.

Sciurus niger Linnaeus, Fox Squirrel. Although statewide in distri- bution, this squirrel is generally least abundant in the upper Piedmont and mountains. It may be locally abundant. Hall (1981) and Golley (1962) suggested that two subspecies— S. n. bachmani Lowery and Davis,

126 Joshua Laerm, et al.

and S.n. shermani Moore — occur in the extreme northwestern and extreme southeastern parts of the state, respectively. However, since specimens from localities in those regions cannot with certainty be assigned to those subspecies, we refer all Georgia specimens to S.n. niger Linnaeus. Laerm is studying the taxonomic status of the subspe- cies in the state. Sciurus n. rufiventer Geoffroy St. Hilaire was intro- duced to Ossabaw Island in the 1920s (J. Jenkins, pers. comm.; Hilliard 1979) and is apparently well established there.

Tamiasciurus hudsonicus abeiticola (Howell), Red Squirrel. Reaches the southern limit of its range in extreme northeastern Georgia, where it is only locally common (Wharton 1968b) in coniferous or mixed conif- erous forests (Howell 1929; Odum 1949) of White, Habersham, Towns, and Rabun counties.

Glaucomys volans (Linnaeus), Southern Flying Squirrel. This squirrel is statewide, but uncommon to locally abundant in a variety of habitats in mixed pine-hardwood forests. Two subspecies are recognized in Georgia (Dolan and Carter 1977): Glaucomys volans querceti (Bangs) is restricted to the southeastern counties, and G. v. saturatus Howell occurs throughout most of the rest of the state.

Geomys pinetis Rafinesque, Southeastern Pocket Gopher. This species occurs in the Sand Hills region south of the Fall Line, between Columbus and Augusta. It is common in a variety of Sand Hill habitats, principally along well drained roadsides. Four species (G. colonus, G. cumberlandius, G. fontanelus, and G. pinetis) were previously recog- nized in Georgia, but recent studies (Williams and Genoways 1980; Laerm 1981; Laerm et al., in press) have shown that the Geomys com- plex is represented in the state by a single species, G. pinetis. Two sub- species are recognized: Geomys pinetis pinetis Rafinesque occurs state- wide south of the Fall Line, and G. p. fontanelus Sherman is known only from its type locality, seven miles northwest of Savannah. Exten- sive surveys by the authors failed to locate G. p. fontanelus, and we believe it is extinct.

Castor canadensis carolinensis Rhoads, Beaver. An important statewide fur bearer in Georgia, the beaver is common in a variety of freshwater aquatic habitats. It is less common in sandy soils of the lower Coastal Plain and the Ridge and Valley province.

Oryzomys palustris palustris (Harlan), Marsh Rice Rat. Statewide and locally abundant in salt and freshwater marshes.

Reithrodontomys humulis humulis (Audubon and Bachman), East- ern Harvest Mouse. Occurs widely throughout the Southeast, but only a few scattered locality records are available for Georgia; distribution is probably statewide. The species may be locally abundant in old-fields, thickets, and meadows.

Georgia Mammals 127

Peromyscus maniculatus nubiterrae Rhoads, Deer Mouse. This very common species reaches the southern limit of its range in Georgia, where it is found in the Blue Ridge Province and extreme northeastern Piedmont, south into Lumpkin, White, and Hall counties. It occurs at higher elevations, generally over 850 m (2800 ft) in mesic hardwood forest.

Peromyscus polionotus (Wagner), Oldfield Mouse. A very common mammal of the Piedmont and Coastal Plain. Taxonomic distinctions of the three recognized subspecies that occur in Georgia (Hall 1981) are poorly defined, but they include: Peromyscus polionotus polionotus (Wagner), south of the Fall Line; P. p. colemani Schwartz, north of the Fall Line; and P. p. subgriseus (Chapman), in the extreme southwestern and south central counties.

Peromyscus leucopus leucopus (Rafinesque), White-footed Mouse. Very common in a variety of woodland habitats north of the Fall Line.

Peromyscus gossypinus (LeConte), Cotton Mouse. This mouse, the largest Peromyscus in Georgia, is common in a variety of habitats below the Fall Line. There are also isolated records from the extreme eastern Piedmont counties, and the Appalachian Valley. Three subspecies are recognized in Georgia (Hall 1981): Peromyscus gossypinus gossypinus (LeConte) inhabits mainland Georgia southeast of a line between Columbus and the extreme northeastern part of the state; P. g. anastasae Bangs is known only from Cumberland and Little Cumberland islands (Neuhauser 1978), but the subspecific affinities of the coastal island populations are suspect and currently under review by Laerm; and P. g. megacephalus (Rhoads) inhabits mainland Georgia northwest of a line between Columbus and the extreme northeastern part of the state.

Ochrotomys nuttalli (Harlan), Golden Mouse. This statewide spe- cies is generally restricted to woodland habitats, where it is common. Two subspecies are recognized in Georgia (Hall 1981): Ochrotomys nut- talli nuttalli (Harlan) inhabits the Piedmont and Coastal Plain, and O. n. aureolus (Audubon and Bachman) the Cumberland Plateau, Ridge and Valley, and Blue Ridge provinces.

Sigmodon hispidus Say and Ord, Hispid Cotton Rat. A statewide species that is very abundant in open fields and along roadsides. Two subspecies are recognized in Georgia (Hall 1981): Sigmodon hispidus hispidus Say and Ord occurs in the extreme southeastern counties, and S. h. komareki Gardner throughout the remainder of the state.

Neotoma floridana (Ord), Eastern Woodrat. Although widely dis- tributed almost statewide, and fairly common throughout a variety of woodland habitats, there are o records from the Piedmont Plateau. Three subspecies are recognized in Georgia (Wiley 1980): Neotoma flor- idana floridana (Ord) occurs in the Coastal Plain; N. f haemotoreia

128 Joshua Laerm, et al.

Howell in the Blue Ridge and Appalachian provinces; and N.f. illinoen- sis Howell on the lookout Plateau. We follow Schwartz and Odum (1957) in recognizing N. f. illinoensis as the subspecies occurring in Dade County.

Clethrionomys gapperi carolinensis (Merriam), Southern Red- backed Vole. Reaches the southern limit of its range in north Georgia, where it occurs only rarely in mesic hardwood habitats, generally above 610 m (2000 ft), in Union, Town, and Rabun counties.

Microtus pennsylvanicus pennsylvanicus (Ord), Meadow Vole. A rare species in north Georgia, where it reaches the southern limit of its range. It is found in the upper Piedmont, Blue Ridge, and Appalachian provinces.

Microtus pinetorum (LeConte), Woodland Vole. A semi-fossorial mouse found statewide except in the extreme southeastern counties. It is most common in wooded areas, but may be locally abundant in old- fields and orchards. Three subspecies are recognized in Georgia (Smolen 1981; Hall 1981): Microtus pinetorum pinetorum (LeConte) occurs throughout the Piedmont and the upper Coastal Plain; M. p. auricularis (Bailey) occurs in the Ridge, Valley, and Lookout provinces; and M. p. parvulus (Howell) occurs in the extreme south central counties.

Neofiber alleni exoristus Schwartz, Round-tailed Muskrat. This species occurs in the Okefenokee Swamp and surrounding areas of Camden, Charlton, and Ware counties, but is common only in the prairie habitat of the Okefenokee swamp.

Ondatra zibethicus zibethicus (Linnaeus), Muskrat. Statewide north of the Fall Line, and locally abundant in appropriate aquatic habitats.

Rattus rattus (Linnaeus), Black Rat. An introduced, statewide spe- cies, most common on the lower Coastal Plain.

Rattus norvegicus (Berkenhout), Norway Rat. An introduced, statewide species, common in association with human habitations, dumps, and similar places, although it also occurs in a variety of natural habitats.

Mus musculus (Linnaeus), House Mouse. An introduced, statewide species, most frequently found in association with human habitations but also known from a variety of natural habitats.

Zapus hudsonius americanus (Barton), Meadow Jumping Mouse. A rare species in Georgia, where it is at the southern limit of its range, and known only from widely scattered locations in the Piedmont, Blue Ridge, and Appalachian provinces. It is generally restricted to moist meadows.

Napaeozapus insignis roanensis (Preble), Woodland Jumping Mouse. Rare in Georgia, where it is at the southern limit of its range and occurs in the extreme northern counties. It generally occurs at ele-

Georgia Mammals 129

vations over 730 m (2400 ft) in cool, moist woodland habitats.

Myocastor coypus bonairiensis (Geoffroy St. Hilaire), Nutria. Dis- tribution in Georgia of this introduced species is poorly known, but it appears restricted to swampy habitats in the south central counties. It is known from Thomas, Brooks, Colquitt and Lowndes counties, and the region of the Chattahoochee River north to Fort Gordon (J. Jenkins, pers. comm.).

Mesoplodon densirostris (Blainville), Tropical Beaked Whale. A single stranding record is known from Cumberland Island (Neuhauser and Ruckdeschel 1978).

Mesoplodon europaeus (Gervais), Gervais' Beaked Whale. Known from a single stranding on Ossabaw Island (Neuhauser and Ruckdeschel 1978).

Ziphius cavirostris (Cuvier), Goose-beaked Whale. Neuhauser and Ruckdeschel (1978) reported six stranding records for this species.

Kogia breviceps (Blainville), Pygmy Sperm Whale. Neuhauser and Ruckdeschel (1978) reported 24 stranding events in Georgia.

Kogia simus (Owen), Dwarf Sperm Whale. Known from eight stranding events on Ossabaw, Cumberland and Little Cumberland Islands (Neuhauser and Ruckdeschel 1978) and from Wassaw Island.

Stenella plagiodon (Cope), Atlantic Spotted Dolphin. This species is known only from sightings offshore.

Steno bredanensis (Lesson), Rough-toothed Dolphin. One strand- ing event involving two animals is known from Little Cumberland Island (Richardson 1973).

Tur slops truncatus (Montague), Bottle-nosed dolphin. Neuhauser and Ruckdeschel (1978) reported 41 stranding events in Georgia.

Pseudorca crassidens (Owen), False Killer Whale. Known from a single stranding record on Tybee Island (Caldwell and Golley 1965).

Globlcephala macrorhynchus (Gray), Short-finned Pilot Whale. Neuhauser and Ruckdeschel (1978) reported 17 stranding events on the Georgia coast.

Balaenoptera edeni Anderso, Bryde's Whale. A 1978 stranding in Chatham County represented the first record of this species for Georgia (Neuhauser and Ruckdeschel 1978).

Megaptera novaeangllae (Borowski), Hump-backed Whale. A sin- gle stranding record on Sapelo Island is recorded (Neuhauser and Baker 1974). Neuhauser and Ruckdeschel (1978) suggested that the hump-backed whale may inhabit Georgia's coastal waters more fre- quently than stranding records indicate.

Balaena glaclalis (Borowski), Black Right Whale. Three stranding records are known, one from "near Savannah" (Neuhauser and Ruckde- schel 1978), one from Little St. Simons Island, and one from Ossabaw

130 Joshua Laerm, et al.

Island. Transients are occasionally observed offshore (Odom et al. 1977).

Canis latrans Woodhouse, Coyote. Wild canids predominantly occur in the western half of the state. It is a matter of conjecture whether they are coyotes introduced by sportsmen as suggested by Gol- ley (1962), or whether they represent the eastern extension of a hybrid swarm of coyote and red wolf (C rufus) as indirectly suggested by Paradiso and Nowak (1971). Identifications are further complicated by interbreeding with domestic dogs, C. familiaris.

Vulpes vulpes fulva (Desmarest), Red Fox. The red fox occurs statewide, but mostly north of the Fall Line, although it is known from widely scattered locations throughout the Coastal Plain. It usually pref- ers open habitat.

Urocyon cinereoargenteus (Schreber), Gray Fox. The gray fox is more common in Georgia than the red fox and is statewide. It is known from a variety of open and woodland habitats. Two subspecies are rec- ognized in Georgia (Hall 1981): Urocyon cinereoargenteus cinereoargen- teus (Schreber) occurs in the northern part of the state, south to the central Piedmont, and U. c. floridanus Rhoads occurs in the southern part of the state.

Ursus americanus Pallas, Black Bear. Statewide, but uncommon and generally restricted to wooded habitat in remote mountainous regions, the Ocmulgee River area, and the Okefenokee Swamp. Two subspecies are recognized in Georgia (Hall 1981): Ursus americanus americanus Pallas is primarily restricted to the mountains, although there are a few scattered locality records from the Piedmont and upper Coastal Plain (Jenkins 1953); and U. a. floridanus Merriam has its larg- est concentrations in the Okefenokee Swamp, with smaller populations in the Ocmulgee River region.

Procyon lotor (Linnaeus), Raccoon. Very common in a wide var- iety of habitats statewide. Four subspecies are recognized in Georgia (Hall 1981): Procyon lotor elucus Bangs occurs in the extreme southern tier of counties, including Okefenokee Swamp; P. I. litoreus Nelson and Goldman occurs in the eastern part of the lower Coastal Plain; P. I. solutus Nelson and Goldman occurs in the tier of counties adjacent to South Carolina; and P. I. varius Nelson and Goldman occurs in the upper Coastal Plain, and north throughout the state.

Mustela frenata Lichtenstein, Long-tailed Weasel. Statewide, gen- erally in brushland and fields. Two subspecies are recognized in Georgia (Hall 1981): Mustela frenata noveboracensis (Emmons) occurs in the northern Piedmont and mountain provinces, and M.f olivacea Howell in the southern Piedmont and Coastal Plain.

Mustela vison Schreber, Mink. Statewide, and generally distributed

Georgia Mammals 131

around freshwater streams and fresh and salt water marshes. Two sub- species are recognized in Georgia (Hall 1981): Mustela vison lutensis (Bangs) in the Coastal counties, and M. v. mink Peale and Palisot de Beauvois in the remainder of the state

Spilogale putorius putorius (Linnaeus), Eastern Spotted Skunk. Statewide, except in the lower Coastal Plain, and uncommon. Distrib- uted around farmlands and rarely occurs in woodlands.

Mephitis mephitis (Schreber), Striped Skunk. The more common skunk in Georgia, occurring statewide and usually around farmlands, less commonly in woodlands. Two subspecies are recognized in Georgia (Hall 1981): Mephitis mephitis elongata Bangs occurs in the northeast- ern, eastern and southern parts of the state, and M. m. nigra (Peale and Palisot de Beauvois) in the northwestern and west central parts.

Lutra canadensis lataxina Cuvier, River Otter. Fairly common in rivers and streams, as well as fresh and salt water marshes of the Coas- tal Plain and lower Piedmont.

Zalophus californianus (Lesson), California Sea Lion. Accidentally or intentionally released into waters of the southeastern United States, this species is reported from offshore Georgia, east of Mcintosh County (Caldwell et al. 1971).

Cystophora cristata (Erxleben), Hooded Seal. Neuhauser and Ruckdeschel (1978) reported on an unpublished note by Ivan Tompkins of a specimen from Bullhead Bluff on the Satilla River, Camden County, that was tentatively identified as C. cristata. There are no recent sightings of the species in southeastern United States waters (Schmidly 1981), and the species is presumed extirpated from this area.

Felis concolor coryi Bangs, Mountain Lion. Jenkins (1971) and Odum et al. (1977) reported recent unconfirmed sightings in Georgia. These are, however, very suspect, and it is very likely that the species has been extirpated in Georgia.

Felis rufus (Schreber), Bobcat. A common but secretive animal statewide in river-bottom swamps, brush and thickets. Two subspecies are recognized in Georgia (Hall 1981): Felis rufus rufus (Schreber) occurs in the mountain provinces; and F. r. floridanus (Rafinesque) in the Piedmont and Coastal Plain.

Trichechus manatus latirostris (Harlan), Manatee. Occurs occa- sionally in Georgia's coastal waters, most commonly south of the Altamaha River, especially during warmer weather.

Sus scrofa, Feral Pig. Feral pigs occur in the lower Coastal Plain and mountain provinces, and are most common in coastal swamps and marshes, less common in boreal habitats.

Cervus dama Linnaeus, Fallow Deer. Originally introduced for hunting, this species is locally abundant in Glynn County (Neuhauser and Baker 1974).

132 Joshua Laerm, et al.

Odocoileus virginianus (Zimmerman), White-tailed Deer. Common in a wide variety of wooded habitats statewide. Three subspecies are recognized as occurring in Georgia (Hall 1981). However, the near extirpation of deer by the later 1800s (Jenkins 1953), and subsequent restocking and interbreeding of individuals from various parts of the species' range, make subspecific distinction unreliable. We can find no records of recent introductions of deer onto Blackbeard Island, the type locality for O. v. nigribarbis (Goldman and Kellogg). This subspecies would appear to be the only recognizably distinct subspecies of deer occurring in Georgia.

DISCUSSION

We recognize here 90 species and 118 subspecies of mammals occurring in Georgia and its coastal waters. Of these, 74 species are non-marine and 16 are marine. Not included in this list are four species that recently ranged into Georgia but have been extirpated — Canis rufus (Red Wolf), Canis lupus (Gray Wolf), Cervus elaphus (Elk), and Bison bison (Bison). Three species and one subspecies listed herein are believed extirpated from Georgia. They are Sylvilagus transitionalis (New Eng- land Cottontail), Geomys pinetis fontanelus (Sherman's Pocket gopher), Cystophora cristata (Hooded Seal), and Felis concolor (Mountain Lion).

The non-game mammals of Georgia are poorly known. At present only very approximate distributional ranges for most species are availa- ble. Certain areas of the state, such as Okefenokee Swamp, the extreme northeastern counties, and the area immediately surrounding Athens, Clarke County, and the barrier islands, have received considerable attention, yet most parts of the state have not. There are glaring gaps in the known ranges of even the most common forms (e.g., Peromyscus). Many of the older collection records for several species (e.g., Sylvilagus transitionalis and Cystophora cristata) probably do not reflect current ranges. A large number of species listed above are known from a single, or at best a few, records (e.g., several marine mammals, Sorex cinereus, and Microsorex hoyi). Most of these species occur in Georgia at the extreme periphery of their natural ranges. Several species (e.g., Sorex dispar, Synaptomys cooperi, and Phoca vitulina) have known distribu- tional ranges that very closely approach Georgia and appropriate habi- tat for them appears to exist in the state. They may well occur in Georgia, but as yet there are no records.

There are a significant number of taxonomic problems (see above), particularly with respect to characterization of subspecies. Attention to these problems is crucial in our attempts to assess the status of Georgia's mammal fauna (see Laerm et al., in press b). We hope that this checklist may provide students of mammalogy a background for further study.

Georgia Mammals 133

ACKNOWLEDGMENTS.— We thank the curatorial staffs of the respective institutions listed under Materials and Methods for access to their mammal collections. This paper was supported by the Department of Zoology, University of Georgia, and in part by grant-in-aid funds under Section 6 of the Endangered Species Act of 1973. This is a con- tribution of the University of Georgia Museum of Natural History.

LITERATURE CITED

Baker, Wilson W. 1965. A contribution to the knowledge of the distribution and movements of bats in North Georgia. Unpubl. M.S. Thesis, Univ. Ga., Athens. 22 pp.

Caldwell, D.K., and F.B. Golley. 1965. Marine mammals from the coast of Georgia to Cape Hatteras. J. Elisha Mitchell Sci. Soc. #7:24-32.

, H. Neuhauser, M.C. Caldwell and H.W. Coolidge. 1971. Recent

records of marine mammals from the coasts of Georgia and South Caro- lina. Cetology5:l-12.

Davis, Wayne H., and C.L. Rippy. 1968. Distribution of Myotis lucifugus and My otis austroriparius in the southeastern United States. J. Mammal. 49:113-117.

Dolan, Patricia G., and D.C. Carter. 1977. Glaucomys volans. Mamm. Species 75:1-6.

Fitch, Henry S., P. Goodrum and C. Newman. 1952. The armadillo in the south- eastern United States. J. Mammal. 53:21-37.

Fitch, John H., and K.A. Schump, Jr. 1979. Myotis keenii. Mamm. Species 727:1-3.

French, Thomas W. 1982. Notes on the distribution and taxonomy of short- tailed shrews (Genus Blarina) in the Southeast. Brimleyana 6:101-1 10.

Gardner, Alfred L. 1973. The systematics of the genus Didelphis (Marsupialia: Didelphidae) in North and Middle America. Spec. Publ. Mus. Texas Tech Univ. No. 4. 81 pp.

Golley, Frank B. 1962. Mammals of Georgia. Univ. Ga. Press, Athens. 218 pp.

Hall, E. Raymond. 1981. Mammals of North America. 2nd Ed. John Wiley and Sons, New York. 1 181 pp.

, and K.R. Kelson. 1959. Mammals of North America. 1st Ed. Ronald

Press, New York. 1083 pp.

Hamilton, William J., Jr. 1940. The biology of the Smoky Shrew (Sorex fumeus fumeus Miller). Zoologica 25:473-492.

Hilliard, Thomas H. 1979. Radio-telemetry of fox squirrels in the Georgia coastal plain. Unpubl. M.S. Thesis, Univ. Ga., Athens. 120 pp.

Howell, Arthur H. 1929. Description of a new red squirrel from North Carolina. J. Mammal. 70:75-76.

Humphrey, Stephen R. 1974. Zoogeography of the nine-banded armadillo (Dasypus novemcinctus) in the United States. BioScience 24:457-462.

Jenkins, James H. 1953. The game resources of Georgia. Ga. Fish Game Comm., Atlanta. 1 14 pp.

134 Joshua Laerm, et al.

1971. The status and management of the Bobcat and Cougar in the

southeastern United States. Symposium on the native cats in North America. USDI Fish Wildl. Serv. U.S. Govt. Printing Office, Washington, D.C. 27 pp.

Jones, Clyde. 1977 Plecotus rafinesquii. Mamm. Species (59:1-4.

Jones, J. Knox, Jr., D.C. Carter and H.H. Genoways. 1979. Revised checklist of North American mammals north of Mexico, 1979. Occas. Pap. Mus. Texas Tech Univ. (52:1-17.

Laerm, Joshua. 1981. The systematic status of Geomys cumberlandius . Brim- leyana 6:141-151.

, B.J. Freeman, L.J. Vitt, J.M. Meyers and L.E. Logan. 1980. Verte- brates of the Okefenokee Swamp. Brimleyana 4:47-73.

, J.C. Avise, J.C. Patton and R.A. Lansman. In press a. Genetic

determination of an endangered species of pocket gopher in Georgia. J.

Wildl. Manage.

, B.J. Freeman, L.J. Vitt, J.H. Rappole and L.E. Logan. In press b.

The status of non-game vertebrates in Georgia. In R. Odom (ed.). Second

Symposium on Non-game/ Endangered Wildlife. Long, Charles A. 1972. Taxonomic revision of the mammalian genus Micro-

sorex Coues. Trans. Kans. Acad. Sci. 74:181-196. 1974. Microsorex hoyi and Microsorex thompsoni. Mamm. Species

55:1-4. Miller, G.C., Jr., and G.M. Allen. 1928. The American bats of the genera

Myotis and Pizonyx. Bull. U.S. Nat. Mus. 744:1-218. Neuhauser, Hans N. 1972. Subnumerary dentition in the least shrew, Cryptotis

parva (Mammalia: Insectivora). Ga. J. Sci. 30:25-26. 1978. Anastasia Island Cotton Mouse, pp. 45-47 in J.N. Layne (ed.).

Rare and Endangered Biota of Florida. Vol. 1. Mammals. Univ. Presses

Fla., Gainesville. 52 pp. , and W.W. Baker. 1974. Annotated list of mammals of the coastal

islands of Georgia, pp. 197-209 in A.S. Johnson, H.O. Hillestad, S.F. Shanholtzer and F.G. Shanholtzer (eds.). An Ecological Survey of the Coastal Region of Georgia. Nat. Park Serv. Sci. Monogr. Ser. 3. 233 pp. , and C. Ruckdeschel. 1978. Whales (Cetacea) of Georgia, pp. 38-53

in R.R. Odom and L. Landers (eds.). Proceeding of the Rare and Endan- gered Wildlife Symposium. Ga. Dep. Nat. Resour. Game Fish Div. Tech. Bull. WL4. 184 pp.

Odom, Ron R., J.L. McCollum, M.A. Neville and D.R. Ettman. 1977. Georgia's protected wildlife. Ga. Dep. Nat. Resour., Atlanta. 51 pp.

Odum, Eugene P. 1949. Small mammals of the Highlands (North Carolina) plateau. J. Mammal. 50:179-192.

Paradiso, J.L., and R.M. Novak. 1971. A report on the taxonomic status and distribution of the red wolf. USDI Bur. Sport Fish. Wildl. Spec. Sci. Rep. Wildl. 745:1-36.

Peterson, Karen E., and T.L. Yates. 1980. Condylura cristata. Mamm. Species 729:1-4.

Georgia Mammals 135

Richardson, James I. 1973. A confirmed occurrence of the rough-toothed dol- phin (Steno bredanensis) on the Atlantic coast of the United States. J. Mammal. 54:215.

Schmidly, David J. 1981. Marine mammals of the southeastern United States coast and the Gulf of Mexico. U.S. Dep. Inter. FWS/OBS80/41. 165 pp.

Schwartz, Albert, and E.P. Odum. 1957. The woodrats of the eastern United States. J. Mammal. 55:197-206.

Smolen, Michael, J. 1981. Microtus pinetorum. Mamm. Species 147:1-1.

van Zyll de Jong, C.G. 1976. Are there two species of pygmy shrews (Micro- sorex)? Can. Field-Nat. 90:485-487.

1979. Distribution and systematic relationships of longeared Myotis

in western Canada. Can. J. Zool. 57:987-994.

Watkins, Larry C. 1972. Nycticeius humeralis. Mamm. Species 25:1-4.

Wharton, Charles H. 1968a. First Records of Microsorex hoyi and Sorex cine- reus from Georgia. J. Mammal. 49:158.

1968b. Distribution of the red squirrel in Georgia. J. Mammal.

49:153-155.

1793. Natural Environments of Georgia. Ga. Dep. Nat. Resour.

Publ., Atlanta. 227 pp.

, and J.J. White. 1967. The red-backed vole, Clethrionomys gapperi,

in North Georgia. J. Mammal. 48:610-612. Wiley, Robert. 1980. Neotoma flohdana. Mamm. Species 759:1-7. Williams, Steven L., and H.H. Genoways. 1980. Morphological variation in the

southeastern pocket gopher, Geomys pinetis (Mammalia: Rodentia). Ann.

Carnegie Mus. 49:405-453. Yates, Terry L., and D.J. Schmidly. 1978. Scalopus aquaticus. Mamm. Species

705:1-4.

Accepted 16 April 1982

Distributional Records for Gastropods and Sphaeriid Clams

of the Kentucky and Licking River and Tygarts

Creek Drainages, Kentucky

Branley A. Branson and Donald L. Batch

Department of Biological Sciences, Eastern Kentucky University, Richmond, Kentucky 40475

ABSTRACT. — Collections from 50 sites in the Kentucky and Licking River drainages and from Tygarts Creek, direct tributary of the Ohio River in eastern Kentucky, produced records of 2 genera and 7 species of sphaeriid clams, and of 6 families, 12 genera, and 22 species of gastropods. Sphaerium corneum and Lymnaea stagnalis are reported from Kentucky for the first time.

INTRODUCTION

In a recent attempt to generate a list of the Endangered, Threat- ened and Rare plants and animals of Kentucky, the authors and asso- ciates (Branson et al. 1981) were hampered by the relative paucity of published information on aquatic gastropods and sphaeriid clams. With the exception of a long series of papers on pleurocerids by Calvin Goodrich (see citations in Bickel 1967 and Branson 1972), very few pap- ers have dealt with the Kentucky molluscan fauna. There are many broad hiatuses in our knowledge of distribution patterns, particularly in the Kentucky, Licking, and Green River drainages, and in the various streams directly tributary to the Ohio River. The present contribution fills some of the gastropod and sphaeriid clam distributional gaps in the Kentucky and Licking rivers, and in Tygarts Creek, a clear tributary of the Ohio River in eastern Kentucky.

COLLECTING STATIONS

The 50 numbered collecting stations and dates of sampling are given below. SR = State Road, US = U. S. Highway, CR = County Road, and KR = Kentucky Highway.

Kentucky River Drainage

1. Kentucky River, Frankfort, Franklin Co.; 5 November 1980.

2. Kentucky River, Lock and Dam 4, Franklin Co.; 20 October 1980.

3. Kentucky River, 0.6 km downstream from Fort Boonesborough State Park, 12.9 km SW Winchester, Madison Co.; 11 October 1980.

4. Kentucky River at mouth of Benson Creek, just above Lock and Dam 4, Franklin Co.; 20 October 1980.

5. Kentucky River, Lock and Dam 8 at CR 1268, Jessamine Co.; 16 June 1973.

Brimleyana No. 7:137-144. July 1981. 137

138 Branley A. Branson and Donald L. Batch

6. Kentucky River, 12 km NE Frankfort on US 127 and 3.2 km W Steeles Mill Rd., Franklin Co.; 20 October 1980.

7. Middle Fork of Kentucky River at SR 30, 16.1 km SSE Jackson, Breathitt Co.; 1 July 1972.

8. South Fork of Kentucky River, Oneida, Clay Co.; 18 July 1970.

9. South Fork of Kentucky River at SR 1 1, 1.2 km S Clay and Owsley cos. line in Clay Co.; 1 August 1970.

10. South Fork of Kentucky River at SR 11, 0.4 km N Clay Co. line in Owsley Co.; 13 September 1970.

1 1. South Fork of Kentucky River, Booneville, Owsley Co.; 16 January 1971.

12. South Fork of Kentucky River, Eversole, Owsley Co.; 19 June 1971.

13. South Fork of Kentucky River, 0.5 km N Booneville, CR 1475, Owsley Co.; 6 June 1971.

14. Middle Fork of Kentucky River, Tallega, Lee Co.; 24 June 1972.

15. Middle Fork of Kentucky River, Hoskinston, Leslie Co.; 17 November 1973.

16. Dix River at KR 52, Hedgeville, Boyle and Garrard cos.; 1 1 October 1980.

17. Elkhorn Creek at CR 1900, 4.8 km NNE Frankfort, Franklin Co.; 19 October 1980.

18. Elkhorn Creek, Frankfort, Franklin Co.; 19 October 1980.

19. Boone Creek at Grimes Mill Road, 19.2 km N Richmond, Fayette Co.; 2 November 1980.

20. Red River, Hazel Green, Wolfe Co.; 15 October 1980.

21. Small pond, Central Kentucky Wildlife Management Area, 2 km SE Kingston, Madison Co.; 9 September 1980.

22. Silver Creek at Barnsmill Road, 9.6 km W Richmond, Madison Co.; 25 October 1980.

23. Silver Creek at SR 52, 19 km E Richmond, Estill Co.; 16 March 1966.

24. Elkhorn Creek at KR 460, 1.2 km E Frankfort, Franklin Co.; 19 September 1980.

25. Elkhorn Creek at KR 227, 0.9 km W Georgetown, Scott Co.; 11 September 1980.

26. Cave Run at confluence with Elkhorn Creek, 3.5 km W Lexington, Fayette Co.; 10 September 1980.

27. Paint Lick Creek, 0.3 km above mouth of Dry Run Creek, 1.0 km SE Paintlick, Garrard Co.; 8 September 1980.

28. Lake Arlington, 0.25 km N Richmond, Madison Co.; 21 March 1972.

29. Farm pond, Eastern Kentucky University, Richmond, Madison Co.; 23 March 1972.

Kentucky Mollusk Distributions 139

30. Eagle Creek at KR 437, 0.7 km E Sparta, Gallatin Co.; 12 Sep- tember 1979.

31. Sturgeon Creek at confluence with Kentucky River, Heidelberg, Lee Co.; 3 July 1980.

32. Little Sturgeon Creek at confluence with Sturgeon Creek, 3.2 km NW Travellers Rest, Owsley Co.; 22 January 1972.

33. Muddy Creek at Doylesville Rd., 1.6 km SE Doylesville, Madison Co.; 16 September 1972.

34. Sinking Creek at SR 52, 3.6 km E Irvine, Estill Co.; 16 March 1966.

35. Goose Creek at SR 718, 0.5 km N Bright Shade, Clay Co.; 3 March 1970.

36. Red Bird Creek at SR 99, 1.0 km N Creekville, Clay Co.,; 23 May 1970.

37. Red Bird Creek at mouth of Little Double Creek and SR 99, Clay Co.; 30 May 1970.

38. Bullskin Creek at CR 1482, 0.5 km E Oneida, Clay Co.; 6 June 1970.

39. Sexton Creek at confluence with South Fork of Kentucky River, 10.5 km S Booneville, SR 11, Owsley Co.; 27 February 1971.

40. Island Creek at SR 1 1, Conklin, Owsley Co.; 6 March 1971.

41. Buck Creek at confluence with South Fork of Kentucky River, Owsley Co.; 10 April 1971.

42. Bear Branch, Nobel, Breathitt Co.; 26 April 1969.

43. Greasy Creek, 1.3 km E Asher, Leslie Co.; 23 October 1976.

Licking River Drainage

44. Licking River at US 60, 3 km SE Farmers, Bath and Rowan cos.; 23 October 1980.

45. Licking River, Butler, Pendleton Co.; 4 August 1964.

46. North Fork of Licking River at SR 165, 5.0 km N Mount Olivet, Robertson Co.; 12 October 1974.

47. Fish ponds, Minor E. Clark Fish Hatchery, Morehead, Rowan Co.; 6 September 1980.

48. Slate Creek at US 60, 3.2 km E Owingsville, Bath Co.; 10 November 1973.

49. Fox Creek, Grange City, Fleming Co.; 10 November 1973.

Tygarts Creek Drainage

50. Tygarts Creek at SR 7, 8 km NW Greenup, Greenup Co.; 5 October 1968.

ANNOTATED LIST

The following list contains data for 2 genera and 7 species of

140 Branley A. Branson and Donald L. Batch

sphaeriids and 6 families, 12 genera and 22 species of aquatic snails. The number of specimens obtained at each site is provided in parentheses following the collecting site number. All specimens were deposited in the Museum of Zoology, Eastern Kentucky University.

Bivalvia: Sphaeridae

The characters used by Clarke (1973) and Burch (1975) to separate Sphaerium and Musculium demonstrate as many relationships as they do differences. Hence, we follow Herrington (1962) in relegating the genus Musculium to subgeneric status under Sphaerium.

Sphaerium corneum (Linnaeus). Collections: 48 (1). The habitat consisted of fine sand admixed with small quantities of silty materials. Considered an exotic by Herrington (1962), this species heretofore has not been reported from Kentucky waters. A single specimen is not very conclusive evidence, particularly because aberrant specimens of S. ni- tidum Westerlund are easily confused with this species. Additional field work needs to be accomplished in order to ascertain the status of S. corneum in Kentucky.

Sphaerium lacustre (Muller). Collections: 28 (4). Specimens were obtained by dredging. The distribution of this and other pond-dwelling sphaeriids is poorly known in Kentucky, principally because of inade- quate sampling.

Sphaerium fabale Prime. Collections: 1 1 (2), 31 (12), 43 (3), 46 (15), 49 (1). This species is not uncommon in clean upland tributaries and main rivers in relatively shallow situations with an abundance of rocks and gravel. Branson and Batch (1981 and 1982) reported thriving popu- lations of this small clam in Dix and Red rivers, both major tributaries of the Kentucky River.

Sphaerium simile Say. Collections: 8(1), 1 1 (1), 14 (10), 32 (2). This rather heavy-shelled form is not uncommon in the Kentucky River drainage (Branson and Batch 1981), but apparently is scarce in most of the Licking River.

Sphaerium striatinum (Lamarck). Collections: 13 (1), 22 (3), 33 (1), 37 (1), 39 (1), 42 (5), 44 (1), 45 (1), 48 (22), 49 (1), 50 (2). The most widespread and abundant sphaeriid in Kentucky, this species sometimes produces prodigious populations in clean (lacking silt), rocky, vegetated riffles.

Sphaerium transversum (Say). Collections: 29 (6). This species was taken in dredge samples from soft mud in water approximately 2 m deep.

Pisidium compressum Prime. Collections: 28 (1), 29 (3). Both col- lections came from mud-bottom ponds. All members of the genus Pi- sidium are poorly known in Kentucky because of a dearth of collecting.

Kentucky Mollusk Distributions 141

Gastropoda: Prosobranchia

Many taxonomic problems still exist among the Pleuroceridae, problems that must be resolved before there is any stability in the family at the species level. Not all of these problems can be resolved by means of shell morphology. Instead, investigators must rely on data from reproductive behavior, particularly the method of egg deposition, and from newer techniques such as electrophoresis. In particular, there is a plethora of problems needing resolution in the Cumberland and Green rivers of Kentucky and adjacent Tennessee. To complicate matters, sev- eral of the species are becoming threatened as we extend domination over progressively larger and larger sections of those streams (Branson et al. 1981). However, in spite of these problems, most of the genera treated in Goodrich (1940) and elsewhere, with some exceptions, are relatively stable and recognizable categories. The generic designations used herein are principally those of Goodrich (1940).

Pleuroceridae

Lithasis obovata (Say). Collections: 1 (1), 2 (10), 27 (2), 45 (1). Not a common species in general riverine collections from Kentucky, this snail is listed as of Special Concern (Branson et al. 1981). There is an apparently healthy population in the pool behind Lock 4 (our Station 2) on the Kentucky River. There is a move by the U.S. Army Corps of Engineers to abandon management and operation of the lock and dam system, although some of the county governments along the river have proposed to take over operating some of the individual locks. These habitats may thus eventually become excellent areas for mollusks and small fishes.

Lithasia plicata Wetherby. Collections: 39 (1). Although not listed as Threatened or Endangered, this species is rare in Kentucky waters and probably deserves such consideration. It cannot withstand settleable- solid or acid-mine pollution. Unfortunately, most of the known Ken- tucky habitats lie in the middle of the Eastern Coal Fields.

Pleurocera canaliculatum (Say). Collections: 2 (5), 4 (1), 5 (1), 10 (1), 24 (1), 45 (11). Most of these specimens are of the morphological type listed under the subspecific designation P. c. undulatum (Say), except those from the Licking River. The population at Station 4 is large.

Pleurocera acuta Rafinesque. Collections: 7(1), 17 (1), 18 (1), 31 (2), 33 (2), 39 (1), 40 (21), 50 (2). Pleurocera acuta is listed as of Special Concern (Branson et al. 1981), although there is a healthy population at Station 45 in the Licking River. The species has nearly disappeared from localities where it was abundant (see Goodrich 1940; Call 1900).

Nitocris trilineata (Say). Collections: 45 (5). Since we made this

142 Branley A. Branson and Donald L. Batch

collection (1964) from the Licking River at Butler, a dam has started releasing very cold water through the habitat. In the spring of 1981 we were unable to find a single specimen of this handsome little snail at the locality. The species is listed as Threatened (Branson et al. 1981).

Goniobasis semicarinata (Say). Collections: 3 (7), 8 (17), 9 (2), 10 (31), 15 (31), 17 (14), 22 (32), 23 (12), 26 (30), 27 (8), 33 (2), 34 (45), 37 (1), 38 (9), 40 (1), 42 (7), 43 (49), 44 (1), 45 (51), 46 (21), 48 (35), 49 (1), 50 (6). This is the characteristic pleurocerid of the entire Kentucky and Licking River drainages, as well as of the Upper Salt River Drainage. Although not formerly reported from the Cumberland River drainage, there are many populations of pleurocerids there (particularly in the Little South Fork and the Rockcastle River) that are not separable by means of shell characteristics from those of the Licking and Kentucky rivers.

Goniobasis costifera (Haldeman). Collections: 11 (13), 12 (9), 13 (15), 14 (1), 25 (24), 36 (1), 41 (3). This species is only locally abundant in the Kentucky River drainage (Branson and Batch 1981).

Viviparidae

The distribution of viviparids is extremely poorly known in Ken- tucky, and practically nothing has been added since the papers of Clench (1962a, b), Clench and Turner (1955), and Clench and Fuller (1965). Two genera and three species are reported here.

Lioplax subcarinata occidentalis Pilsbry. Collections: 30 (1), 45 (1). The habitat at both sites was mud and rocks with rooted vegetation. Clench and Turner (1955) considered this a synonym of L. sulculosa (Menke).

Campeloma integrum (Say). Collections: 8 (3), 45 (1). This species was taken from mud at the base of water willow along the margins of riffles.

Campeloma crassula Rafinesque. Collections: 11 (1), 17 (3), 24 (1), 48 (1). Some of the older literature reported this species as C. ponderosa (Say).

Hydrobiidae

Pomatiopsis lapidaria(Say). Collections: 3 (1), 6 (1), 16 (2), 19 (1), 25 (3). Although most authors consider this species amphibious, the specimens from Station 16 were removed from dead leaves in the water. The others were found at the water's edge on wet soil and rocks.

Pulmonata: Basommatophora

Other than the exceptions noted below, we basically follow the classification scheme of Taylor and Sohl in the Basommatophora.

Lymnaeidae

We follow the rationale of Hubendick (1951, 1978) in our treatment

Kentucky Mollusk Distributions 143

of this family of snails. Distributional data for lymnaeids in Kentucky are sparse to lacking, particularly in the western half of the state.

Lymnaea columella Say. Collections: 21 (4), 44 (4). The specimens were taken from backwaters over mud bottoms, and most of them were heavily laden with fluke larvae.

Lymnaea palustris (Muller). Collections: 28 (3). Although the occur- rence of this pond snail in Kentucky was implied by Baker (1911), this is the first record of its occurrence supported by definite locality data.

Lymnaea stagnalis Linnaeus. Collections: 28 (5). The pond from which these specimens were taken supports a large community of cool- water vegetation and attracts many migratory aquatic birds, which may account for the rather speciose snail population at this station. Aquatic snails, particularly young specimens, are easily transported in mud and plant debris on the feet of these birds. This species was heretofore unknown from Kentucky.

Lymnaea humilis Say. Collections: 3 (1), 28 (1), 29 (1). This is by far the most commonly encountered pond snail in Kentucky. The spec- imen from Station 3 is of the obrussa form.

Physidae

This family is badly in need of comprehensive monographic treat- ment. Many of the so-called species are doubtless ecotypes, and still other species probably await discovery by means of electrophoretic and other biochemical techniques. Identifications based on shell features alone, such as the three species reported here, are only tentative.

Physa heterostropha (Say). Collections: 22 (1), 28 (2).

Physa gyrina Say. Collections: 32 (5).

Physa Integra Haldeman. Collections: 22 (3), 25 (3), 29 (23).

Ancyloplanorbidae

We follow Hubendick (1978) in assigning this family name, which includes the Planorbidae and the Ancylidae. Bulininae: Physastrini Ferrissia rivularis (Say). Collections: 44 (3), 45 (1). The specimens were removed from dead unionid valves in deep, swift riffles. Bulininae: Camptoceratini Helisoma anceps (Menke). Collections: 15 (1), 17 (2), 20 (6), 28 (2), 29 (17), 42 (1), 43 (1), 50 (1). In the eastern highlands of Kentucky, Helisoma anceps appears to be more common and abundant than the following species.

Helisoma trivolvis (Say). Collections: 21 (5), 22 (1), 25 (1), 28 (1), 29 (12), 47 (7), 48 (2). The principal habitats of this widespread species are lowland, mostly in vegetated backwaters over mud bottoms. Planorbinae: Planorbini

Gyraulus parvus (Say). Collections: 29 (1). There are very few

144 Branley A. Branson and Donald L. Batch

published records for this and other small planorbid species, doubtless because of inadequate collecting in vegetated standing waters.

LITERATURE CITED

Baker, Frank C. 1911. The Lymnaeidae of North and Middle America. Chic.

Acad. Sci. Spec. Publ. 3. 539 pp. Bickel, David. 1967. Preliminary checklist of Recent and Pleistocene Mol-

lusca of Kentucky. Sterkiana 28:1-20. Branson, Branley A. 1972. Checklist and distribution of Kentucky aquatic

gastropods. Ky. Fish. Bull. 54:1-20. , and D. L. Batch. 1981. The gastropods and sphaeriacean clams of the

Dix River system, Kentucky. Trans. Ky. Acad. Sci. 42:54-61. , and 1982. The Gastropoda and sphaeriacean clams of Red

River, Kentucky. Veliger 24:200-204. , D. F. Harker, Jr., J. M. Baskin, M. E. Medley, D. L. Batch, M. L.

Warren, Jr., W. H. Davis, W. C. Houtcooper, B. Monroe, Jr., L. R. Phil- lippe and P. Cupp. 1981. Endangered, Threatened, and Rare animals and plants of Kentucky. Trans. Ky. Acad. Sci. 42:77-89.

Burch, John B. 1975. Freshwater sphaeriacean clams (Mollusca: Pelecypoda) of North America. Malacol. Publ., Hamburg, MI. xii + 95 pp.

Call, Robert E. 1900. A descriptive catalogue of the Mollusca of Kentucky.

Clarke, Arthur H. 1973. The freshwater molluscs of the Canadian Interior Basin. l:165-183.Malacologia 75:1-509.

Clench, William J. 1962a. New records for the genus Lioplax. Occas. Pap. Mollusks Mus. Comp. Zool. Harv. Univ. 2:288.

1962b. A catalogue of the Viviparidae of North America with

notes on the distribution of Viviparus georgianus Lea. Occas. Pap. Mol- lusks Mus. Comp. Zool. Harv. Univ. 2:1-20.

, and S. L. H. Fuller. 1965. The genus Viviparus in North America

Occas. Pap. Mollusks Mus. Comp. Zool. Harv. Univ. 2:385-412 , and R. D. Turner. 1955. The North American genus Lioplax in the

family Viviparidae. Occas. Pap. Mollusks Mus. Comp. Zool. Harvard

Univ. 2:1-20. Goodrich, Calvin. 1940. The Pleuroceridae of the Ohio River drainage. Occas.

Pap. Mus. Zool. Univ. Mich. 417:1-21. Herrington, H. B. 1962. A revision of the Sphaeriidae of North America

(Mollusca: Pelecypoda). Misc. Publ. Mus. Zool. Univ. Mich. 118:1-81. Hubendick, Bengt. 1951. Recent Lymnaeidae. Kungl. Svenskavetenska. Handl.

Bd. 3:1-222. 1978. Systematics and comparative morphology of the Basommato-

phora. pp. 1-47 in V. Fretter and J. Peake (eds.). Pulmonates. Academic

Press, N.Y. 540 pp. Taylor, Dwight W., and N. F. Sohl. An outline of gastropod classification.

Malacologia 7:7-32.

Accepted 1 December 1981

Rediscovery and Distribution of Bembidion plagiatum Zimmermann (Coleoptera: Carabidae)

Richard L. Hoffman

Department of Biology, Radford University, Radford, Virginia 24142

ABSTRACT. — Bembidion plagiatum Zimmermann, heretofore consid- ered one of the scarcest American members of the genus, is reported from new localities in New Jersey, Pennsylvania, Virginia, and North Carolina. At the two known Virginia localities the species seems to prefer silt-coated sandbars up to several meters distant from the stream side, a habitat not shared with other bembidiids. Collection dates sug- gest that the species is active from April to early July; adults have not been found after that time. Both external details and structure of the penis suggest that plagiatum is closely related to B. lacunarium rather than the species with which Lindroth (1963) associated it on the basis of color pattern.

At the time Prof. C. H. Lindroth revised the bembidiid fauna of boreal North America (1963) and re-established Bembidion plagiatum as a valid species, he had seen only two specimens: the original type from "Maryland" and a female that he collected at Long Point, Ontario. Other specimens must have been found in between these two, however, to judge from Hay ward's remark (1897:82) that "Specimens with a sub- marginal pale spot {plagiatum Zimm.) bear some resemblance to scopul- inum . . . "; nonetheless Lindroth quite justifiably stated that plagiatum was "Apparently strictly eastern and very rare.".

Despite a renaissance of interest in American bembidiids in recent years, I am not aware of any subsequent discoveries of plagiatum. Hav- ing had the good fortune to recently find this species in Virginia (as well as in museum collections and other people's notes), I feel that some remarks on its apparent favored biotope and the enumeration of new localities will be of interest to carabidologists.

The first specimen that came to my attention entered through the back door, so to say, as the result of my curiosity about the Southern Pines, North Carolina, record for B. scopulinwn Kirby cited in Brim- ley's Insects of North Carolina (1938:117). That this northern species would occur naturally at Southern Pines seemed utterly implausible so I obtained on loan all of Brimley's Bembidion material for a personal examination. Although the tray labeled scopulinum does contain a spec- imen of that species from New Hampshire, the single female from

Brimleyana No. 7:145-150. July 1981. 145

146 Richard L. Hoffman

Southern Pines (A. H. Manee, leg.) lacks the coarse temporal puncta- tion of scopulinum and keys out readily to plagiatum in Lindroth's synopsis.

In June, 1969, Dr. Thomas C. Barr, Jr. collected a single male of plagiatum on Blackrock Creek in Horse Cove, Transylvania County, North Carolina, and this record in connection with those for Southern Pines and "Maryland" rendered the eventual discovery of plagiatum in Virginia almost certain.

The first known Virginia specimen was a female obtained amongst a variety of common bembidiids along Cobb's Branch, a tributary to Smith River (about 2 km northwest of Irisburg on Virginia Highway 750), Henry County, Virginia, on July 14, 1980. These beetles were not identified until a week later, so that the immediate return visit to the site occurred only near the end of July. It did not produce any further spec- imens of plagiatum, nor did subsequent spot-checks made even later in the summer. However, sampling on April 18, 1981, yielded an adult male, and two females were found on May 2, 1981. The yellow elytral spots were conspicuous enough that the species could be recognized in the field without magnification, and it was possible to associate individ- uals with their precise biotope. On May 28, 1981, another male was found along the Sandy River (upstream of its crossing by Virginia Highway 855), Pittsylvania County, about 14 km northeast of the Cobb's Branch locality. Sampling at both these localities later in the summer of 1981 produced no additional specimens.

Two additional new localities came to my attention serendipitously. In reporting my Virginia finds to Dr. Terry Erwin, he recalled having located a specimen in the California Academy of Sciences collected at Phillipsburg, New Jersey, June 25, 1915, by J. W. Green. While examin- ing the type specimen of plagiatum at Harvard on my behalf, Dr. A. F. Newton, Jr. located an individual — overlooked by Lindroth — in the Horn Collection labeled only "Allegheny, Pa." and identified as this species in Horn's handwriting. The original hamlet of Allegheny no longer occurs on most maps, the place having long since been consumed in the urban spread of Pittsburgh.

At the two Virginia localities, plagiatum was found only on gravel- sand bars with i surface coating of fine damp silt, about 0.5 to 1 m above water le ?' and 2 or 3 m removed from the edge. The only asso- ciated member v the genus here was B. inaequale but the biotope was shared by thf tachyine Elaphropus vivax (LeConte) and the staphylinids Geodromicus brunneus Say, Homaeotarsus bicolor (Gravenhorst), Phi- lonthus sp., Scopaeus sp., and Lissobiops serpentinum (LeConte). Along

Bembidion Distribution

147

the water's edge B. nigrum (Cobb's Branch) and B. honestum (Sandy River) were extremely abundant. The sandbars were at the time devoid of vegetation and were exposed to full sun at one place and afternoon insolation at the other. Both streams were about 2 to 3 m wide and entrenched in sandy floodplain at both sites.

Dr. Barr informed me (in litt.) that his specimen from Horse Cove was likewise found on a sandbar, near the mouth of Blackrock Creek, and that repeated visits during the summer of 1969 failed to produce another. This observation parallels my own lack of success at Cobb's Branch after mid-July. Perhaps the species is active as an adult only during spring and early summer, at least in the south. Manee's capture at Southern Pines, North Carolina, was made in April. Lindroth's Ontario specimen was found June 7, 1956. By late July the sandbars at the two Virginia sites were considerably grown up in rank weedy vegeta- tion that considerably altered the former appearance and doubtless also affected the microhabitat conditions as well.

Examination of my material revealed an interesting structural fea- ture perhaps diagnostic of this species. As shown by Figure 1 the 1st elytral interval (sutural) is provided with one or two adventitious setae near the apical end, in all four specimens. Dr. Newton kindly examined the male holotype at Harvard and reported one such seta on each side,

Fig. 1. Posterior end of abdomen of specimen of Bembidion plagiatum (Henry County, Virginia) showing accessory setae on 1st elytral intervals apparently characteristic of this species.

148 Richard L. Hoffman

slightly posterior to the posititions shown in my drawing.

In most other respects, this species is extremely similar in body form and minute details of structure to B. lacunarium, which was also named by Zimmermann at the same time as plagiatum. Aside from the setae just noted, the obvious external difference evident to me is the apical yellow elytral spot, which appears to be the result of local trans- lucence of the integument that allows the folded wing tips to show through, and not the reflection of yellow pigment per se.

Contrary to Lindroth's remark that "It is difficult to understand how this species could be regarded as a synonym (spotted form) of lacunarium ..." I can see every justification for postulating a close rela- tionship. Lindroth placed plagiatum in his striola group, so far as I can perceive, solely on the basis of elytral spots, against the more solid evi- dence of both body form and penial structure. His own drawings (Figs. 155f, 159f) show that the internal armature of the penis sac allies plagia- tum with lacunarium at the same time it shows these two species to be disjunct in the groups to which he assigned them. Perhaps both, along with B. texanum, merit recognition in a separate species group.

Several seasons of fairly intensive field work in southwestern Virgin- ia have failed to disclose plagiatum in the mountains, where it seems to be replaced by lacunarium. The collective localities now known suggest a wide distribution at low to moderate elevations, extending from New Jersey south to North Carolina, east of the Blue Ridge, and then, pre- sumably, northward on the west side of the Appalachians as far as Lake Ontario. The map (Fig. 2) graphically represents this apparent "Upper Austral" distribution.

ACKNOWLEDGMENTS.— C. S. Brimley's material of Bembidion was kindly loaned by Mr. James E. Greene, N. C. Department of Agri- culture, Raleigh; Dr. Alfred F. Newton, Jr., Museum of Comparative Zoology, examined the type specimen of plagiatum for me and also identified the staphylinids taken at Cobb's Branch; Dr. Thomas C. Barr, Jr., University of Kentucky, and Dr. Terry L. Erwin, National Museum of Natural History, generously permitted me to report unpublished information from their files. Mr. Robert Davidson, Carnegie Museum, read an early draft of the manuscript and provided information about the location of "Allegheny, Pa.". Roberta R. Hoffman provided diligent and skillful assistance on collecting trips. I am very much indebted to all of these persons for their contributions to the knowledge of an interest- ing and generally overlooked ground beetle.

Bembidion Distribution

149

Fig. 2. Known localities for Bembidion plagiatum in eastern North America. The spot for Maryland is arbitrarily centered in the state as the specimen it represents lacks precise data.

150 Richard L. Hoffman

LITERATURE CITED

Brimley, C. S. 1938. The Insects of North Carolina. N. C. Dep. Agric. Div.

Entomol., Raleigh. 560 pp. Hayward, Roland. 1897. On the species of Bembidium of America north of

Mexico. Trans. Am. Entomol. Soc. (Phila.) 24:32-158 Lindroth, Carl H. 1963. The ground-beetles (Carabidae, excl. Cicindelinae) of

Canada and Alaska. Opusc. Entomol. Suppl. 24:201-408.

Accepted 16 October 1981

Incisor Malocclusion in a Specimen of Sylvilagus floridanus (Mammalia: Lagomorpha) from North Carolina

Gary W. Woodyard

Wayne Community College, Goldsboro, North Carolina 27530

ABSTRACT. — A cottontail rabbit with incisor malocclusion was examined, but no clear reason for the unusual dental condition was found. This is only the second report of the phenomenon in wild Sylvi- lagus floridanus; most such reports deal with domestic or laboratory animals.

Malocclusion, with the consequential overgrowth of incisors, pre- molars or molars, has been reported in rodents and, to a much lesser extent, in lagomorphs. Most reports concerning lagomorph malocclu- sion deal with domestic or laboratory animals (Zeman and Fielder 1969). Thorpe (1930), Lincoln (1938), and others, reported malocclusion and overgrowth of incisors in Marmota. In 1968 I collected but have not previously reported a Scuirus carolinensis with incisor malocclusion and hypertrophied incisors. The only prior published report of incisor malocclusion in a wild cottontail rabbit was that of Gregory (1952).

A female Sylvilagus floridanus, apparently a young-of-the year, was brought from the Seven Springs area of Wayne County, North Carolina, to Wayne Community College for examination because of its unusual dental condition (Fig. 1). It weighed approximately 1 kilogram and had a remarkable amount of body and kidney fat considering the severity of its incisor condition. The lower left incisor extended 1.8 cm beyond the mandible, where it interesected the lower right incisor. The tip of the left incisor was rounded and smooth and the tooth appeared to be correctly positioned relative to the symphysis of the mandibles. Although the incisor rested against the upper lip with the mouth closed, no skin irritation was evident. The lower right incisor extended 2.5 cm beyond the mandible and curved to the left to a point 0.3 cm from the symphysis of the mandibles. Its tip was rounded and rested against the lower part of the left nostril when the mouth was closed, but there was no apparent irritation to the nostril. There was a slight notch approxi- mately 0.7 cm from the tip of the right incisor, caused by the intersec- tion of the lower left incisor.

The upper left first incisor extended 1.4 cm from the premaxillary and had grown into a short arc. Its tip appeared to have been broken anteriorly at an angle of approximately 20 degrees sometime prior to the animal's death. The smaller incisor was of equal length and lay in a

Brimleyana No. 7:151-153. July 1981. 151

152

Gary W. Woodyard

Fig. 1. Maloccluded incisors in a cottontail rabbit, Sylvilagus floridanus.

right lateral position to the larger one. The tips of these incisors were 0.4 cm to the left of the symphysis of the premaxillary. The upper right first incisor extended 3.4 cm from the premaxillary and had grown in an arc of approximately 320 degrees. The tip of the incisor, smooth and rounded, was offset 1.2 cm to the right of the premaxillary symphysis. The smaller incisor extended 0.2 cm from the premaxillary and appeared to have been broken off prior to the animal's death. This tooth lay in its normal position.

The premolars and molars appeared normal, with two exceptions. The first molar of the lower left jaw was reduced to a splinter approxi-

Incisor Malocclusion in Sylvilagus 153

mately 0.15 cm in diameter. The alveolus appeared normal size and was filled with soft, cartilagelike tissue. At the bottom of the alveolus, on the lingual side of the jaw, a small pit opened into the mouth cavity. The second molar showed excessive wear on the posterior cusp; the anterior cusp measured 0.2 cm in height from the maxillary, and the posterior cusp 0.1 cm.

Although the cause of these dental deformities was not clearly evi- dent, the good physical condition of the rabbit suggested recent occur- rence. Gregory (1952) postulated that the malocclusion he described may have been caused by an infection that resulted after a possible for- cible extraction of the right lower incisor. The condition of the lower left first and second molars of my specimen suggests that this area caused misalignment of the jaws and the resultant hypertrophied inci- sors. There was no evidence of any fractured or malformed bones, except for the eroded area found at the base of the first left lower molar.

LITERATURE CITED

Gregory, Joseph T. 1952. Incisor malocclusion in a cottontail rabbit. J. Mam- mal. 34:394.

Lincoln, A., Jr. 1938. Malocclusion in a woodchuck skull. J. Mammal. 79:107.

Thorpe, Malcolm R. 1930. A remarkable woodchuck skull. J. Mammal. 77:69-70.

Zeman, W. V., and F. G. Fielder. 1969. Dental malocclusion and over- growth in rabbits. J. Am. Vet. Med. Assoc. 755:1 1 15-1 1 19.

Accepted 31 October 1981

A Survey of the Freshwater Mussels (Mollusca: Unionidae)

of Middle Island Creek, West Virignia

Ralph W. Taylor and Beverly D. Spurlock

Department of Biological Sciences, Marshall University, Huntington, West Virginia 25701

ABSTRACT. — Twenty-two native species of freshwater mussels plus the introduced Asiatic clam, Corbicula fluminea, were collected from six stations along the seventy-five mile course of Middle Island Creek, a tributary of the Ohio River in West Virginia. Eight species are reported from this creek for the first time. The occurrence of Villosa fabalis, Pleurobema clava, and Alasmidonta marginata is significant because of the rarity of these species within the state. The more com- monly found species include Lampsilis radiata luteola, Lampsilis ven- tricosa, Elliptio dilatata, Amblema plicata plica ta, and Tritogonia verrucosa.

INTRODUCTION

As a result of pollution, impoundment, clear-cutting, and other fac- tors, many of North America's larger streams have experienced severe reductions in plant and animal diversity (Starrett 1971). Freshwater mussels are among the species that have been most seriously affected by habitat changes in "such streams. In numbers of animals and species composition, current mussel populations and communities in most major waterways no longer resemble those that were present at the turn of the twentieth century (Taylor 1980a). Smaller streams have not been altered quite so severely (Taylor 1980b,c). Thus, since it may already be too late to save the mussel faunas of large rivers, a logical conservation approach might be to concentrate on protecting smaller streams in which populations persist and that may serve as refugia for mussels. Later, if water quality in larger waterways can be sufficiently improved, the smaller tributaries could provide mussels with which to restock the major systems.

A first step in implementing a plan of this nature must be to thor- oughly document the native mussel species that inhabit tributaries of major rivers. Only by such studies can the baseline data be obtained that will allow future investigators to monitor trends in mussel popula- tions and communities. Toward this end, we conducted surveys of the mussel fauna of Middle Island Creek, a small tributary of the Ohio River in West Virginia. Mussel populations in the Ohio have declined drastically in numbers of individuals and diversity over the last seventy- five years (Taylor 1980a).

Brimleyana No. 7:155-158. July 1981. 155

156 Ralph W. Taylor and Beverly D. Spurlock

Middle Island Creek flows for approximately 120 km (75 mi.) through the rolling hills of Doddridge, Tyler, and Pleasants counties, and confluences with the Ohio River (RM 154) at St. Marys, Pleasants County. Since there are only a few towns, little industry, and no agricul- ture in these areas, most of the creek's watershed is relatively undis- turbed. Hillside vegetation along the creek may generally be classified as second growth mixed mesophytic forest, 75 to 100 years old.

No extensive survey of the mussels of Middle Island Creek has been previously conducted. The first malacologist to visit the creek (ca. 1900) was A. E. Ortmann of the Carnegie Museum, who listed two spe- cies from there (1919). Bates (1971) reported eight species from two localities on the creek.

METHODS AND PROCEDURES

During October 1980, six sites along Middle Island Creek were extensively sampled for freshwater mussels. Specimens were hand- picked from banks and shallow water. Representative specimens were acces- sioned into the Marshall University Malacological Collections, and additional specimens were placed with the Ohio State University Museum of Zoology. Material from Middle Island Creek taken in ear- lier studies is housed at the Ohio State University Museum of Zoology (OSU); Museum of Comparative Zoology, Harvard University (MCZ); and Carnegie Museum of Natural History (CM).

Species names are those used by Stansbery (1971).

Sampling Sites

In the following list, SR = state route and CR = county road. All sites were on Middle Island Creek. Doddridge County:

1. West Union, approx. 200 m above Main St. bridge at SR18 and old route 50.

Tyler County:

2. 11.3 km N of West Union on SR18.

3. intersection of SR18 and SR74, 20 km N of West Union.

4. SR18 at "The Jug," 3.3 km SE of Middlebourne.

5. bridge near southern boundary of Middlebourne, SRI 8 at CR26.

6. 1.3 km W of Little, along CR14.

RESULTS

Twenty-two species of freshwater mussels were collected in Middle

Island Creek (Table 1). The Asiatic clam, Corbicula fluminea, was

found at all collecting stations. All specimens of mussels housed at the

previously mentioned museums were collected at least fifty years ago.

Freshwater Mussels Middle Island Creek 157

Additional data on the museum specimens are included in Table 1. Over two hundred specimens were collected during this study.

DISCUSSION

Middle Island Creek apparently is a stream of good water quality. The large number of mussel species and individuals indicate thriving populations. Eight species were newly found in the creek, and all species previously reported (Ortmann 1919; Bates 1971) were still present. Most of the species are very common and widespread throughout the upper Ohio River basin, but Villosa fabalis (Lea, 1831) and Pleurobema clava

Table 1. List of mussel species of Middle Island Creek, West Virginia, and sam- pling sites where each occurred. Museums housing historical specimens are indi- cated in parentheses. Asterisk (*) indicates first time collected in Middle Island Creek.

Species Sampling site

12 3 4 5 6

Anodonta grandis grandis Say (OSU) X X

Strophitus undulatus undulatus (Say) (MCZ) X X

Alasmidonta marginata Say*

Simpsonaias ambigua (Say)* X

Lasmigona complanata (Barnes) (OSU, CM) X

Lasmigona costata (Rafinesque) (OSU)

Tritogonia verrucosa (Rafinesque) (OSU)

Quadrula quadrula (Rafinesque) (OSU, CM)

Quadrula pustulosa pustulosa (Lea)*

Amblema plicata plicata (Say) (OSU)

Fusconaia flava (Rafinesque) (MCZ, OSU)

Pleurobema clava (Lamarck)*

Elliptio dilatata (Rafinesque) (OSU)

Ptychobranchus fasciolaris (Rafinesque) (OSU)

Actinonaias ligamentina carinata (Barnes)*

Obovaria subrotunda (Rafinesque)

Villosa fabalis (Lea)*

Villosa iris iris (Lea)*

Lampsilis radiata luteola (Lamarck) (MCZ)

Lampsilis ventricosa (Barnes) (MCZ)

Lampsilis fasciola Rafinesque (OSU)

Epioblasma triquetra (Rafinesque)*

				X
	X	X	X X	X X X
X	X	X	X	X
X	X X	X		X
X	X	X	X	X
X	X X
X	X X X	X	X
X	X	X	X	X
X	X X	X	X X	X

158 Ralph W. Taylor and Beverly D. Spurlock

(Lamarck, 1819) are presently considered rare (Stansbery 1971). Alas- midonta marginata Say, 1818, normally is not found in the western half of West Virginia. The more commonly found species in Middle Island Creek include Lampsilis radiata luteola (Lamarck, 1819), Lampsilis ven- tricosa (Barnes, 1823), Elliptio dilatata (Rafinesque, 1820), and Trito- gonia verrucosa (Rafinesque, 1820).

ACKNOWLEDGMENTS.— We thank David H. Stansbery for con- firming identification of some of our specimens. An anonymous reviewer also provided helpful comments on the manuscript.

LITERATURE CITED Bates, John M. 1971. Mussel investigations State of West Virginia, Part I. Cent.

Aquatic Biol., Eastern Michigan Univ., Ypsilanti. 91 pp. Ortmann, A. E. 1919. A Monograph of the Naiads of Pennsylvania. Ann.

Carnegie Mus. VIII. 384 pp. Stansbery, David H. 1971. Rare and endangered mollusks in eastern United

States, pp. 5-18 in S. E. Jorgensen and R. W. Sharp (eds.). Proceedings of

a Symposium on Rare and Endangered Mollusks (Naiads) of the U. S. Bur.

Sport Fish. Wildl., Twin Cities, MN. 79 pp. Starrett, William C. 1971. A survey of the mussels (Unionacea) of the Illinois

River: A polluted stream. 111. Nat. Hist. Surv. Bull. 30(5): 267-403. Taylor, Ralph W. 1980a. A survey of the freshwater mussels of the Ohio River

from Greenup Locks and Dam to Pittsburgh, PA. Huntington/ Pittsburgh

Districts, U. S. Army Corps of Engineers. 71 pp. 1980b. Freshwater bivalves of Tygart Creek, northeastern Kentucky.

Nautilus 94(2):89-91. 1980c. Mussels of Floyd's Fork, a small northcentral Kentucky

stream (Unionidae). Nautilus 94(\):\3-\5.

Accepted 15 April 1982

159 SUBSCRIPTIONS AND EXCHANGES

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Address all subscriptions and requests for information on purchase and exchange to Managing Editor, Brimleyana, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. Back issues are available for $4.50 each.

DATE OF MAILING

Brimleyana No. 6 was mailed on 16 April 1982.

ERRATA

The following errors appeared in Brimleyana No. 6:

Page 10, Notropis petersoni account, line 3: change at to a^.

Page 21, paragraph 2, line 1 1 should read: "This may explain absence of . . . ."

160

MANUSCRIPT REVIEWERS

The editor and editorial staff are indebted to the following biolo- gists who kindly reviewed manuscripts published or submitted for pub- lication in Brimleyana Nos. 3 through 6 (1980 and 1981):

Ronn Altig, Mississippi State University

Thomas C. Barr, Jr., University of Kentucky

Joseph A. Beatty, Southern Illinois University at Carbondale

Arthur C. Benke, Georgia Institute of Technology

E. L. Bousfield, National Museums of Canada

Branley A. Branson, Eastern Kentucky University

Alvin L. Braswell, North Carolina State Museum

E. E. Brown, Davidson, North Carolina

Brooks M. Burr, Southern Illinois University at Carbondale

Archie F. Carr, University of Florida

Bruce B. Collette, National Marine Fisheries Service NOAA

Roger Conant, University of New Mexico

David C. Culver, Northwestern University

Raymond D. Dueser, University of Virginia

James L. Dobie, Auburn University

William R. Elliott, Texas Tech University

Kenneth M. Fahey, Auburn University

Dorothea D. Franzen, Illinois Wesleyan University

John B. Funderburg, North Carolina State Museum

L. L. Gaddy, Walhalla, South Carolina

Hugh H. Genoways, Carnegie Museum of Natural History

J. Whitfield Gibbons, Savannah River Ecology Laboratory

Carter R. Gilbert, Florida State Museum

Charles O. Handley, Jr., National Museum of Natural History

Julian R. Harrison, III, College of Charleston

Marvin M. Hensley, Michigan State University

Richard L. Hoffman, Radford University

John R. Holsinger, Old Dominion University

Francis G. Howarth, Bernice P. Bishop Museum

Dale R. Jackson, University of South Florida

Robert Jaeger, University of Southwestern Louisiana

Robert E. Jenkins, Roanoke College

J. Eric Juterbock, Ohio State University at Lima

Eugene P. Keferl, Brunswick Junior College

Howard I. Kochman, National Fish and Wildlife Laboratory,

Gainesville Christopher P. Kofron, Texas A&M University Robert A. Kuehne, University of Kentucky

161

Joshua Laerm, University of Georgia

David S. Lee, North Carolina State Museum

Alicia T. Linzey, Blacksburg, Virginia

Donald W. Linzey, Virginia Polytechnic Institute and State

University Charles S. Manooch, National Marine Fisheries Service NOAA

C. J. McCoy, Carnegie Museum of Natural History Robert H. Mount, Auburn University

William B. Muchmore, University of Rochester

Helmut C. Mueller, University of North Carolina at Chapel Hill

Henry R. Mushinsky, University of South Florida

Jerry W. Nagel, East Tennessee State University

Wilfred T. Neill, New Port Richey, Florida

John F. Pagels, Virginia Commonwealth University

William M. Palmer, North Carolina State Museum

John L. Paradiso, U. S. Fish and Wildlife Service

Patricia Parker, University of North Carolina at Chapel Hill

Norman I. Platnick, American Museum of Natural History

Thomas L. Poulson, University of Illinois at Chicago Circle

Peter W. Price, Northern Arizona University

John S. Ramsey, Auburn University

Clayton E. Ray, National Museum of Natural History

Robert K. Rose, Old Dominion University

Duane A. Schlitter, Carnegie Museum of Natural History

Frank J. Schwartz, UNC Institute of Marine Sciences

Raymond D. Semlitsch, Savannah River Ecology Laboratory

William A. Shear, Hampden-Sydney College

Rowland M. Shelley, North Carolina State Museum

Stephen E. Stancyk, University of South Carolina

Jon D. Standing, University of California at Berkeley

Wayne C. Starnes, University of Tennessee at Knoxville

D. L. Stoneburner, University of Georgia Stephen G. Tilley, Smith College

Walter R. Tschinkel, Florida State University

Priscilla Tucker, Texas A&M University

Amy S. VanDevender, Boone, North Carolina

Laurie J. Vitt, University of Georgia

Marvalee H. Wake, University of California at Berkeley

S. David Webb, Florida State Museum

Frederick Whittaker, University of Louisville

Gerald K. Williamson, Savannah Science Museum

Lawrence A. Wilson, Clemson University

Ralph W. Yerger, Florida State University

162

TABLE OF CONTENTS

1981

Number 5

Anderson, Elaine (see Ray, Clayton E.) 1

Andre, John B. Habitat Use and Relative Abundance of the Small

Mammals of a South Carolina Barrier Island 129

Barr, Thomas C, Jr. Pseudanophthalmus from Appalachian Caves (Coleoptera: Carabidae): The Engelhardti Complex 37

Blem, Charles R. Reproduction of the Eastern Cottonmouth Agkistrodon piscivorus (Serpentes: Viperidae) at the Northern Edge of its Range 117

Gibbons, J. Whitfield and Julian R. Harrison, III. Reptiles and Amphib- ians of Kiawah and Capers Islands, South Carolina 145

Harrison, Julian R., Ill (see Gibbons, J. Whitfield) 145

Hoffman, Richard L. On the Taxonomic Status, Distribution and Sub- species of the Milliped Pseudotremia fracta (Chamberlin) (Chordeu- matida: Cleidogonidae) 135

Lee, David S. and William M. Palmer. Records of Leatherback Turtles, Dermochelys coriacea (Linnaeus), and Other Marine Turtles in North Carolina Waters 95

Palmer, William M. (see Lee, David S.) 95

Ray, Clayton E., Elaine Anderson and S. David Webb. The Blancan carnivore Trigonictis (Mammalia: Mustelidae) in the Eastern United States 1

Roush, Mary Beth and Donald C. Tartar. Ecological Life History of Ptilostomis postica (Walker) (Trichoptera: Phryganeidae) in Green- bottom Swamp, Cabell County, West Virginia 107

Tartar, Donald C. (see Roush, Mary Beth) 107

Webb, S. David (see Ray, Clayton E.) 1

Number 6

Andre, John B. and Larry West. Nesting and Management of the Atlantic Loggerhead, Caretta caretta caretta (Linnaeus) (Testudines: Cheloni-

idae) on Cape Island, South Carolina, in 1979 73

Bennett, Charles R. (see Birkhead, William S.) Ill

Birkhead, William S. and Charles R. Bennett. Observations of a Small Population of Estuarine-inhabiting Alligators near Southport, North

Carolina Ill

Bozeman, Luke L. (see Camp, Carlos D.) 163

Burr, Brooks M. (see Walsh, Stephen J.) 83

Camp, Carlos D. and Luke L. Bozeman. Food of Two Species of Pleth-

odon (Caudata: Plethodontidae) from Georgia and Alabama 163

French, Thomas W. Notes on the Distribution and Taxonomy of Short- tailed Shrews (Genus Blarina) in the Southeast 101

163

Gaddy, L. L. Observations on Some Spiders of Maritime Forests on Four

South Carolina Barrier Islands 1 59

Laerm, Joshua. Systematic Status of the Cumberland Island Pocket

Gopher, Geomys cumberlandius 141

Lindquist, David G. (see Shute, John R.) 1

Link, Garnett W., Jr. (see Ross, Steve W.) 61

MacPherson, Kerry A. (see Ross, Steve W.) 61

Rose, Robert K. Small Mammals in Openings in Virginia's Dismal

Swamp 45

Ross, Steve W., Garnett W. Link, Jr. and Kerry A. MacPherson. New Records of Marine Fishes from the Carolinas, with Notes on Addi- tional Species 61

Schiller, David H. (see Tardell, Janice Heard) 153

Seidel, Michael E. A Taxonomic Analysis of Pseudemyd Turtles (Testu- dines: Emydidae) from the New River, and Phenetic Relationships in

the Subgenus Pseudemys 25

Shelley, Rowland M. A New Milliped of the Genus Brevigonus from South Carolina, with Comments on the Genus and B. shelf ordi (Loo- mis) (Polydesmida: Xystodesmidae) 51

Shute, John R., Peggy W. Shute and David G. Lindquist. Fishes of the

Waccamaw River Drainage 1

Shute, Peggy W. (see Shute, John R.) 1

Tardell, Janice Heard, Richard C. Yates and David H. Schiller. New Records and Habitat Observations of Hyla andersoni Baird (Anura:

Hylidae) in Chesterfield and Marlboro Counties, South Carolina 153

Travis, Joseph. A Key to the Tadpoles of North Carolina 119

Warren, Melvin L., Jr. New Distributional Records of Eastern Kentucky

Fishes 1 29

Yates, Richard C. (see Tardell, Janice Heard) 153

164

INDEX TO SCIENTIFIC NAMES

(New names in italics)

Numbers 5: and 6: (1981)

New Names

Brevigonus arcuatus 6:56-60 Pseudanophthalmus assimilis 5:65-66 Pseudanophthalmus calcareus 5:85-86 Pseudanophthalmus cordicollis 5:82-83 Pseudanophthalmus deceptivus 5:43-46 Pseudanophthalmus/tfsf/ga/u.s 5:50-5 1 Pseudanophthalmus/r/g/V/w5 5:86-87 Pseudanophthalmus georgiae 5:90-91 Pseudanophthalmus hypolithos 5:83-84 Pseudanophthalmus longiceps 5:79-80 Pseudanophthalmus nickajackensis 5:51-52 Pseudanophthalmus nortoni 5:48-49 Pseudanophthalmus pallidus 5:78-79 Pseudanophthalmus paradoxus 5:70-7 1 Pseudanophthalmus paulus 5:63 Pseudanophthalmus paynei 5:56-57 Pseudanophthalmus praetermissus 5:87-88

Pseudanophthalmus pusillus 5:56 Pseudanophthalmus rogersae 5:75-76 Pseudanophthalmus sanctipauli 5:67-69 Pseudanophthalmus scholasticus 5:84-85 Pseudanophthalmus scutilus 5:73-75 Pseudanophthalmus seclusus 5:76-77 Pseudanophthalmus sequoyah 5:52-53 Pseudanophthalmus sericus 5:62 Pseudanophthalmus steevesi 5:53-54 Pseudanophthalmus thomasi 5:80-82 Pseudanophthalmus unionis 5:57-58 Pseudanophthalmus ventus 5:64-65 Pseudanophthalmus wallacei 5:46-47 Pseudotremia fracta ingens 5:141-143 Pseudotremia fracta nantahala 5:143-144 Pseudotremia fracta paynei 5:139-141

Acacesia hamata 6:161 Acantharchus pomotis 6:15 Acanthepeira spp. 6:160,161,162 Acer rubrum 6:47,157 Acris

crepitans 6:121

gryllus 6:121 Agkistrodon

contortrix 5:150

piscivorus 5:150,157,158,159 conanti 5:118 leucostoma 5:118,124 piscivorus 5:117-128 Alligator 5:29

mississippiensis 5:148,150,156; 6: 111-117 Alosa

pseudoharengus 6:8

sapidissima 6:8,19 Alternanthera philoxeroides 6:4 Ameroduvalius 5:37,40,41

Amia calva 6:8

Ammocrypta pellucida 6:133-134 Andropogon sp. 5:130 Anguilla rostrata 6:8 Anisotremus surinamensis 6:66 Anolis carolinensis 5:149,156 Apeltes quadracus 6:64 Aphanotrechus virginicus 5:91 Aphredoderus sayanus 6:12,93 Araneus 6:160

bicentenarius 6:160,161

miniatus 6:160,161

pegnia 6:160,161

pratensis 6:160,161 Argiope 6:160

aurantia 6:161

trifasciata 6:161 Argyrodes

fictilium 6:162

furcatus 6:162

nephilae 6:162

165

Aristida stricta 6:154 Arundinaria gigantea 6:47,157

Bidens laevis 6:4 Blarina 6:49,101-110

brevicauda 6:46,49,101-1 10 churchi 6:106 shermani 6:106 telmalestes 6:46,49 carolinensis 6:49,101-1 10 sp. 6:48,104

telmalestes 6:101,106,108 Brevigonus 6:51-60 arcuatus 6:51,52,53,54,55,56-60 shelf ordi 6:51,52,54,55-56,57, 59,60 Brotula barbata 6:62 Bufo6:126 americanus 6:121 quercicus 6:121,126 terrestris 5:149,152,157; 6:121 woodhousei fowled 6:121

Canimartes 5:1

(?) cookii 5:2

cumminsii 5:6,9,21,27

(?) idahoensis 5:2 Caretta

caretta 5:149,160; 6:73 caretta 5:104-106; 6:73-82 Cemophora coccinea 5:150 Centrarchus macropterus 6:15 Cephalanthus occidentalis 5:108 Chaetodipterus faber 6:20 Chaetoduvalius 5:41 Chelonia mydas mydas 5:103-104 Chelydra serpentina 5:150 Chironomus sp. 5:114 Chologaster cornuta 6:12 Chrysemys 5:29

picta marginata 6:41 picta 6:41

picta X marginata 6:41 Cleidogona 5:136 Cleptoria 6:51,52,54,59

abbotti 6:59

macra 6:59

shelf ordi 6:51,54,55 Clethra alnifolia 6:157 Cnemidophorus sexlineatus 5:149,157 Coluber constrictor 5: 1 50, 1 57 Croatania 6:59 Crotalus 5:29

atrox 5:119

horridus 5:150

viridis 5:119 Croton punctatus 5:130 Cryptotis

parva 6:48,49 parva 6:47 Cyanea 5:98

capillata5:102 Cyclops sp. 5:111,114 Cyclosa sp. 6:161 Cynorca proterva 5:5 Cyprinus carpio 6:9 Cyrilla racemiflora 6:154,157

Daramattus 6:63

americanus 6:63-65

armatus 6:63

barnardi 6:63 Darlingtonea 5:37,40,41 Dermochelys 5:29

coriacea coriacea 5:95-103,105 Dolomedes triton 6:162 Dormitator maculatus 6:20 Dorosoma

cepedianum 6:8

petenense 6:8,19

Eira 5:15,22,27

barbara 5:1,10,18,19,20,21,22,26 Elaphe

guttata 5:150

obsoleta 5:150 Elassoma

evergladei 6:15

species 6:15,21

zonatum 6:15,93 Enhydrictis 5:23,27

ardea 5:23

166

Enneacanthus

chaetodon 6:16

gloriosus 6:16

obesus 6:16 Eremotherium 5:29 Eretmochelys imbricata imbricata

5:103 Erimyzon

oblongus 6:10,93

sucetta 6: 1 1 Erogala formosa 6:10 Esox

americanus 6:9,93

niger 6:9 Etheostoma

camurum 6:135

cinereum 6:134

fusiforme 6:17,20 barratti 6:17,20,21 fusiforme 6: 17,20,21

gracile 6:93

kennicotti 6:133

maculatum sanguifluum 6:135

nigrum 6:93,134 susanae 6:134

olmstedi 6:18

parvipinne 6:93

perlongum 6:1,18,21

serriferum 6:18

sp. 6:93

swaini 6:93

tippecanoe 6:134-135

zonale 6:93 Eucinostomus argenteus 6:20 Eumeces

inexpectatus 5:149

laticeps 5:149,150 Eustala anastera 6:161

Folsomia Candida 5:75 Fundulus

chrysotus 6:13,19

diaphanus 6:13,19

lineolatus 6:13

olivaceus 6:93

waccamensis 6:1,13,19,21 Furcillaria 6:59

Galera macrodon 5:1,2,4,5,21,22 Galictis 5:1,2,15,23,27

macrodon 5:2,21

vittata 5:10,20,21,22,26 Gambusia affinis 5:159; 6:14,93 Gasteracantha cancriformis 6:161 Gasterosteus aculeatus 6:64 Gastrophryne carolinensis 5:149,152,

157; 6:121,126 Gea heptagon 6:161 Geomys

bursarius 6:150

colonus 6:141,143

cumberlandius 6:141-151

fontanelus 6:141

pinetis 6:141,142,143,144,150 Glaucomys volans 5:130 Graptemys 6:39 Grison 5:23 Grisonella 5:10,15,20,21,23,27

cuja 5:10,18,19,20,21,22

Hemanthias leptus 6:66 vivanus 6:66

Heterandria formosa 6: 14

Hybognathus regius 6:9

Hybopsis species 6:19

Hyla 5:152 andersoni 6:123,124,153-158 chrysoscelis 6:120,123 cinerea 5:149,157,159; 6:125 crucifer 6:1 19,122,123,124,125 femoralis 6:120,123 gratiosa 6:120,123,125 squirella 5:149,157,158; 6:123

Ichthyomyzon fossor 6:130 Icatalurus

catus 6: 1 1

melas 6:1 1,19,93

natalis6:ll,93

nebulosus 6:11

platycephalus 6: 1 1

punctatus 6:12,19 Ilex

coriacea 6: 1 57

167

opaca 6:159

vomitoria 5:130,147; 6:159 Iva imbricata 5:130

Juncus

effusus 6:47

roemerianus 6:112 Justicia americana 6:134,136

Kinosternon subrubrum 5:149,152,156 Kleptochthonius affinis 5:78

Lagodon rhomboides 6:20 Lampetra

aepyptera 6:83-100

appendix 6:87

lamottenii 6:87

lethophaga 6:96

zanandreai 6:87,96 Lampropeltis getulus 5:150 Larus

argentatus 6:80

atricilla 6:79 Lasiotrechus discus 5:41 Latrodectus mactans 6:162 Lemna perpusilla 6:4 Lepidochelys kempi 5:104 Lepisosteus osseus 6:8 Lepomis

auritus 6:16

cyanellus 6:93

gibbosus 6:16

gulosus 6:16,93

macrochirus 6:16,93

marginatus 6:17,93

megalotis 6:93

microlophus 6:17

punctatus 6:17

sp. 5:158 Lepophidium jeannae 6:62 Lethenteron meridionale 6:83,87-88 Leucage venusta 6:160,161 Limnaoedus ocularis 6:121,123 Liriodendron tulipifera 6:157 Lutravus

cookii 5:1

(?) idahoensis 5:1

Lyncodon 5:27

Lynx rufus floridanus 6:45

Magnolia

grandiflora 5:130,147; 6:159

virginiana 6:154,157 Malaclemys terrapin 5:149,160 Mammuthus 5:7 Mangora 6:160

gibberosa 6:160,161

maculata 6:160,161

placida 6:161 Masticophis flagellum 5:150 Mecynogea lemniscata 6:161 Melanogrammus aeglefinus 6:62 Menidia extensa 6:1,14,19,21 Micrathena 6:160

gracilis 6:161

sagittata 6:161 Microdesmus longipinnis 6:66 Micropterus salmoides 5:158; 6:17 Microspora 5:114 Microtus

pennsylvanicus 5:132; 6:47,48 nigrans 6:46

pinetorum scalopsoides 6:47 Minytrema melanops 6:1 1 Morone

americana 6:14

saxatilis 6:14,19 Myrica cerifera 5:130,147 Mus 5:131

musculus 5:133 domesticus 6:47 Myripristis jacobus 9:63

Najas guadalupensis 6:3 Nannippus

minor 5:11

phlegon 5:8 Natrix fasciata 5:150 Neaphaenops 5:37,40,41 Nelsonites 5:37,40,41 Neoscona 6:160

arabesca 6:160,161

domiciliorum 6:160,161 Nephila clavipes 6: 1 60, 1 6 1 , 1 62

168

Nerodia taxispilota 5:127

Nitella sp. 6:3

Notemigonus crysoleucas 6:9,93

Notropis

ariommus 6:134

camurus 6:93

chalybaeus 6:9

chrysocephalus 6:134

cummingsae 6:10

fumeus 6:93

galacturus 6:130-131

hudsonius 6:10,19

hypselopterus 6:10,19

maculatus 6:10

petersoni 6:1,10

rubellus 6:134

sp. 6:131-132

spectrunculus 6:132

umbratilis 6:93

waccamanus 6:10 Noturus

gyrinus 6:12

hildebrandi 6:93

insignis 6:12

leptacanthus 6:12

phaeus 6:93

species 6:12,19,21 Nuphar luteum sagitifollium 6:2 Nyssa sylvatica 6:157

Ochrotomys

nuttalli 6:48 nutalli 6:46 Ocypode quadrata 6:76 Okkelbergia 6:88

Ondatra zibethicus macrodon 6:46 Oostethus

brachyurus 6:65 lineatus 6:65-66 Opheodrys aestivus 5:150 Ophisaurus ventralis 5:149 Oryzomys 5:131

palustris 5:132-133 Osmunda cinnamomea 6:157 Oxydendrum arboreum 6:157

Panicum

amarum 5:130

hemitomum 6:2,14 Pannonictis 5:23,27

pilgrimi 5:17,23,26

pliocaenica 5:23 Paraconger caudilimbatus 6:62 Perca flavescens 6:18 Percina

caprodes 6:136

copelandi 6:135,136

evides 6:136

oxyrhyncha 6:135-136,137

phoxocephala 6:135,136-137

sciera 6:93,136

squamata 6:135,137 Percopsis omiscomaycus 6:133 Peromyscus 5:131

leucopus 6:48 easti 6:46

gossypinus 5:132-133 gossypinus 6:46 Persea borbonia 5:130; 6:157,159 Phenacobius mirabilis 6:93 Phoxinus cumberlandensis 6:132-133 Pinus

elliottii 6:154,159

palustris 6:154

sp. 5:147

taeda5:130;6:154,157,159 Plesippus 5:29 Plethodon

cinereus 6:163

glutinosus 5:149,157,158,160

serratus 6:163-166

websteri 6:163-166 Pomoxis nigromaculatus 6:17 Prionotus

ophryas 6:67

stearnsi 6:67,69 Procyon lotor 6:74 Pseudacris

brachyphona 6: 1 1 9, 1 25

brimleyi 6: 119,123

nigrita 6: 1 25

169

ornata 6:123,125 triseriata 6:122,123,125 Pseudanophthalmus 5:37-94 alabamae 5:53,66,88,89,90 assimilis 5:53,61,65-66 calcareus 5:68,72,85-86,87 cordicollis 5:71,81,82-83,89 deceptivus 5:43-46,50,80 delicatus 5:47,59,60-61,62 digitus 5:49,50,5 1 ,59,6 1 ,63-64,65,66 egberti 5:38,66,69,70 engelhardti 5:38,42-43,46,47,50,60,78 fastigatus 5:44,45,50-51 frigidus 5:68,72,86-87 fulleri 5:38,48,49-50,5 1 ,52,53,54,64 fuscus 5:91,92 georgiae 5:72,81,90-91 gracilis 5:92 grandis 5:91 hadenoecus 5:92 higginbothami 5:92 hirsutus 5:58-60,61,62,63,64,65 hoffmani 5:92 holsingeri 5:38,46,47,48,61 hortulanus 5:92 hubrichti 5:38,66-67,69 hypertrichosis 5:91 hypolithos 5:68,72,83-84,85,86 jonesi 5:71,72-73,75,76,89,91 krekeleri 5:91 loedingi 5:38,50

longiceps 5:46,71,79-80,81,83,89 meridionalis 5:50 montanus 5:91

nickajackensis 5:45,49,50,5 1-52 nortoni 5:44,45,48-49 pallidus 5:47,54,71,74,77,78-79,89 paradoxus 5:66,68,70-71,72 paulus 5:59,63 paynei 5:47,56-57,58 petrunkevitchi 5:92 praetermissus 5:72,83,87-88 pusillus 5:44,56,57,58 quadratus 5:69-70 rogersae 5:73,74,75-76,85,91

rotundatus 5:47,50,61,78 sanctipauli 5:67-69,72 scholasticus 5:68,72,84-85,87 scutilus 5:73-75,76,89,91 seclusus 5:71,74,76-77,79,88,89 sequoyah 5:45,50,52-53,66 sericus 5:59,61,62 sidus 5:47-48,50 steevesi 5:45,53-54 sylvaticus 5:40,91,92 tennesseensis 5:54,55,56,57,58 thomasi 5:62,80-82,83,89 unionis 5:57-58 ventus 5:61,64-65,66 vicarius 5:66,70,7 1,92 virginicus 5:91 wallacei 5:44,45,46-47

Pseudemys 6:25-44

alabamensis 6:27,29,32,35

concinna 6:25,27,29,32,34,36,37,38, 39,40

concinna 6:27,29,31,34,35,36,40,41 hieroglyphica 6:27,29,3 1 ,32,34,35,

36,39,40,41 mobilensis 6:27,29,32,34,35,36 suwanniensis 6:27,29,34,35,36,40 texana 6:27,29,32,34,35,36,39

floridana 6:25-27,29,32,34,36,38, 39,40

floridana 6:27,29,31,32,34,35,36 hoyi 6:27,29,32,34,35,36,39 peninsularis 6:27,29,34,35,36,38

floridana X rubriventris 6:27

nelsoni 6:27,29,32,35,39

rubriventris 6:25,27,29,3 1 ,32,35,38,39

scripta 5:149,157,158,160 Pseudotremia 5:85,135-144

cocytus 5:137,138

cottus 5:135,136,137,139,144

fracta 5:135-144 fracta 5:138-139,143 ingens 5:140,141-143,144 nantahala 5:140,142,143-144 paynei 5:139-141,142,143

170

hobbsi 5:136

minos 5:137

nodosa 5:78

scrutorum 5:137,138,143 Ptilostomis

ocellifera 5:110,112

postica 5:107,116 Putorius ardeus 5:20

Quercus laevis 6:154 laurifolia 5:130; 6:159 marilandica 6:154 sp. 5:147 stellata 6:154 virginiana 5:130,147; 6:159

Rana 5:29; 6:126

areolata 6:125

catesbeiana 6: 1 25, 1 26

clamitans 6:1 19,120,126

heckscheri 6:1 19,126

palustris 6:125

pipiens 5:149; 6:125

sphenocephala 5:149; 6:125

sylvatica 6:125

utricularia 5:149,152,157

virgatipes 6:126 Rattus

norvegicus 6:76

rattus 6:76 Rhomphaea lacerta 6:162 Rhus

copallina 6:157

vernix 6:154,157 Riethrodontomys humulis humulis 6:46 Rubus allegheniensis 6:47

Sabal

palmetto 5:130; 6:159

sp. 5:147 Salix

caroliniana 5:130

nigra 5:108 Scaphiopus holbrooki 5:139,152,157;

6:121,126

Scincella laterale 5:139,156 Scirpus sp. 5:130 Semotilus

atromaculatus 6:93

lumbee 6:19 Sigmodon 5:131

hispidus 5:132-133 Sigmoria 6:51,52,54,55,56,59,60

laticurvosa 6:59

latior 6:52,56,59

quadrata 6:59

tuberosa 6:59 Smilax

laurifolia 6:157

spp. 6:47,154 Sminthosinus 5:27

bowleri 5:6,9,19,20,22,23 Sorex

longirostris 6:48 fisheri 6:46,49 longirostris 6:49 Spartina

alterniflora 5:130; 6:111

patens 6:1 16 Sphagnum spp. 6:157 Synaptomys

cooperi 6:48 helaletes 6:46,49 Syngnathus elucens 6:66

Tantilla coronata 5:150 Tayra 5:23

Terrapene Carolina 5: 139 Thamnophis

sauritus 5:150,157,158,159

sirtalis 5:150 Tidarren sisyphoides 6:162 Tillandsia usneoides 6:160 Trechoblemus 5:41 Trechus

cumberlandus 5:40,138

hydropicus 5:40

schwartzi 5:138

tennesseensis 5:138 tauricus 5:138 Trigonictis 5:1-36

cookii 5:1,2,3,4,6,7,8,11,15,17.

18,19,20,22,23,24,26,27-30,36 idahoensis 5:1,2,4,7,12,14,15,19 kansasensis 5: 1 ,2,3,4,7,22,23,24,35 macrodon 5:2,3-27,29,30,35-36 sp. 5:6,36

Trinectes maculatus 6:18

Trochictis 5:23,27

Typhasp. 5:130

171

Umbra

limii 6:93

pygmaea 6:8 Uniola paniculata 5:130; 6:162 Ursus americanus americanus 6:45

Vitis spp. 6:47

Xenolepidichthys americanus 6:63 Xenotrechus 5:37,41

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CONTENTS

Terrestrial Drift Fences With Pitfall Traps: An Effective Technique for Quantitative Sampling of Animal Populations. J. Whitfield Gibbons and Raymond D. Semlitsch 1

Marine and Freshwater Fishes of the Cape Fear Estuary, North Caro- lina, and Their Distribution in Relation to Environmental Factors. Frank J. Schwartz, William T Hogarth and Michael P. Weinstein ... 17

Nexting of the Green Turtle, Chelonia mydas (L.), in Florida: Historical Review and Present Trends. C. Kenneth Dodd, Jr 39

Thermal Preferenda and Diel Activity Patterns of Fishes from Lake Waccamaw. W. W. Reynolds, M. E. Casterlin and D. G. Lindquist 55

Helminths of Some Seabirds from North Carolina. Ronald W. Mobley and Grover C. Miller 61

Life History of a Coastal Plain Population of the Mottled Sculpin, Cottus bairdi (Osteichthyes: Cottidae), in Delaware. Fred C. Rohde and Rudolf G. Arndt 69

Distribution and Ecology of the Seepage Salamander Desmognathus aeneus Brown and Bishop (Amphibia: Plethodontidae) in Tennessee. R. L. Jones 95

Dental and Cranial Anomalies in the River Otter (Carnivora: Musteli- dae). Thomas D. Beaver, George A. Feldhamer and Joseph A. Chapman 101

Life History and Ecology of Chauliodes rastricornis Rambur and C. pectinicornis (Linnaeus)(Megaloptera: Corydalidae) in Greenbottom Swamp, Cabell County, West Virginia. Pamela S. Dolin and Donald C. Tarter Ill

Annotated Checklist of the Mammals of Georgia. Joshua Laerm, Lloyd E. Logan, M. Elizabeth McGhee and Hans N. Neuhauser 121

Distributional Records for Gastropods and Sphaeriid Clams of the Kentucky and Licking River and Tygarts Creek Drainages, Kentucky. Branley A. Branson and Donald L. Batch 137

Rediscovery and Distribution of Bembidion plagiatum Zimmermann (Coleoptera: Carabidae). Richard L. Hoffman 145

Incisor Malocclusion in a Specimen of Sylvilagus floridanus (Mamma- lia: Lagomorpha) from North Carolina. Gary W. Woody ard 151

A Survey of the Freshwater Mussels (Mollusca: Unionidae) of Middle Island Creek, West Virginia. Ralph W. Taylor and Beverly D. Spurlock 155

Errata and Miscellany 1 59

Manuscript Reviewers 160

Table of Contents, Nos. 5 and 6 ( 198 1 ) 162

Index to Scientific Names, Nos. 5 and 6 (1981) 164

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