I

N.C. DOCUMENTS

CLEARINGHOUSE

DEC 2 0 REC'D

STATE UBRAP.^Oe-

I

number 9

June 1983

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, 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

James W, Hardin, Department Rowland M. Shelley, Chief

of Botany, N.C State Curator of Invertebrates, N.C

University State Museum

Brimleyana, the Journal of the North Carolina State Museum of Natural His-

tory, will appear at irregular intervals in consecutively numbered issues. Con-

tents will emphasize zoology of the southeastern United States, especially North

Carolina and adjacent areas. Geographic coverage will be limited to Alabama,

Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North

Carohna, South Carolina, Tennessee, Virginia, and West Virginia.

Subject matter will focus on taxonomy and systematics, ecology, zoo-

geography, evolution, and behavior. Subdiscipline areas will include general

invertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and

paleontology. Papers will stress the results of original empirical field studies, but

synthesizing reviews and papers of significant historical interest to southeastern

zoology will be included.

Suitability of manuscripts will be determined by the Editor, and where neces-

sary, the Editorial Board. Appropriate specialists will review each manuscript

judged suitable, and final acceptability will be determined by the Editor.

Address manuscripts and all correspondence (except that relating to subscrip-

tions and exchange) to Editor, Brimleyana, N. C. State Museum of Natural

History, P. O. Box 27647, Raleigh, NC 27611.

In citations please use the full name — Brimleyana.

North Carolina State Museum of Natural History

North Carolina Department of Agriculture

James A. Graham, Commissioner

CODN BRIMD 7

ISSN D193-4406

A New Species of Woodland Salamander

of the Plethodon glutinosus Group

from the Southern Appalachian Mountains

Richard Highton

Department of Zoology,

University of Maryland, College Park, Maryland 20742

ABSTRACT. — A new species of woodland salamander, Plethodon

aureolus, occurs between the Little Tennessee and Hiwassee rivers on

the western slopes of the Unicoi Mountains and nearby lowlands in

southeastern Tennessee and adjacent North Carolina. It is a small-

sized member of the P. glutinosus group and was discovered by a study

of electrophoretic variation in 22 genetic loci. It is sympatric with the

white-spotted form of P. glutinosus (here recognized as a distinct spe-

cies, P. teyahalee) at 28 localities, and at one of these it is also sympat-

ric with typical brassy-spotted P. glutinosus. Plethodon aureolus

hybridizes with Unicoi Mountain P. Jordani on Sassafras Ridge, but

there is no evidence of significant hybridization between P. aureolus

and P. teyahalee or P. glutinosus.

Two unpublished electrophoretic studies of geographic genetic

variation in eastern woodland salamanders of the Plethodon glutinosus

group, one by Peabody (1978) and the other in preparation by Highton,

have revealed the existence of an undescribed species of the group. Its

range appears to be largely restricted to the western slopes of the Unicoi

Mountains and adjacent lowlands, between the Little Tennessee and

Hiwassee rivers, in Monroe and northern Polk counties, Tennessee, and

adjacent Graham and Cherokee counties, North Carolina.

Highton (1970) called attention to the presence of three distinct

geographically parapatric color pattern variants of P. glutinosus in the

southern Appalachian Mountain region: (1) populations in the moun-

tains of western North Carolina are characterized by having small dor-

sal white spots; (2) in populations from northeastern Georgia many

individuals lack dorsal spotting; and (3) populations to the west and

south of the above areas are characterized by having brassy-colored

dorsal spots. Highton (1972) mapped the distribution of three parapat-

ric dorsal pattern variants of P. glutinosus in Pennsylvania, Maryland,

Virginia and West Virginia. Two of these resemble the first and the

third southern Appalachian types in the color of their dorsal spots,

while a third, smaller. Coastal Plain variant is characterized by its very

small dorsal brassy-colored spots. I suggested that there may be limited

gene exchange between some of these parapatric forms and some pairs

may be at or close to the species level of evolutionary divergence. Our

unpublished genetic studies have shown that hybridization often occurs

Brimleyana No.9:l-20. June 1983.

2 Richard Highton

in the narrow overlap zones where the ranges of some of the above

parapatric forms are in contact. The species described here differs genet-

ically from all of the above types and throughout its range it is sympat-

ric with the white-spotted southern Appalachian variant of P. glutino-

sus. The two forms appear to be both morphologically and genetically

distinct at all 28 localities where they have been taken sympatrically.

The new species is characterized by possessing abundant brassy-

colored dorsal spots and by its small size. It is very distinct from the

sympatric, white-spotted, large-sized populations of P. glutinosus, but

some other nearby brassy-spotted populations of P. glutinosus are very

similar to it in appearance. Using the same electrophoretic methods and

genetic loci described in Highton and MacGregor (1983), the new spe-

cies was compared genetically to 11 samples of P. kentucki, 128 samples

of P. glutinosus and 41 samples oi P. jordani taken from localities scat-

tered throughout their ranges. It was also compared with a single sam-

ple of each of the other four species of the group (yonahlossee, cad-

doensis, ouachitae and fourchensis). Geographic genetic variation in the

latter three species was studied by Duncan and Highton (1979) and the

remaining results are being prepared for publication. The new species is

genetically distinguishable from samples of all of these species, just as it

is from sympatric P. glutinosus. However, I have failed to find any

morphological characters that may be used to distinguish it from some

allopatric types of brassy-spotted P. glutinosus. The diagnosis presented

here is therefore valid only for comparisons with the three forms of

eastern large Plethodon with which it is sympatric. This is the second

cryptic species of Plethodon discovered by electrophoretic studies of

genetic variation in proteins, the first being P. websteri Highton (1979).

The new species is named for its brightly-colored brassy dorsal

spots. The name is from the Latin word meaning gilded, ornamented or

very beautiful.

Plethodon aureolus, nevj species

Diagnosis. — An eastern Plethodon of the P. glutinosus group

(Highton and Larson 1979). It differs from sympatric white-spotted P. glutin-

osus by its smaller size, its relatively larger dorsal spots, the presence of

abundant brassy flecking in the dorsal iridophore spots, and more

abundant lateral white or yellow spotting. It differs from most nearby

populations of brassy-spotted P. glutinosus by its smaller size and light-

er chin. It differs from Unicoi Mountain P. jordani by the presence of

dorsal spots and by its more abundant white iridophore spotting on the

sides and legs.

Holotype.— USNM 238341, an aduh male collected at Farr Gap

(locality 1, Table 1), Unicoi Mountains, Monroe County, Tennessee, on

30 June 1979, by Richard Highton and Jeffrey K. Streicher.

New Species of Plethodon 3

Paratypes. — USNM 238342-51, topotypes, same collecting data as

the holotype.

Other material. — Plethodon aureolus from 31 localities (Table 1)

have been identified electrophoretically, and preserved specimens from

all of these sites will be deposited in the National Museum of Natural

History (USNM).

Description of holotype. — Before preservation, the length from the

tip of the snout to the anterior angle of the vent was 54 mm, to the

posterior angle of the vent 58 mm, and the total length 122 mm. There

are 16 costal grooves (equivalent to 17 trunk vertebrae) and the vomer-

ine teeth number 8 on the right side and 9 on the left. In life there were

abundant dorsal white iridophore spots with much associated brassy

flecking scattered on a black ground color. Similar spots were also pres-

ent on the dorsal surfaces of the legs and the top of the head. There

were abundant yellow iridophore spots on the sides of the head and

body and a few yellow iridiphore spots were also present on the chin

and belly. The chin is lighter than the belly.

Distribution. — Plethodon aureolus is known from southern and east-

ern Monroe and northeastern Polk counties, Tennessee, and also occurs

in northwestern Cherokee and western Graham counties, North Caro-

lina (Fig. 1).

Variation in P. aureolus. — There is little morphological variation in

P. aureolus throughout its small range, except that at higher elevations

in the northeastern part of its range the dorsal spotting may be very

reduced or absent in some individuals.

Remarks. — Although I had collected several P. aureolus during my

earlier study of variation in southern Appalachian large Plethodon

(Highton 1970), it was not recognized as distinct from other brassy-

spotted P. glutinosus until recently when we obtained new material for

our electrophoretic studies. The results have indicated that the large-

sized, brassy-spotted, dark-chinned populations from eastern Tennessee

are closely related to P. glutinosus from the northern part of its range

(New York west to Illinois and south through Kentucky, West Virginia,

western Virginia and eastern Tennessee). Since the type locality (Prince-

ton, New Jersey) of P. glutinosus is within this area, this form will

retain the name P. glutinosus regardless of the eventual taxonomic sta-

tus of the other geographic variants. These northern populations of P.

glutinosus are characterized by possessing much darker chins than those

of the white-spotted populations (Highton 1962), P. kentucki (Highton

and MacGregor 1983) and P. aureolus. However, in the immediate vi-

cinity of the range of P. aureolus, many individuals of otherwise geneti-

cally typical P. glutinosus possess unusually light chins, making it very

difficult to distinguish the two species without an analysis of their pro-

teins. The two have been found sympatrically only at locality 4, along

Richard Highton

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New Species of Plethodon 7

Ellis Branch of Spring Creek, near Springtown, in northern Polk

County, Tennessee, with no genetic evidence of current hybridization

between them (see below).

At 28 of the 3 1 P. aureolus localities, white-spotted P. glutinosus

has been taken in sympatry. Highton (1972) pointed out that the name

Plethodon jordani teyahalee Hairston (1950) is available for the latter

form. The population at the type locality, Teyahalee Bald, Graham-

Cherokee County Une, North Carolina, is probably of hybrid origin, but

is much more like white-spotted P. glutinosus than P. jordani (Highton

1970). As shown below, this form occurs sympatrically with both P.

aureolus and P. glutinosus at locality 4, where apparent reproductive

isolation exists between all three forms. On the basis of this evidence, I

suggest that the white-spotted form should also be recognized as a dis-

tinct species, P. teyahalee. Its distribution and genetic relationships will

be discussed in a later paper.

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Fig. 1. Distribution of P. aureolus in southwestern North Carolina and south-

eastern Tennessee between the Hiwassee and Little Tennessee rivers. Crosses

represent localities (46-48) where P. aureolus and P. jordani hybrids occur.

8 Richard Highton

Plethodon jordani, another Ught-chinned species, occurs at higher

elevations in the Unicoi Mountains (Highton 1962, 1970). An electro-

phoretic analysis of geographic genetic variation in P. jordani and P.

teyahalee in the southern Appalachian Mountains by Peabody (1978)

showed that the latter species is more closely related to some popula-

tions of P. jordani than it is to most other P. glutinosus. Plethodon

jordani and P. teyahalee hybridize extensively in a number of contact

zones (Highton 1970; Highton and Henry 1970; Peabody 1978), includ-

ing the entire periphery of the range of P. jordani in the Unicoi Moun-

tains, but in many areas the two species overlap extensively without

evidence of hybridization. Although we have no evidence of current

hybridization between P. aureolus and P. teyahalee, a transect through

the only known contact zone between P. jordani and P. aureolus on

Sassafras Ridge provided evidence of hybridization between these two

species (unpublished data).

The small size of P. aureolus is one of its most distinctive features.

As discussed in Duncan and Highton (1979), size is a difficult character

to use in salamander taxonomy. In Plethodon, however, mean adult size

and maximum length are sometimes quite consistent among genetically

closely related populations. At the type locality of P. aureolus, 356 indi-

viduals were collected for a study of the life history of the species. The

largest specimen is 67 mm from snout to anterior angle of the vent.

Only three other individuals are over 61 mm in snout-vent length. All of

the other species of southern Appalachian large Plethodon attain much

larger sizes (Highton 1970). The mean adult size of both P. glutinosus

and P. teyahalee is usually at least 70 mm and large adults are often

over 80 mm (record size, a P. teyahalee from Davis Ridge in the Great

Smoky Mountains, Sevier Co., Tennessee, 94 mm). In a sample of 78 P.

teyahalee from the type locality of P. aureolus the 10 largest females

range from 75-90 mm (mean 81.6) and the 10 largest males range from

74-90 mm (mean 78.5).

I suggest that an appropriate common name for P. aureolus is the

Tellico salamander. Tellico Plains is located centrally in its range and

much of this region is drained by the Tellico River and its tributaries.

Electrophoretic Genetic Analysis of Proteins.

Once collections from the type locality of P. aureolus were found to

differ genetically from all other Plethodon species, extensive field work

was done in the Unicoi Mountains and adjacent areas to determine the

geographic distribution of each of the four species and to study their

geographic and genetic interactions. Salamanders were obtained from 84

localities, and individuals from each of these were compared using the

same 22 genetic loci and methods described in Highton and MacGregor

(1983). When compared with a sample of P. glutinosus from near the type

New Species of Plethodon

Fig. 2. Sites of samples of P. glutinosus (hollow triangles), P. teyahalee (solid

circles) and P. jordani (solid squares) identified electrophoretically in southwestern

North Carolina, southeastern Tennessee and northern Georgia. Dotted lines outline

known range of P. aureolus.

locality in New Jersey (locality 10), 32 samples are genetically very similar

and are referred to this species. All are from areas west, north or south of

the range of P. aureolus (Fig. 2) and represent large, brassy-spotted

animals similar in appearance to those from New Jersey, although some

have chins that are much lighter than those in northern populations of P.

glutinosus. Plethodon teyahalee was found at 44 localities, and at 28 of

these P. aureolus was taken in sympatry. Plethodon teyahalee occurs at

low and intermediate elevations east of the range of P. glutinosus (Fig. 2).

Since P. teyahalee may be distinguished from the other two species by its

color pattern in life, its distribution west of the French Broad River is

probably accurately indicated by the map in Highton (1970, Fig. 5).

Two samples from populations of Unicoi Mountain P. jordani are

included in order to compare this species with the three other forms. These

10 Richard Highton

are the same salamanders studied by Peabody (1978) and are from near

Junction, on Snowbird Creek (locality 13) and Johns Knob (locality 14).

The distribution of P. jordani in the Unicoi Mountains was mapped by

Highton (1970) and its range supposedly includes all of the higher areas of

this mountain range. It was therefore surprising to discover that at four

localities in the northernmost part of the Unicoi Mountains, populations

resembling P. jordani in coloration (no dorsal spotting and reduced lateral

spotting), are assigned genetically to P. aureolus. These are from near

Cherry Log Gap (localities 22 and 23), Naked Ground (locality 38), and

Stratton Bald (locality 39). Thus the northernmost known sites for P.

jordani in the Unicoi Mountains are now in the vicinity of Johns Knob.

Highton (1970) called attention to the apparent hybridization between P.

jordani and P. teyahalee all around the periphery of the range oi P. jordani

in the Unicoi Mountains. Two transects reported by Peabody ( 1 978) (from

Johns Knob west along the North River, and from Junction east along

Snowbird Creek) have confirmed this hybridization genetically.

At some of the 84 localities few animals were collected. Voucher

specimens were preserved from all of the sites, and tissue samples (usually

viscera and tail muscle) from some were used for electrophoresis instead of

the material used in my previous work on Plethodon (whole animal

homogenates). The three general protein loci usually cannot be scored

from homogenates of viscera and tail muscle. Rather than use many with

small sample sizes and/ or incomplete genetic data, I present here the

results of a complete genetic analysis of only 1 7 populations in addition to

the sample of P. glutinosus from New Jersey (locality 10). These are from

scattered sites throughout the local ranges of the four species: 4 P. aureolus

(localities 1-4), 6 P. glutinosus (localities 4-9), 5 P. teyahalee (localities 1 , 2,

4,11,12) and 2 P. jordani (localities 1 3 and 1 4). Three species are sy mpatric

at locality 4, and P. aureolus and P. teyahalee are sympatric at localities 1

and 2. Material from all other localities shows no unusual genetic variation

beyond that observed in the 18 samples for which complete genetic analysis

is presented.

Table 2 provides the frequency data of genie variation of 18 popula-

tions from 14 localities. Of the 22 presumed genetic loci evaluated, 3

(Mdh-1, Pep, and Pt-3) show no variation. Three loci are monomorphic

except for a single population: a-Gpd has a rare slower allelomorph in P.

glutinosus (.02) at locality 8, Gdh has a slower allelomorph (.29) in P.

teyahalee at locality 1, and Mdh-2 has a rare faster allelomorph (.02) in P.

aureolus at locality 2. Table 3 gives Nei standard genetic distances {D) and

normalized identity of genes (/) (Nei 1972) for all comparisons and the

mean heterozygosity (//) estimated from allelomorph frequencies. The /

values are clustered by the UPGM A method (Sneath and Sokal 1973) in a

phenogram in Figure 3.

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The 18 samples cluster into 4 groups, each representing a taxonomic

species. There is very little geographic genetic variation within P. teyaha-

lee, but there is a considerable amount in both P. glutinosus and P.

aureolus. Indeed, some samples within both of the latter species are as

different genetically as are some comparisons of P. teyahalee and P.

jordani. The southwestern populations oi P. jordani from the Unicoi and

Nantahala Mountains are genetically very similar to P. teyahalee (Pea-

body 1978); in fact, these two species are more closely related than any

other two species of Plethodon yet examined (see Highton and Larson

1979). It is therefore not surprising that they hybridize so extensively

(Highton 1970).

Plethodon jordani is the most variable species (mean //=.21), P.

aureolus (mean H-.\2) and P. glutinosus (mean H-. 1 1) are intermediate,

while P. teyahalee (mean H=.01) is the least variable. Compared to more

northern populations of P. glutinosus and P. jordani (Highton and Mac-

Gregor 1983), these southern samples are much more variable. The P.

glutinosus, however, have slightly lower average H values than Arkansas

and Oklahoma P. glutinosus (Duncan and Highton 1979).

In light of my unpublished evidence that P. aureolus and P. jordani

hybridize at localities 46-48 on Sassafras Ridge (the only known area of

contact between the two species), the proper taxonomic relationship

between the two forms is difficult to decide. The average D of the 8

comparisons between the two forms (.3 1 ) is not very different from that of

the 28 comparisons between P. aureolus and P. glutinosus (.29) or the 20

comparisons between P. aureolus and P. teyahalee (.43), two species with

which P. aureolus is sympatric and is not known to hybridize. In the

southwestern isolates of P. jordani (in the Great Smoky Mountains,

Cowee Bald, the Nantahala Mountains, Cheoah Bald, and in the Unicoi

Mountains), P. jordani is always a high altitude species, whereas P. aureo-

lus is mostly a lower altitude form. The color pattern is very different

(except in the northern Unicoi Mountains where hybridization between

the two has occurred). None of the above mentioned populations of nearby

P. jordani has as abundant lateral and dorsal yellow, white or brassy

spotting. Plethodon aureolus is not significantly more similar genetically

to adjacent samples of Unicoi Mountain P. jordani than it is to other

populations of P. jordani throughout its range (Peabody 1978). The only

similarity between P. jordani and P. aureolus is that they are both smaller

than Appalachian populations of P. glutinosus and P. teyahalee. I there-

fore regard the interbreeding between the two on Sassafras Ridge as a case

of hybridization between species rather than intergradation between con-

specific populations. Considering the very extensive hybridization between

P. jordani and P. teyahalee throughout their contact zone in the Unicoi

Mountains, it is curious that P. teyahalee does not appear to hybridize with

the hybrid populations of P. aureolus and P. jordani on Sassafras Ridge.

16 Richard Highton

The isolating mechanisms that keep P. aureolus from interbreeding with P.

teyahalee apparently are present in the aureolus-jordani hybrids in suffi-

cient degree to prevent the usual interbreeding of P. jordani and P.

teyahalee at all 3 Sassafras Ridge sites (localities 46-48).

Since without genetic data it is extremely difficult to correctly identify

many individuals of this complex of southern Appalachian large Pleth-

odon, particularly some P. aureolus and P. glutinosus, there is a problem

in assigning individuals to species before examining the genetic data. After

a long search for a site where the two widely sympatric species, P. teyahalee

and P. aureolus, contact the parapatric species, P. glutinosus, a localilty

was discovered along Ellis Branch, near Springtown, Polk County, Ten-

nessee (locality 4), where the three forms are sympatric. Nei genetic identi-

ties between all 24 individuals from this localilty are clustered in a

UPGM A phenogram (Fig. 4). The results clearly show that the 24 animals

are separable into three groups consisting of 1 3 P. teyahalee, 5 P. aureolus

and 6 P. glutinosus. Four additional very small animals from locality 4

were also examined at some of the diagnostic loci and were identified as

P. teyahalee, but are not included in the genetic analysis because of

incomplete genetic data for several loci. Each of these samples clusters

with others of its own species (Fig. 3), and only the P. glutinosus sample

has a higher than average H value (Table 3). The alleUc data in Table 4

indicate that there is only one locus (Ldh-muscle) in which there are

fixed differences between all three species. At the other differential loci,

sometimes two of the species have identical electromorphs and some-

times there are rare electromorphs of the same kind found in one or

both of the other species. This latter pattern of variation is also present

in sympatric populations of P. glutinosus and P. kentucki (Highton and

MacGregor 1983) and could result from inheritance of the same elec-

tromorphs from their common ancestor, or occasional hybridization

between the species after complete differentiation had occurred. The

relationships in Figures 3 and 4 and the data in Tables 2, 3 and 4 are

considered strong evidence for the recognition of all three forms as dis-

tinct species. The Ldh-muscle data clearly show that there is not a single

F| hybrid between any of the three species at the Ellis Branch locality,

as does the pattern of variation at the other differential loci.

I have no explanation as to why in three cases an electromorph from

another species appears as a rare homozygote instead of in the expected

heterozygous condition [P. teyahalee #9, Alb; P. aureolus #21, Idh-2; and

P. glutinosus #16, Ldh (heart)] (see Table 4).

The Pep electromorphs of P. aureolus are faster than those of the

other two species at locality 4 and are indicated as different in Table 4. This

difference could not be consistently detected on comparison gels of sam-

ples of the three species at other localities and is therefore not regarded as a

New Species of Plethodon

17

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New Species of Plethodon 19

polymorphic locus in Table 2. Thus the D values between P. aureolus and

the other two species indicated in Figure 4 are slightly higher than they

would be if this Pep difference had not been detected (see Fig. 3.)

Although there is little or no evidence for hybridization between P.

teyahalee and P. glutinosus in the samples from locality 4, at locality 44

there are 4 individuals in a sample of 17 that have dorsal spots of interme-

diate coloration. In addition, there is a low frequency of P. glutinosus

electromorphs at all of the loci that differentiate the two species (Alb, Est,

Ldh (muscle), Pt-2, and Trf). This is interpreted as evidence for hybridiza-

tion between the two species at this locality. It is not surprising that P.

teyahalee and P. glutinosus hybridize in some areas and not in others, since

this same pattern occurs between P. teyahalee and P. jordani in the

southern Appalachian Mountains.

ACKNOWLEDGMENTS.— I wish to thank the many persons who

aided in the field and laboratory work, especially Ivette Aguirre, Debra

Glover, Karol Jackson, Scrap Oktay, Robert Peabody, Joshua Schwartz,

Jeffrey Streicher and Susan Fogle. Richard L. Hoffman suggested the

scientific name. I am also grateful for the financial support of the work by

the National Science Foundation (grants DEB-79-03858 and DEB-81-

17973) and the Computer Science Center of the University of Maryland.

David B. Wake made helpful comments on the manuscript.

LITERATURE CITED

Duncan, Robert, and R. Highton. 1979. Genetic relationships of the eastern large

Plethodon of the Ouachita Mountains. Copeia 1979(1): 95-1 10.

Hairston, Nelson G. 1950. Intergradation in Appalachian salamanders of the genus

Plethodon. Copeia, 1950(4): 262-273.

Highton, Richard. 1962. Revision of North American salamanders of the genus

Plethodon. Bull. Fla. State Mus. Biol. Sci. 6;235-367.

1970. Evolutionary interactions between species of North American

salamanders of the genus Plethodon. Part 1. Genetic and ecological rela-

tionships of Plethodon jordani and P. glutinosus in the southern Appa-

lachian Mountains. Evol. Biol. 4:2\\-24\.

1972. Distributional interactions among eastern North American

salamanders of the genus Plethodon. Pp. 139-188 in P.C. Holt (ed.). The

distributional history of the biota of the southern Appalachians, Part III:

Vertebrates. Res. Div. Monogr. 4, Va. Polytech. Inst. State Univ., Blacks-

burg. 306 pp.

1979. A new cryptic species of salamander of the genus Plethodon

from the southeastern United States (Amphibia: Plethodontidae). Brim-

leyana 1:31-36.

, and S. A. Henry. 1970. Evolutionary interactions between species of

North American salamanders of the genus Plethodon. Part 2. Variation in the

electrophoretic migration of plasma proteins of Plethodon Jordani, P. gluti-

nosus, and their natural hybrids. Evol. Biol. -^.-241-256.

20 Richard Highton

and A, Larson. 1979. The genetic relationships of the salamanders of

the genus Plethodon. Syst. Zool. 25;579-599.

, and J. R. MacGregor. 1983. Plethodon kentucki Mittleman: a

valid species of Cumberland Plateau woodland salamander. Herpetologica

39.189-200.

Nei, Masatoshi. 1972. Genetic distance between populations. Am. Nat. 706;283-292.

Peabody, Robert B. 1978. Electrophoretic analysis of geographic variation of

two Appalachian salamanders, Plethodon jordani and Plethodon glutinosus.

Ph.D. Dissert., Univ. of Maryland, College Park. Ill pp.

Sneath, Peter H. A., and R. R. Sokal. 1973. Numerical Taxonomy. W. H.

Freeman and Co., San Francisco. 573 pp.

Accepted 31 December 1983

Lower Wilson Creek, Caldwell County, North Carolina:

A Thermal Refugium for Reptiles?

Robert Wayne Van Devender

Department of Biology, Appalachian State University,

Boone, North Carolina 28608

AND

Paul F. Nicoletto

Department of Biology, Virginia Polytechnic Institute and

State University, Blacksburg, Virginia 24061

ABSTRACT. — Fourteen species of reptiles were found in a year-long

study of lower Wilson Creek, Caldwell County, North Carolina. Of

these, four — Anolis carolinensis, Scincella laterale, Eumeces inexpecta-

tus, and Tantilla coronata — represented significant range extensions

into the northwestern mountains of the state. Climatological studies in

the area revealed a relatively equable temperature regime, with lower

daytime highs and higher daytime and winter lows than nearby Lenoir.

A combination of exposed rock for basking, deep fissures, and warmer

night and winter temperatures is probably responsible for the con-

tinued presence of these species well outside their heretofore known

ranges. These populations may represent relicts dating from late in the

Hypsithermal Interval, approximately 7000 years ago.

INTRODUCTION

Wilson Creek and its tributaries drain most of the southeastern

slope of Grandfather Mountain in Avery, Burke and Caldwell counties,

North Carolina. These streams descend steeply from over 1220 m

(4000') to about 330 m (1080') where Wilson Creek flows into the Johns

River on its way to the Catawba River. Wilson Creek is one of the more

rugged and scenic streams of the eastern Blue Ridge front, with water-

falls and rapids in the main gorge. The discovery of several green

anoles, Anolis carolinensis Voigt, in the lower part of the Wilson Creek

gorge in August 1980 indicated that this area might also harbor other

species normally found only in lower, warmer parts of North Carolina.

We therefore set out to determine which species of reptiles occur in the

area, whether they represent natural populations, and what aspects of

the area are responsible for maintaining the reptile community.

MATERIALS AND METHODS

The study was carried out in the gorge of lower Wilson Creek

approximately 18 km (1 1 mi) west of Lenoir. The site was in the Pisgah

National Forest and included the riparian zone along Wilson Creek and

the steep, northeastern side of the gorge up to about 765 m (2510') on

Brimleyana No.9:21-32. June 1983. 21

22 Robert Wayne Van Devender and Paul F. Nicoletto

the southern and western slopes of Adams Mountain and the western

slope of Loose Mountain (USGS Collettsville 7.5' topographic map).

The site was centered near 81°44'W 35°36'N along county road 1328

three to five km north of county road 1337 and about 2.5 km north of

Brown Mountain Beach resort.

In this area, Wilson Creek cuts a gorge that drops off from about

700 m (2300') to the streambed at 365 m (1200'). The northeastern side

of the gorge, which includes the study area, consists of steep, rugged

slopes with exposed granite bedrock interdigitating with woods domi-

nated by stunted oaks {Quercus ssp.), maples {Acer ssp.), sweet gum

{Liquidambar), and Virginia pines (Pinus virginiana). Occasional small

patches of vegetation are isolated in extensive areas of bare rock. Deep

crevices and fissures are common throughout the area and abundant in

the streambed.

The site was searched for reptiles several times in August 1980 and

weekly from September 1, 1980 until March 15, 1981. Sporadic visits

were then made through August 1981. Thirty-six site visits were made,

and over ninety hours were spent searching for reptiles. During warmer

periods visual searches for active animals were supplemented by turning

rocks and logs. Winter searches were mostly visual searches of the fis-

sured granite along the creek. Species and activity were recorded when-

ever a reptile was encountered. Voucher specimens of most species were

deposited in the vertebrate collection at Appalachian State University.

In order to learn more about the local environment and to deter-

mine whether it was unique in any way, temperature and rainfall data

were obtained for the Wilson Creek area and compared with similar

data for comparable elevations at nearby Lenoir (U. S. Weather Ser-

vice). A thermograph was placed at Brown Mountain Beach resort on

September 1, 1980 and checked weekly until March 15, 1981. Data were

recorded for 186 days when the thermograph functioned properly. Rainfall

data for Wilson Creek consisted of fifteen years of data from Mortimer,

a short distance upstream from the study area (U. S. Forest Service

environmental statement for the Wilson Creek area, pers. comm., Mr.

Pat Cook, District Ranger). Lenoir was chosen for comparison because

of its proximity and because Nicoletto's experience in the area indicated

that none of the unusual species occurred there.

Since temperature, in its various guises, potentially limits reptile

distributions, daily maximum and minimum temperatures at Wilson

Creek were used separately as independent variables in linear regres-

sions against paired Lenoir data (MINITAB computer program pack-

age; Ryan et al. 1976). These regressions and long term data for Lenoir

(49 years, U. S. Weather Service) were used to estimate mean monthly

minimum temperatures at Wilson Creek. Monthly and annual rainfall

Lower Wilson Creek NC Reptiles 23

data at the two sites were compared with paired /-tests. Statistical sour-

ces were Sokal and Rohlf (1969) and Rohlf and Sokal (1969). Signifi-

cance levels of P < 0.05 and P < 0.01 are indicated throughout this

paper by one (*) and two (**) asterisks, respectively.

RESULTS

Fourteen species of reptiles were found at the study site (Table 1).

Local residents also reported that rattlesnakes, Crotalus horridus Lin-

naeus, were occasionally seen in the area. A medium-sized black snake

seen but not captured was probably a black rat snake, Elaphe obsoleta

(Say). Most species found along Wilson Creek are those to be expected

in the mountains, but four species represent significant range extensions

into this part of North Carolina of forms that are usually in lower,

warmer areas. These species are the anole; the ground skink, Scincella

laterale (Say); the southeastern five-lined skink, Eumeces inexpectatus

Taylor; and the southeastern crowned snake, Tantilla coronata Baird

and Girard.

Lower Wilson Creek is apparently an unusual microclimate in the

mountains. It is wetter and warmer than the Lenoir area, where the

unusual species do not occur. Annual precipitation averaged 136.9 cm

(53.9 in) at Wilson Creek and 125.3 cm (49.3 in) at Lenoir. In matched

data for 15 years prior to 1980, annual and monthly averages were

higher at Wilson Creek (**), with differences of 0.23 to 1.7 cm (0.09-

0.67 in).

Figure 1 shows the relationship between daily minimum tempera-

tures at Wilson Creek and Lenoir. The regression is highly significant

(**) and indicates not only that Wilson Creek is warmer but also that

the magnitude of the difference is greatest when the temperature is low-

est. At 0°C in Lenoir, Wilson Creek should be about 3.8° C. Wilson

Creek was cooler than Lenoir on only six of the 186 days (**, sign test).

Daily maximum temperatures at the sites are presented in figure 2. The

regression is highly significant (**), and the two sites are rather differ-

ent. Wilson Creek had lower maximum temperatures 169 of 178 days

(**, sign test). There is about 2°C difference between the sites at 0°C in

Lenoir, but this difference gets larger as do maximum temperatures.

Daily temperature ranges are narrower at Wilson Creek than at Lenoir.

Monthly means of minimum daily temperature at Lenoir (49 years) and

estimates for Wilson Creek are presented in figure 3. Wilson Creek is

generally warmer and does not have a month with a mean low tempera-

ture below freezing; even in winter there are numerous warmer days.

Coldest temperatures recorded during the study were -12°C (10° F) at

Wilson Creek and -15° C (5° F) at Lenoir.

24

Robert Wayne Van Devender and Paul F. Nicoletto

Lower Wilson Creek NC Reptiles

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DISCUSSION

Four of Wilson Creek's reptiles represent significant range exten-

sions into the northwestern mountains of North Carolina (Conant 1975;

Martof et al. 1980; pers. comm., W. M. Palmer and A. L. Braswell) as

discussed below.

Anolis carolinensis, — Anoles are common in the Coastal Plain of

North Carolina almost to Virginia and are known from a couple of

Piedmont Plateau and Mountain areas (Martof et al. 1980; pers.

comm., Palmer and Braswell). There are only a few published records

for this species in the mountains. Weller (1930) found them not

uncommon at about 457 m (1500') near Chimney Rock, Rutherford

County. Bruce (1965) found several individuals in the gorges of the

southern Blue Ridge escarpment, Transylvania County, where they

reached an altitude of 670 m (2200'). Anoles have also been reported at

elevations below 550 m (1800') in the Tennessee part of the Great

Smoky Mountains National Park (King 1939; Huheey and Stupka

1967). Another population near Townsend, Tennessee, just north of the

Park, has been studied recently (Wade and Echternacht 1980, and pers.

comm.). All montane sites for this species are associated with exposed,

fractured rock with deep fissures, usually near the head of rather steep

gorges. These conditions apparently ameHorate the cool conditions that

exclude anoles from most of the mountains. The exposed rocks allow

behavioral thermoregulation in the face of the cold, while the crevices

provide a means to escape severe cold weather, for which they seem

otherwise ill adapted (Mount 1975).

The Wilson Creek population represents a range extension of about

66 km (41 mi) north and 50 km (31 mi) east of the Rutherford County

site, the closest published record. Wilson Creek is also about 161 km

(100 mi) west of a Randolph County population in the northern Pied-

mont. The northernmost Tennessee population is near Pigeon Forge,

which is about 48 km (30 mi) south and 170 km (105 mi) west of the

Wilson Creek site. The anoles in Caldwell County are one of the most

northern populations known, and are probably the population that

experiences the most severe winter conditions encountered by the species.

Eumeces inexpectatus. — Southeastern five-lined skinks are not

found in the higher mountains of North Carolina, even though they do

extend to the north on either side of the mountains (Conant 1965; Mar-

tof et al. 1980; pers. comm.. Palmer and Braswell). As with anoles, there

are few records for this species in the mountains. Bruce (1965) reported

them at elevations up to 550 m (1800') in the gorges along the south-

eastern escarpment of the Blue Ridge Mountains, Transylvania

County. The species has been found infrequently in the Great Smoky

Mountains National Park up to about 610 m (2000') (King 1939;

Lower Wilson Creek NC Reptiles 29

Huheey and Stupka 1967). Western Piedmont records from North

Carolina are from Gaston, Randolph and Stokes counties (pers. comm.,

Palmer and Braswell). The Wilson Creek population apparently fills in

a rather large hiatus between known populations. The site is about 225

km (140 mi) east-northeast of Swain County sites, 141 km (90 mi) north-

east of Transylvania County records, 105 km (65 mi) north-northwest

of Gaston County sites, 160 km (100 mi) west of Randolph County

populations, and 140 km (85 mi) west-southwest of the Stokes County

site. The closest Tennessee population is at least 160 km (100 mi) to the

west. These lizards seemed to prefer rather xeric situations compared to

other skinks at the site, as reported by Mount (1975) in Alabama and

Minton (1972) in Indiana. Otherwise, there was little difference in the

sites where this species and Eumeces fasciatus were found.

Scincella laterale. — Ground skinks are widely distributed in south-

eastern United States except at higher elevations (Conant 1965; Mar-

tof et al. 1980; pers. comm.. Palmer and Braswell). Bruce (1965) found

this species up to 945 m (3100') in southwestern North Carolina. It has

been reported from the Great Smoky Mountains National Park up to

about 793 m (2600') (King 1939; Huheey and Stupka 1967). Weller

(1930) also found the species near Chimney Rock, Rutherford County,

North Carolina. Several general works suggest that this species is rela-

tively tolerant of dry, hot places (Mount 1975; Minton 1972), but it can

apparently also live in some cooler, higher places.

The Wilson Creek site is about 40 km (25 mi) north-northeast of

the closest record in McDowell County (pers. comm.. Palmer and

Braswell) and represents the most northern montane population yet

reported.

Tantilla coronata. — The distribution of the southeastern crowned

snake includes many areas in the Coastal Plain and Piedmont Plateau

and several areas in the mountains (Conant 1975; Martof et al. 1980;

pers. comm.. Palmer and Braswell). The highest record for North Carol-

ina is 428 m (1400') in Transylvania County (Bruce 1965). In the Great

Smoky Mountains National Park this species has been recorded only on

the Tennessee side at elevations up to 610 m (2000') (King 1939; Huheey

and Stupka 1967). The Wilson Creek specimen extends the known

range of this species in North Carolina 40 km (32 mi) north-northeast of

a Burke County site and about 80 km (50 mi) west of an Alexander

County locality. It also extends slightly the known elevational range in

North Carolina.

Co-occurrence of four species in one area outside their previously

known distributions argues fairly strongly that the populations are

native to the site and brings up questions about why they persist at the

site and when and how they arrived there. Their survival seems to be tied

to the microenvironment of Wilson Creek gorge. Exposed rocks provide

30 Robert Wayne Van Devender and Paul F. Nicoletto

abundant basking sites to overcome locally cooler daytime tempera-

tures. Warmer evening and winter temperatures provide a relatively sta-

ble environment, and deep fissures permit escape from the occasional

severe surface conditions. This combination of conditions probably

occurs in other protected montane areas that could also support popula-

tions of these and/ or other species that prefer warmer conditions.

Arrival of these species in the Wilson Creek area may be a rela-

tively recent event or a more ancient one. Recent arrival would proba-

bly be the result of immigration by populations expanding up the

Catawba and Johns rivers into the area. Since there is no other evidence

for recent range expansions of native reptiles in the eastern United

States, it is unlikely that this hypothesis is accurate. We consider these

populations to be relicts and offer the following hypothesis about their

origins.

During the last glacial maximum, about 18,000 years ago (yr BP),

essentially all of the Coastal Plain of the Carolinas was occupied by a

relatively Boreal forest of spruce and jack pines, indicative of a much

cooler climate that excluded most southern plants and many southern

animals (e.g. Watt 1979, 1980; Watt and Stuiver 1980; Flint 1971; Del-

court 1980; Wright 1976). The reptile species now characteristic of the

Coastal Plain, including the four relicts at Wilson Creek, were found

only much farther south. The warming trend that led to the retreat of

the glaciers also resulted in the immigration of southern plants and

animals into the Carolina Coastal Plain (Delcourt 1980; Webb 1981;

Watts 1979, 1980; Watts and Stuiver 1980). The warming trend reached

its climax in the Hypsithermal Interval when conditions were warmer

and drier than they are today (Webb 1981; Wright 1976; Deevey and

Flint 1957; Watts 1980; Watts and Stuiver 1980). In a few areas the

Hypsithermal lasted until about 5000 yr BP (Delcourt 1980; Watts 1979)

when the cooler, wetter climate of today was established.

By the end of the Hypsithermal, animals and plants should have

spread throughout climatically suitable areas. Reptiles are somewhat

limited by climate, so their distributions should have been more exten-

sive at the end of the Hypsithermal than they are today. The '^ooling

trend of the last 5000 to 7000 years should have resulted in local extinc-

tion of reptile populations as conditions became too severe, and in a

general contraction of species' ranges to those seen today. Special local

conditions, such as those of lower Wilson Creek or the gorges of the

Southern Blue Ridge escarpment, could forestall local extinctions of

reptiles by limiting the impact of the cooling trend (Billings and Ander-

son 1966; Anderson and Zander 1973; Bruce 1965). The result of this

protection would be isolated, relict populations in refugia well separated

from the main range of the species, as seen in the reptiles of lower Wil-

son Creek.

Lower Wilson Creek NC Reptiles 31

ACKNOWLEDGMENTS.— We are pleased to thank those whose

help was invaluable to the successful completion of this study: Messrs.

William Palmer and Alvin Braswell, North Carolina State Museum of

Natural History, gave us access to their unpublished notes on species

distributions. Dr. J. Frank Randall was supportive in the early stages of

the project. Mr. Bob Green allowed us to place the thermograph in a

protected area at Brown Mountain Beach. Mr. Pat Cook, District

Forest Ranger, supplied us with weather data for Wilson Creek. Deborah

Nicoletto, assistant city planner, City of Lenoir, provided weather data

for Lenoir. Computer time was donated by the Appalachian State Uni-

versity Computing Center. Janice Ashley typed the final draft of the

manuscript. An anonmyous reviewer provided helpful criticisms of the

manuscript.

LITERATURE CITED

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Blue Ridge Province and their phytogeographic relationship. J. Elisha

Mitchell Sci. Soc. 59(1-2): 15-60.

Billings, W. D., and L. E. Anderson. 1966. Some microclimatic characteristics

of habitats of endemic and disjunct bryophytes in the Southern Blue Ridge.

The Bryologist 69:76-95,

Bruce, Richard C. 1965. The distribution of amphibians and reptiles on the

southeastern escarpment of the Blue Ridge Mountains and adjacent pied-

mont. J. Elisha Mitchell Sci. Soc. 57(1): 19-24.

Conant, Roger. 1975. A Field Guide to Reptiles and Amphibians of Eastern and

Central North America. 2nd edition. Houghton Mifflin Co., Boston. 429

pp.

Delcourt, Paul A. 1980. Goshen Springs: Late Quaternary vegetation record for

southern Alabama. Ecology <57(2):37 1-386.

Deevey, E. S., Jr., and R. F. Flint. 1957. Postglacial hypsithermal interval.

Science 725:182-184.

Flint, Richard Foster. 1971. Glacial and Quaternary geology. John Wiley and

Sons, Inc., New York, xii + 892 pp.

Huheey, James E., and Arthur Stupka. 1967. Amphibians and Reptiles of Great

Smoky Mountains National Park. U. Tenn. Press, Knoxville. xii + 98 pp.

King, Willis. 1939. A Survey of the Herpetofauna of Great Smoky Mountains

National Park. Am. Midi. Nat. 27(3):53 1-582.

Martof, Bernard S., William M. Palmer, Joseph R. Bailey, Julian R. Harrison,

III and Jack Dermid. 1980. Amplibians and Reptiles of the Carolinas and

Virginia. U. North Carolina Press, Chapel Hill. 264 pp.

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

Acad. Sci. Monogr. No. 3. 346 pp.

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

Univ. Agric. Exp. Stn., Auburn. 347 pp.

Rohlf, F. James, and Robert R. Sokal. 1969. Statistical Tables. W. H. Freeman

and Co., San Francisco, xi + 253 pp.

32 Robert Wayne Van Devender and Paul F. Nicoletto

Ryan, Thomas A., Brian L. Joiner and Barbara F. Ryan. 1976. MINITAB Stu-

dent Handbook. Duxbury Press, North Scituate, MA. ix + 341 pp.

Sokal, Robert R., and F. James Rohlf. 1969. Biometry. W. H. Freeman and

Co., San Francisco, xxi + 776 pp.

Wade, Julia, and A. C. Echternacht. 1981. A comparative study of reproductive

cycles in two populations of Anolis carolinensis. Abstract. ASB (Assoc.

Southeast. Biol.) Bull. 25(2):96-97.

Watts, W. A. 1979. Late Quaternary vegetation of central Appalachia and the

New Jersey Coastal Plains. Ecol. Monogr. -^9(4): 427-469

1980. Late Quaternary vegetation history at White Pond on the inner

Coastal Plain of South Carolina. Quat. Res. 75:187-199.

, and M. Stuiver. 1980. Late Wisconsin climate of northern Florida

and the origin of species-rich deciduous forests. Science 270:325-327.

Webb, Thompson, IIL 1981. The past 11,000 years of vegetational change in

eastern North America. BioScience i7(7):501-506.

Weller, W. H. 1930. Records of some reptiles and amphibians from Chimney

Rock Camp, Chimney Rock, N. C. and vicinity. Proc. Junior Soc. Nat. Sci

7(8-9):(5 l-54)(pages unnumbered).

Wright, H. E., Jr. 1976. The dynamic nature of Holocene vegetation, a problem

in paleoclimatology, biogeography, and stratigraphic nomenclature. Quat.

Res. 5:581-596.

Accepted 15 April 1983

Comparative Food Studies of Yellowfin Tuna, Thunnus

albacares, and Blackfin Tuna, Thunnus atlanticus

(Pisces: Scombridae) from the Southeastern

and Gulf Coasts of the United States

Charles S. Manooch, III

AND

Diane L. Mason

National Marine Fisheries Service,

Southeast Fisheries Center, Beaufort Laboratory,

Beaufort, North Carolina 28516-9722

ABSTRACT. — Two hundred and six yellowfin tuna, Thunnus alba-

cares, and 98 blackfin tuna, T atlanticus, were sampled from sport

fisheries in the South Atlantic and Gulf of Mexico, April 1980 to July

1982. Stomach contents were analyzed by frequency of occurrence,

number of food items, and volume. Invertebrates (85%) and fish (77%)

occurred in the diet of yellowfin relatively equally. Major invertebrates

by frequency of occurrence were cephalopods (62%) and crustaceans

(52%). Fishes were represented primarily by the families Scombridae

(12.2%), Balistidae (11.2%), and Syngnathidae (8.2%). Yellowfin also

ingested floating materials such as plastic, feathers, seagrasses, and

balls of tar. Invertebrates occurred in 82% of the blackfin stomachs

with food, and represented 75% and 31% of the foods by number and

volume, respectively. Fish were found in 67% of the stomachs and con-

stituted 26% and 68% of the food number and volume, respectively.

The most frequently occurring invertebrates were crustaceans (67.4%)

and cephalopods (36.0%). Fishes were represented primarily by the

families Balistidae (10.1%), Trichiuridae (5.6%), and Carangidae (4.5%).

Blackfin also consumed floating materials, such as plastic and sea-

grasses. Statistical comparisons of the diets of the two species indicated

no significant correlation. Overall, their diets appear to reflect those of

fast, aggressive predators, and also of fish that use their gill apparatus

to strain small, near-surface items from the water.

INTRODUCTION

The family Scombridae includes many species of pelagic fish that

are very important to the world's fisheries. Some, such as the mackerels

Scomberomorus spp. and Scomber spp., are primarily coastal, migrat-

ing north in spring and summer and south in fall and winter. Others,

including members of the genus Thunnus, are usually much larger than

the mackerels and are reputed for their more complex, often transoce-

anic migrations.

Two species of Thunnus, the yellowfin tuna, T. albacares, and the

blackfin tuna, T. atlanticus, are highly esteemed food and sport fishes

whose distributions include the southeastern and Gulf coasts of the Uni-

Brimleyana No.9:33-52. June 1983. 33

34 Charles S. Manooch, III and Diane L. Mason

ted States. The yellowfin is the largest and more prized of the two,

attaining a weight of at least 176 kg (compared with 19 kg for blackfin).

On the whole, tuna landings in the western Atlantic are sporadic

and are much smaller than those made by the large-scale, international

hook and line and seine tuna fisheries that operate in the eastern Atlan-

tic and Pacific. The total United States commercial landings of all tunas

was 341,149,000 pounds in 1981, 326,860,000 pounds from the Pacific

and 14,289,000 from the Atlantic. Only 131,000 pounds were landed in

the South Atlantic Region — North Carolina, South Carolina, Georgia,

and the east coast of Florida (D. S. Fitzsgibbon, pers. comm.). Of the

South Atlantic total, only 5,000 pounds were identified as yellowfin

tuna, and none as blackfin, although the 55,000 pounds of unclassified

tunas undoubtedly included blackfin. Recreational catches of yellowfin

and blackfin tunas tend to be greater than the commercial catches for

the southeastern United States. In North Carolina, for instance, anglers

fishing from charter boats in 1978 caught approximately 151,000 pounds

of yellowfin tuna and 38,000 pounds of blackfin tuna (Manooch et al.,

1981). No information is available for 1981.

Considering the disproportionately large commercial catch of tunas

in the Pacific, it is not surprising that many publications pertaining to

life histories, population dynamics and exploitation have resulted from

research on species in that region. Relatively few studies have been con-

ducted on Atlantic stocks. Dragovich (1969) in his review of food stud-

ies on Atlantic tunas mentioned that the papers he read emphasized

the need for additional research on the foods and feeding habits of

Atlantic stocks. The limited information available from the western

Atlantic usually resulted from fish collected aboard scientific vessels

that did not operate along the southeastern or Gulf coasts of the United

States, or that operated well offshore of the normal sport fishing

grounds (Dragovich 1969, 1970).

To obtain more data pertinent to the management of pelagic

stocks, studies were initiated on oceanic species important to fisheries

along the southeastern and Gulf coasts of the United States. Our study

is the result of a cooperative effort that included the Oceanic Pelagic

Program, SEFC, Miami Laboratory, and the Bioprofiles Task, SEFC,

Panama City Laboratory. The objectives were to 1) identify the food

habits of yellowfin and blackfin tunas; 2) compare the diets of the spe-

cies collected from the same geographic area during the same period of

time; and 3) determine if changes in the diets occur for different sizes of

fish.

METHODS

Of the 206 yellowfin and 98 blackfin stomachs examined, 169 and

55, respectively, were from fish landed at Oregon Inlet or Hatteras,

Foods of Yellowfin and Blackfin Tuna 35

North Carolina during the spring, summer and fall of 1980, 1981 and

1982. A few additional samples, indicated in parentheses as yellowfin

and then blackfin, were obtained from locations along the southeast

Atlantic and Gulf of Mexico coasts: South Carolina (31,8), Georgia

(3,1), east coast of Florida (0,2), northwest Florida (3,1), Mississippi-

Louisiana (0,6), and south Texas (0,25).

Samplers at all locations apportioned their efforts to coincide with

local charter boat activities, primarily April through October. Port

samplers met boats at the docks as a day's catch was being unloaded.

Most fishermen either wanted to save their fish whole for mounting, or

to have them filleted and packed on ice or frozen upon returning to the

dock. Data were obtained only from the latter group, either in exchange

for cleaning the fish, or from fish cleaners who worked at local markets.

Fish were measured to the nearest millimeter (FL) and weighed to the

nearest tenth of a kilogram. Stomachs and gonads were placed in

labeled cloth bags or cheese cloth and preserved in 10% formaUn.

In the laboratory, stomach contents were identified to the lowest

possible taxon and were enumerated, thus providing the relative number

of each food type in the stomachs. Frequency of occurrence of materials

was determined by counting every stomach that contained at least one

specimen or part of a specific item (taxon). Empty stomachs were

excluded. The volume of each taxon was obtained by water displace-

ment and was later converted to weight by a linear regression equation.

Larval and juvenile fish in the stomachs were identified after they had

been cleared and stained following the methods discussed by Dingerkus

and Uhler (1977) and Taylor and Van Dyke (1978). Crustaceans were

identified by Steven G. Morgan and Joseph W. Goy, Duke University

Marine Laboratory, Beaufort, North Carolina. Parasites, encountered

only occasionally, were separated from food items, counted, identified

and preserved. A stomach containing only parasites was considered

empty.

All data were analyzed as percent frequency of occurrence, percent

of total number, and percent of food volume. Once frequencies, volumes

and numbers of the various foods were obtained, an index of relative

importance (IRI) was used to estimate the contribution of major food

groups to the diet (Pinkas et al., 1971). The index was calculated as: IRI

= (N + V) F, where N = numerical percentage of a food, V = its volumet-

ric percentage, and F = its percentage frequency of occurrence.

The Spearman rank correlation (r^) was used to evaluate differences

in diets of the two species based on IRI values of foods from fish col-

lected in the same geographic area and over approximately the same

period of time. Two different equations may be used. One, where there

are no ties (rankings are equal for two or more food categories), and the

36 Charles S. Manooch, III and Diane L. Mason

other where ties do occur. The equation for tied food categories (Fritz

1974) was used:

"^ - where Sx^ = — STx; Sy2 = zTy;

2 Sx2 Xy^ N -^ N

J - ^^ ' ^ ; N = numbers of ranks; d = difference between ranks; T =

N

correlation factor for ties and t = number of observations tied at a given

rank. Pearson and Kendall's Tau B Correlation Coefficients, in addition

to the Spearman rank, were also derived to evaluate differences in the

diets.

RESULTS AND DISCUSSION

Composition of Stomach Contents

Stomach contents of both species could be grouped into four prin-

cipal categories: fish, cephalopods, crustaceans and miscellaneous non-

food items (Tables 1, 2; Fig. 1). Major representatives of each group will

be discussed below under separate headings and will also be analyzed

later to identify differences in diets related to the species of predator and

its size. A graphic presentation of the overall contribution of selected

foods to the diet (IRI plots) is presented in Figure 2.

Fish. — Fishes occurred in 77% of yellowfin and 67% of blackfin stom-

achs that contained food (Tables 1, 2; Fig. 2) and consisted primarily of

older larvae and juveniles often associated with floating Sargassum. In

all, 23 famiHes were identified. Adult exocoetids, scombrids and syngna-

thids were found occasionally in yellowfin, as were syngnathids, serran-

ids, sciaenids and stromateids in blackfin. For all life stages, fish that

occurred most frequently in yellowfin tuna were Scombridae (12.2%),

Balistidae (11.2%), Syngnathidae (8.2%), Diodontidae (5.1%) and Exo-

coetidae (4.6%). Fifty-three percent of stomachs with food contained

unidentifiable fish remains. Fish that occurred most often in blackfin

tuna stomachs were Balistidae (10.1%), Trichiuridae (5.6%), Carangidae

(4.5%) and Syngnathidae (4.5%). Unidentifiable fishes were found in

44.9% of the stomachs containing food.

Cephalopods. — Cephalopods constituted almost all the molluscan

food of both species. One exception was unidentifiable mollusk tissue,

possibly cephalopod, from a yellowfin captured in the Gulf of Mexico.

Two groups were represented: Teuthidida and Octopodida. Teuthoids

(squids) were the most important by frequency of occurrence and by

volume: 50.5% and 41.0% for yellowfin, 31.5% and 21.5% for blackfin.

By comparison, octopodids, represented by the paper nautilus, Argo-

nauta argo, appeared in only 7.7% of the yellowfin tuna and 3.4% of the

Foods of Yellowfin and Blackfin Tuna

37

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46 Charles S. Manooch, III and Diane L. Mason

blackfin tuna. Percent volumes of these mollusks were less than 2% for

both predators, and whereas over 430 squid were consumed by the

tunas, less than 30 paper nautilus were eaten. At least three genera of

squids were ingested: Loligo, Sepioteuthis, and Illex. Generic identifica-

tions were obtained by comparing saved, pooled samples with reference

collection specimens and therefore do not appear in the tables.

Crustaceans. — Crustaceans, important foods of both species and

second only to fish in overall frequency of occurrence, were identified in

52% of the yellowfin and in 67.4% of the blackfin. The majority were

immature stages (larvae, megalopa and glaucothoe). Due to the small

sizes of the animals, the relative percentages of the total food volume —

5.9% for yellowfin and 8.4% for blackfin — were comparatively small.

Major taxa in the diet of yellowfin by frequency of occurrence were

Raninidae (27.5%), Penaeidae (12.2%), Stomatopoda (7.7%), Portuni-

dae (7.1%), and Dromiidae (6.1%). For blackfin tuna, the most fre-

quently encountered were Stomatopoda (34.8%), Diogeninae (16.9%),

Raninidae (15.7%), Penaeidae (14.6%), and Dromiidae (12.4%). In all,

over 5,000 individuals were enumerated, and on one occasion a single

predator contained hundreds of these small, mesopelagic invertebrates.

Our findings of the overall food habits seem to agree closely with

those of Dragovich (1970), who described fish, cephalopods and crusta-

ceans as the major foods of yellowfin and skipjack, Katsuwonus pela-

mis, tunas in the Atlantic. He also mentioned that larval and juvenile

stages were prevalent for ingested fishes and macrozooplanktonic crus-

taceans.

Miscellaneous. — The very nature of tuna feeding, near-surface strain-

ing as well as actively pursuing and capturing larger animals, results in a

variety of items being consumed that are probably ingested by accident

along with natural foods. Yellowfin tuna had the most diverse assemb-

lage of non-food items (31.6% frequency): plants {Sargassum, Zostera,

Thalassia and Spartina), feathers, globs of tar, and plastic. Miscellane-

ous items occurred in only 15.7% of the blackfin, represented by Sargas-

sum, Zostera and plastic. Sargassum was found in 26.5% of the stom-

achs with food, and usually occurred in tunas captured off North

Carolina. This percentage is similar to the 37.8% reported for Sargas-

sum removed from the digestive tracts of skipjack tuna captured earlier

from approximately the same geographical area (Batts 1972).

Other studies also revealed a dominance of fish, squid and crusta-

ceans in tuna diets for the Atlantic and Pacific. Reintjes and King

(1953) investigated the food habits of 1,097 yellowfin from the Central

Pacific and found that fish occurred in 70.4% of the stomachs; squid in

55.4%; and crustaceans (mostly immature, pelagic stages) in 66.9%.

Alverson (1963) found fish, squid and crustaceans occurring in 53.8%,

Foods of Yellowfin and Blackfin Tuna 47

23.9% and 76.1%, respectively, of the yellowfin he examined from the

Pacific. Similar occurrences were reported for yellowfin from the Atlan-

tic (Dragovich 1970), and for skipjack tuna (Alverson 1963; Nakamura

1965; Batts 1972); bluefin tuna, T. thynnus (Pinkus et al., 1971); and

albacore, T. alalunga (Pinkas et al., 1971) from the Pacific.

Comparative Diets

Since temporal and spatial variations in the diets were so great

(data collected over a period of three years, and from several widely

different geographical locations), we believed that only by analyzing

small, discrete samples could we detect important differences in them.

To achieve this, we used only stomach contents of the two species col-

lected together off Oregon Inlet on 10 different days from May through

September 1981 (Table 3).

Index of Relative Importance. — Indexes of Relative Importance

(IRI), which present the combined contributions of volume, frequency

of occurrence, and numbers of each food item to the diet (Table 3),

showed that, surprisingly, invertebrates were very important foods for

both species. The first five categories (ranks) for yellowfin were Teuthid-

ida (squids), unidentifiable fish, Raninidae, Scombridae, and unidentifi-

able crustaceans. For blackfin they were unidentifiable fish, Teuthidida,

Raninidae, Stomatopoda, and unidentifiable crustaceans. Obvious dif-

ferences were more clupeids and unidentifiable diogenid crabs in black-

fin, and more scombrids and squids in yellowfin. Other items were also

different, but their respective IRI values were relatively small (i.e., exo-

coetids for yellowfin = 9.7, for blackfin = 0.0).

Correlation Coefficients. — Data from Table 3, ranked by IRI values,

were used to obtain quantitative comparisons of local food habits of the

two species. Three different measures were used: Spearman Rank Corre-

lation Coefficient (Fritz 1974); Kendall Rank Correlation Coefficient

(Bray and Ebeling 1975); and Pearson Product-moment Correlation

Coefficient (Goodall 1973). The first two require no assumption of

normality with regard to the distribution of the two predator species,

whereas the latter does. Cailliet and Barry (1978), who compared the

three methods of analyzing diets that have different distributions of prey

items, found that Spearman and Kendall correlation coefficients are

somewhat unpredictable when there are 1) a large number of ties, 2) a

considerable nonoverlap of prey items, and 3) high prey richness and

evenness (i.e., diversity). They felt that the Pearson method was best.

Although our data have a fairly low richness and evenness, there are

relatively few ties (2 for yellowfin, 3 for blackfin) and there is a fairly

good overlap in the diets. For these reasons all three methods of meas-

uring diet similarity are probably appropriate. Qualitatively, both spe-

cies feed extensively on epipelagic and mesopelagic fishes and inverte-

48

Charles S. Manooch, III and Diane L. Mason

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Foods of Yellowfin and Blackfin Tuna 49

brates. Eleven of the 28 food categories occurred in the stomachs of

both species, and 6 of the 10 most important categories to blackfin also

ranked in the top 10 for yellowfin. The obvious conclusion is that both

species have similar diets when they occur together off the coast of

North Carolina. Statistically, however, the correlation coefficients were

all nonsignificant at the 0.05 (0.344; 29 dO level. The correlation coeffi-

cients were: Spearman, 0.2273; Kendall, 0.1451; and Pearson, 0.2273.

Comparative Diets by Predator Size

Differences in stomach contents by fish size may of course be

attributable merely to the availability of food in the environment, but

they may also be attributable either to a change in food preference, or

to the ability of the predator to capture and swallow certain organisms

as it increases in size. Our objectives of comparing diets by tuna size

were to determine if near-surface feeding was related to tuna size and to

ascertain if basic changes in the diets occurred as the fish grew larger.

Different studies throughout the world's oceans generally suggest

that as tunas grow larger, their diets change. Reintjes and King (1953)

reported that the overall high occurrence of crab larvae, stomatopod

larvae, squid, and juvenile fishes indicates a preference by Pacific yel-

lowfin tuna for small food items. These authors further explained that

small tuna feed predominantly on crustacean larvae; medium-size fish

feed on fish, crustacean larvae, and squid; and large yellowfin mainly

consume fish and squid. These findings were substantiated by Nakam-

ura (1965) and Batts (1972) for skipjack tuna whose diets reflected a

decline in crustaceans and a subsequent higher percentage of fish, as

tuna size increased.

To accomplish our evaluations we first grouped the fish into size

classes (mm FL) (Tables 4, 5). Next, selected food groups — fish, adult

fish, juvenile fish, invertebrates, squid, larval crustaceans and plants —

were established to demonstrate food size (i.e., adult fish vs. larval crus-

Table 4. Selected food items consumed by different sized yellowfin tuna,

expressed as percent frequencies of occurrence.

Fish size (mm FL)

Contents 501-700 701-900 901-1100 >1100

Fish

Adult fish

Juvenile fish

Invertebrates

Squid

Larval crustaceans

Plants

50 Charles S. Manooch, III and Diane L. Mason

Table 5. Selected food items consumed by different sized blackfin tuna,

expressed as percent frequency of occurrence.

taceans) and materials that we believed to be consumed on or near the

surface (i.e., floating plants). Contents are presented as percent fre-

quency of occurrence (Tables 4, 5).

Yellowfin Tuna. — Size of food items showed little change as fish size

increased or decreased (Table 4). The three key food categories — adult

fish, juvenile fish, and larval crustaceans — neither steadily increased

nor decreased in occurrence as tuna size increased. This finding is con-

trary to that of Dragovich (1970), who found that the frequency occur-

rence offish in stomachs of yellowfin increased with fish size. However,

he discovered no relationship between squid in the diet and tuna size. In

our study, the occurrence of floating plants decreased for the larger size

classes, indicating that perhaps smaller individuals fed more extensively

near the surface.

Blackfin Tuna. — The size of prey items and feeding proximity to the

surface appeared to change with fish size. As fish size increased, large

food items (i.e., adult fish) generally occurred more frequently, and

small food items (i.e., larvel crustaceans and juvenile fish) occurred less

frequently (Table 5). Surface feeding, as suggested by the incidental

ingestion of floating plants, decreased as fish attained larger sizes.

Volumes of Contents Related to Species and Fish Body Weight

Since the quantity and types of foods ingested by fishes are often

converted into caloric equivalents for energetics studies, we present fre-

quencies of the range of food volumes for the two species (Table 6). The

displacement volume for yellowfin averaged 67.9 ml (72.2 g), compared

with 28.6 ml (29.6 g) for blackfin tuna. Volumes of stomach contents of

yellowfin and blackfin varied from 0.1 to 745.0 ml and from 0.1 to 257.5

ml, respectively. The largest volumes were found in a 40 kg yellowfin

and an 8.8 kg blackfin. The volume range for yellowfin from the Pacific

was similar, 0.1 to 1,000 ml (Reintjes and King, 1953). The extremes in

our data were much greater than those described by Dragovich and

Potthoff (1972): 0.1 to 20.0 for skipjack, and 0.1 to 60.0 ml for yellow-

fin tunas collected off the west coast of Africa. In our study, approxi-

Foods of Yellowfin and Blackfin Tuna 51

Table 6. Frequencies of food volumes by species of tuna.

mately 33% of the yellowfin had food volumes exceeding 50 ml, a pro-

portion similar to that of the 29% found by Reintjes and King (1953).

By comparison, Dragovich (1970) noted volumes of less than 20 ml for

85% of the yellowfin from the Atlantic. We found that only 19% of the

blackfin, a much smaller species, had contents over 50 ml.

To determine the relationship of volume to fish body weight, we

first derived the following equation for converting volume in ml to

volume in grams:

Volg = -1.4009 + 1.0846 (Volj^i), N=25, r=0.999.

Comparisons were then made between estimates of stomach contents

and the body weights of some of the tunas selected at random. Percent-

ages of food weight to fish weight varied from trace (<0.002) to 2.02

for yellowfin, and from 0.02 to 3.20 for blackfin tuna. Qnly 10% of the

yellowfin had contents exceeding 1% offish body weight, whereas 20%

of the blackfin tuna had contents exceeding this percentage. Usually our

observations were well below 1%, as were those of Dragovich (1970).

In summary, yellowfin and blackfin tuna appear to be fast, aggres-

sive predators capable of capturing swift, relatively large prey. On the

other hand, they use their gill apparatus to strain small, near-surface

items from the water. During feeding, non-food materials (inorganic as

well as organic) are ingested, probably incidental to normal prey. The

variability of specific food organisms within the major categories (fish,

cephalopods, and crustaceans) in the diets suggests that tunas are non-

selective feeders. This is undoubtedly a factor in their wide geographic

52 Charles S. Manooch, III and Diane L. Mason

distribution, and one would expect, therefore, for the diets of such well-

traveled fish to be rather cosmopolitan.

LITERATURE CITED

Alverson, Franklin G. 1963. The food of yellowfin and skipjack tunas in the

eastern tropical Pacific Ocean. Inter-Am. Trop. Tuna Comm. Bull.

7:295-396.

Batts, Billy S. 1972. Food habits of the skipjack tuna, Katsuwonus pelamis, in

North Carolina waters. Chesapeake Sci. 7i: 193-200.

Bray, Richard N., and A. W. Ebeling. 1975. Food, activity and habitat of three

"picker-type" microcarnivorous fishes in the kelp forests off Santa Barbara,

California. Fish. Bull. U.S. 75:815-829.

Cailliet, Gregor, and J. P. Barry. 1978. Comparison of food array overlap

measures useful in fish feeding habit analysis. Pp. 67-79 in S. Lopovsky and

C. Simenstad (eds.). Fish food habits studies. Proc. Second Pacific

Northwest Tech. Workshop, Univ. Washington, Seattle.

Dingerkus, G., and L. D. Uhler. 1977. Enzyme clearing of alcian blue stained

whole small vertebrates for demonstration of cartilage. Stain Technol.

52:229-232.

Dragovich, Alexander. 1969. Review of studies of tuna food in the Atlantic

Ocean. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 593. 21 pp.

1970. The food of skipjack and yellowfin tunas in the Atlantic

Ocean. Fish. Bull. U.S. 65:445-460.

, and T. Potthoff. 1972. Comparative study of food of skipjack and

yellowfin tunas off the coast of West Africa. Fish. Bull. U.S. 70:1087-1101.

Fritz, Eugene S. 1974. Total diet comparison in fishes by Spearman rank

correlation coefficients. Copeia 1974(1):210-214.

Goodall, David W. 1973. Sample similarity and species correlation. Pp. 106-156

in R. H. Whittaker (ed.). Ordination and Classification of Communities.

Junk, The Hague.

Manooch, Charles S., Ill, L. E. Abbas and J. L. Ross. 1981. A biological and

economic analysis of the North Carolina charter boat fishery. U.S. Natl.

Mar. Fish. Serv. Mar. Fish. Rev. ^i(8):l-ll.

Nakamura, Eugene L. 1965. Food and feeding habits of skipjack tuna {Katsu-

wonus pelamis) from the Marquesas and Tuamota Islands. Trans. Am.

Fish. Soc. 9^:236-242.

Pinkas, Leo, M. S. OHphant and I. L. K. Iverson. 1971. Food habits of alba-

core, bluefin tuna, and bonito in California water. Calif. Dep. Fish Game

Fish Bull. 752:1-105.

Reintjes, John W., and J. E. King. 1953. Food of yellowfin tuna in the Central

Pacific. U.S. Fish Wildl. Serv. Fish. Bull. 5¥:91-110.

Taylor, William R., and G. C. Van Dyke. 1978. Unpublished manuscript.

Staining and clearing small vertebrates for bone and cartilage study.

Smithsonian Institution, Washington, DC. 19 pp.

Waldron, Kenneth D., and J. E. King. 1963. Food of skipjack in the Central

Pacific. FAO Fish. Rep. 6(3): 143 1-1457.

Accepted 17 September 1983

Benthic Macroinvertebrates of Cane Creek, North Carolina,

and Comparisons with Other Southeastern Streams

David R. Lenat

Biological Monitoring Group,

N.C. Division of Environmental Management,

Archdale Building, Raleigh, North Carolina 27611

ABSTRACT.— The macrobenthos of Cane Creek, in the Piedmont

Plateau of North Carolina, have been sampled by several investigators.

This information was combined to generate a list of 272 invertebrate

taxa. Cane Creek is compared to other unstressed Piedmont streams to

define characteristics of a "normal" stream in this geographic area. If

used cautiously, this data set can provide control information for bio-

logical monitoring. Average taxa richness appears to be the best tool

for environmental assessment work. It shows little variability across a

wide range of North Carolina streams, even outside the Piedmont.

Such a pattern suggests a constant number of niches in stream ecosys-

tems.

INTRODUCTION

The study of pollution in freshwater ecosystems is a complex prob-

lem. Water quality degradation may be caused by an immense number

of pollutants, many of which have an alarming degree of temporal

and/ or spatial variability. To deal with this variability, water quality

monitoring often includes some biological sampling.

The North Carolina Division of Environmental Management

(DEM) has used biological monitoring to analyze a wide variety of

water quality problems (Penrose et al. 1980). Specifically, the Division's

biologists use the structure of the benthic macroinvertebrate community

to detect stress in aquatic systems. There are many ways to examine

such data (Lenat et al. 1980), but all are based on comparisons of actual

data with some expected pattern. The expected pattern is often derived

from a control area, but in many situations it may be difficult to locate

good control stations. This difficulty can often be overcome by using

control data sets. The Division's Biological Monitoring Group has

attempted to generate control data sets by compiling information from

many unpolluted North Carolina streams and rivers. An earlier contri-

bution examined the benthos of a Mountain river system (Penrose et al.

1982). This paper describes the benthic macroinvertebrates of a typical

Piedmont stream.

Brimleyana No.9:53-68. June 1983. 53

54

David R. Lenat

STUDY SITE

Cane Creek is a third-order stream located in Orange County,

North CaroHna (Fig. 1). The total watershed is about 90 km^ and aver-

age discharge is roughly 0.7 m^s (N.C. Division Environmental Man-

agement 1975). Cane Creek is classified as A-II water, i.e. suitable for

drinking (after treatment), body contact, recreation, and "fish and wild-

life propagation".

ci:

c^

Bear Creek

Saxapahaw

White Cross

N

Orange Co.

Chatham c7. -

Fig. 1. Sampling Stations, Cane Creek, North CaroHna.

Benthic Macroinvertebrates Cane Creek NC 55

Fish collections from Cane Creek (unpublished data on file with

N.C. Wildlife Resources Commission) include 25 species. The list appears

typical of Piedmont streams in North Carolina.

The watershed contains both forest and agriculture, principally

dairy farming and row crops. According to the water basin plan (N.C.

Division Environmental Management 1975), there are no point source

discharges in this area, but runoff often causes high turbidity in Cane

Creek. In 1978 the Soil Conservation Service (unpubUshed) cited Cane

Creek as a high priority area for land treatment to reduce erosion rates.

However, little accumulation of coarse bedload sediment was noted in

biological surveys, which may be due to local geology. Cane Creek is

located in the "slate belt" of North Carolina, a zone of metamorphosed

volcanic rock (Simmons and Heath 1979). DEM surveys within this

land type (unpublished data) suggests that little "sandy" stream sedi-

ment is produced through erosion.

METHODS

Several investigators have collected benthic macroinvertebrates from

Cane Creek (Smock and Hughes 1975; Mozley 1978; Penrose et al.

1980). Extensive collections have also been made by the author and by a

limnology class at North Carolina State University. Collection methods

included Hester-Dendy muhiple plate samplers (FuUner 1971), "kicks"

(Frost et al. 1971), and various qualitative techniques. The most inten-

sive collections have been at Station 1 (Lower Cane Creek), but all areas

of the Cane Creek watershed have been sampled (see Fig. 1). Areas

sampled included temporary streams and stream orders 1 through 3. All

records were vertified by the author.

RESULTS AND DISCUSSION

Taxa List

Table 1 lists 272 taxa collected from Cane Creek. A list of taxo-

nomic references used for identification of these organisms is available

from the author. This table also contains data on frequency (rare, com-

mon, or abundant), a classification that is somewhat subjective since

most collections were qualitative. The list contains few species not col-

lected in other Piedmont streams (DEM, unpubl. surveys). The most

unusual record was Mystacides alafimbriata Hill-Griffin, a common

edge species found at several stations in July 1979. This caddisfly had

not been collected east of the Mississippi River. Identification was based

on young larvae and should be confirmed by collection of adults.

Another unusual caddisfly record was Dibusa angata Ross, a species

strongly associated with red algae (Wiggins 1977).

The turbellarian Hydrolimax grisea Haldeman was collected sev-

eral times in Cane Creek. Pennak (1978) listed this species as rare and

56 David R. Lenat

Table 1. List of benthic macroinvertebrates from Cane Creek. Under frequency,

A = abundant, C = common, R = rare.

Taxon Frequency

EPHEMEROPTERA

Baetis amplus A

B. flavistriga A

B. intercalaris A

B. pluto C

B. propinquus R

Callibaetis sp. R

Centroptilum sp. R

Cloeon alamance R

Pseudocloeon spp. C

Caenis cf. diminuta C

Ameletus lineatus C

Isonychia spp.' C

Siphloplectron basale R

Leptophlebia sp. C

Paraleptophlebia sp. A

Hexagenia munda C

Ephemerella (E. ) catawba^ R

E. {Attenella) attenuata R

E. (Dane I la) simplex R

E. {Seratella) deficiens C

E. (Eurylophella) bicolor C

E. (E.) temporalis C

E. (E.) funeralis \ R

Heptagenia aphrodite C

Stenonema modestum^ A

S. smithae C

S. vicarium C

S. (femoratum) R

Stenacron interpunctatum A

S. pallidum C

PLECOPTERA

Allocapnia spp. C

Leuctra sp. R

Acroneuria abnormis C

/I. evoluta R

Eccoptura xanthenes R

Perlesta placida C

Taeniopteryx metaqui A

T. burksi A

Strophoteryx fasciata A

Amphinemura sp. R

Isoperla clio C

Isoperla namata R

Hastaperla brevis R

Benthic Macroinvertebrates Cane Creek NC 57

HEMIPTERA

Belastoma fluminea R

Sigara spp. A

Gerris remigis C

Limnogonus sp. R

Trepobates sp. R

Metrobates hesperius R

Rheumatobates palosi R

Mesovelia mu Isanti R

Rhagovelia obesa C

Microvelia americana C

NEUROPTERA

Climacia sp. R

MEGALOPTERA

Corydalus cornutus C

Nigronia serricornis C

Chauliodes pectinicornis R

5/fl/w A

ODONATA

/4r^/fl spp. C

A. sedula

A. trans lata

A. moesta

A. tibialis

Enallagma spp."* C

Ischnura spp. C

Calopteryx sp. C

Baesiaeschna Janata R

Boyeria vinosa C

Cordulegaster sayi R

Helocordulia selysii R

Neurocordulia obsoleta R

Epitheca cynosura R

Libellula sp. R

Perithemis tenera R

Macromia allegheniensis C

Didymops transversa R

Gomphus spp C

Lanthus parvulus R

Stylogomphus albistylus R

Hagenius brevistylus . R

Dromogomphus spinosus C

TRICHOPTERA

Diplectrona modesta C

Cheumatopsyche spp. A

Hydropsyche betteni A

Macronema Carolina C

Nectopsyche sp. R

58 David R. Lenat

Ceraclea ancylus C

C. tarsipunctata R

Oecetis spp.^ C

Triaenodes injustus R

Triaenodes tardus R

r. cf. sp. b C

Mystacides alajimbriata C

Dibusa angata R

Stactobiella sp. R

Pycnopsyche guttifer R

P. gentilis R

Hydatophylax argus R

Neophylax cf. oligius R

Ptilostomis sp. R

Brachycentrus sp. R

Polycentropus spp.^ C

Phylocentropus sp. C

L>'/7e diversa R

Molanna blenda R

Chimarra cf. aterrima C

Wormaldia sp. R

Psilotreta sp. R

Lepidostoma sp. R

Rhyacophila Carolina R

/?. acutiloba R

/?. /^^ro R

COLEOPTERA

Helichus fastigiatus C

Ancyronyx variegata C

Macronychus glabratus C

Stenelmis spp. C

Oulimnius latiusculus R

Optioservus ovalis R

Dubiraphia quadrinotata C

Ectopria nervosa R

Psephenus herricki C

Anchy tarsus bicolor R

Hydroporus spp. C

Hydrovatus sp. II R

Rhantus sp. R

Tropisternus sp. R

Helophorus sp. R

Laccophilus sp. R

Copelatus glyphicus R

Dineutes sp. C

Gyrinus sp. R

Benthic Macroinvertebrates Cane Creek NC 59

DIPTERA (Miscellaneous)

Palpomyia (complex) C

Anopheles punctipennis C

Culex restuans R

Chaoborus punctipennis R

Dolichopodidae R

Empididae R

Simulium vittatum A

Prosimulium mixtum A

P. rhizophorum C

Chrysops sp. C

Tabanus sp. R

Antocha sp. - C

Dicranota sp. R

Hexatoma sp. R

Limonia sp. R

Pseudolimnophila sp. R

Tipula sp. R

r. abdominalis C

£)ixa sp. R

DIPTERA: CHIRONOMIDAE

Chironomus sp. R

Cryptochironomus blarina R

C fulvus gr. R

Demicryptochironomus sp. R

Dicrotendipes nervosus R

D. neomodestus A

Glyptotendipes sp. R

Kiefferulus dux R

Microtendipes pedellus A

A/, nr. rydalensis R

Paratendipes albimanus C

Phaenopsectra sp. R

P. flavipes C

Polypedilum aviceps R

P. convictum C

P.fallax R

P. illinoense C

P. scalaenum R

Stenochironomus sp. C

Stictochironomus sp. R

Thbelos jucundus C

Xenochironomus xenolabis R

Cladot any tarsus spp. R

Const empellina sp. R

Micropsectra sp. R

60 David R. Lenat

Paratanytarsus sp. ' R

Rheot any tarsus spp. A

Tanytarsus spp. A

T. guerlus gr. C

T. nr.glabrescens C

T. glabrescens R

Zavrelia sp. R

Ablabesmyia mallochi R

A. ornata C

A. parajanta R

Clinotanypus pinguis C

Conchapelopia group C

Labrundinia neopilosella R

L. nr. virescens , C

Larsia sp. R

Natarsia sp. R

Nilotanypus sp. R

Procladius bellus R

P. sublettei R

Psectrotanypus dyari R

Zavrelimyia sp. R

Sympotthastia sp. R

Briilia spp. '' R

Xylotopus par C

Corynoneura spp. C

Cardiocladius sp. R

Cricotopusj Orthocladius %r.

Cricotopus (C) bicinctus C

C (C) tremulus gr. sp. 1 R

(=C infuse at us)

C. (C.) tremulus gr. sp. 2 R

C. (C.) cf. cylindraeeus C

Orthoeladius (O.) robaeki R

O. (O.) nr. dorenus C

O. (O.) cf. obumbratus €

O. (O.) cf. nigritus R

O. (O.) nr. c/arA:e/ R

O. (Euorthoeladius) sp. 1 R

a (£.) sp. 2 » C

Diploeladius eultriger C

Eukiefferiella elaripenis gr. R

Tretenia bavariea gr. R

r. diseoloripes gr. R

Heterotrissoeladius marcidus R

Hydrobaenus spp. R

Nanoeladius spp. C

Genus nr. Nanoeladius R

Benthic Macroinvertebrates Cane Creek NC

61

Parakiefferiella sp. 1

P. sp. 3

P. nr. triquetra

Paraphaenocladius sp. T

Paracricotopus sp.

Parachaetocladius sp.*

Pseudosmittia sp.

Psectrocladius sp.

Rheocricotopus cf. robacki

Synorthocladius sp.

Thienemaniella sp.

MOLLUSCA

Somatogyrus sp.

Ferrissia rivularis

Phy sella sp.

Stagnicola sp.

Gyraulus sp.

Heliosoma anceps

Elimia sp.

Campeloma decisum

Eupera cubensis

Pisidium spp.

Sphaerium simile

Elliptio camplanata

E. icterina

Strophitus undulatus

CRUSTACEA

Cambarus acuminatus

Procambarus acutus

Palaemonetes paludosus

Lirceus sp.

Crangonyx spp.

Hyallela azteca

OLIGOCHAETA

Aulodrilus pigueti

A. pluriseta

Limnodrilus hoffmeisteri

Ilyodrilus templetoni

Peloscolex variegatus

Branchiura sowerbyi

Nais bretscheri

N. variabilis

Slavinia appendiculata

Stylaria lacustris

HIRUDINEA

Helobdella elongata

Placobdella multilineata

62 David R. Lenat

P. papillifera I R

Mooreobdella tetragon R

TURBELLARI

Cura foremanii C

Dugesia tigrina C

Hydrolimax grisea R

BRYOZOA

Plumatella repens C

Fredericella sultana R

MISCELLANEOUS

Porifera: Eunapius sp. C

Nemertea: Prostoma graecens R

Hydracarina C

Nematoda C

' Includes Isonychia bicolor.

^ Subgenera listed for Ephemerella are considered as genera by some authors.

^ Old identifications of Lewis (1974) have been revised in accordance with Bed-

narik and McCafferty (1979).

■* Includes Enallagma divergens.

^ Includes Oecetis cf. cinerascens.

^ Probably includes Cernotina sp.

^ This species listed as Parametriocnemus in many other surveys.

Near Pseudorthocladius; identified by Len Ferrington.

"reported from New Jersey and east Pennsylvania". This interpretation

of range probably results from taxonomic difficulties rather than rarity.

I have found this species to be widespread in Piedmont and Coastal

Plain streams, especially in the latter.

Several common Piedmont macroinvertebrates were rare or absent

in Cane Creek. Ephemerella catawba Traver, which I have found to be a

highly abundant organism in most Piedmont streams, was collected

only once. This mayfly prefers sand-gravel areas, a rare habitat in Cane

Creek. The lack of sand substrates probably also accounts for the

absence of Baetisca Carolina Traver, Progomphus obscurus Rambur,

and Robackia demeijerei (Kruseman).

Taxa collected in first-order tributaries were markedly different

from the fauna at lower stream stations. Many of these first-order

stream taxa were more typical of small mountain streams: Diplectrona

modesta Banks, Ephemerella funeralis McDonnough, Amphinemura

sp., Micropsectra sp., and Heterotrissocladius sp. This may reflect the

colder water temperature normally found in headwater areas (Vannote

and Sweeney 1980). Other taxa collected only in first-order tributaries

included Paraleptophlebia sp., Eccoptura xanthenes (Newman), Pyc-

nopsyche gentilis (McLachlan), and Molanna blenda Sibley.

Benthic Macroinvertebrates Cane Creek NC

63

Total Taxa Richness

Table 2 presents Cane Creek taxa richness by group, and includes

data from several other southeastern Piedmont stream investigations.

These data show a remarkable degree of consistency if allowances are

made for geographic area, collection methods, and special interest of the

investigators. Some differences, as seen in South Carolina and Georgia

samples, are attributable to collections of nonaquatic adult insects. This

usually increases estimates of species richness, as greater taxonomic pre-

cision may be attained with adult specimens.

Collections in semiaquatic areas also may increase the number of

species collected, especially for Coleoptera. Holeski and Graves (1978)

found that 30+ species of "shore beetles" were usually found at most

stations, but that this type of data was not useful in assessing environ-

mental stress. Eighty chironomid taxa were identified from Cane Creek,

about 30% of the total fauna. Use of adults or pupal exuviae might have

doubled this figure. Coffman (1973), using pupal exuviae, identified 143

chironomid taxa from Linesville Creek, while larval sampHng from the

same area produced only 77 taxa.

Table 2. Total taxa richness at Cane Creek, with selected data from other intensive stream

investigations. Numbers in parentheses omit data based only on nonaquatic

adults. NI = Not Identified.

This study.

White and Fox (1980), based largely on the collections of Paul Carlson.

White and Fox (1980).

Morse etal. (1980).

Coffman etal. (1971).

Includes many semiaquatic species.

64 David R. Lenat

Table 3. Average taxa richness by group for North Carolina Piedmont streams. Ranges

are rounded to integer values.

Average Taxa Richness/ Percent Density

Environmental assessment is often based on quantitative data from

a single collection or on data averaged over several collections. It may

be difficult to relate these data to water quality if good control informa-

tion is lacking. Tables 3 and 5 present average taxa richness and density

values from Cane Creek and make comparisons with five other rela-

tively unstressed Piedmont streams. All samples were taken during

DEM investigations. Collection methods (kicks) and identification

techniques were identical for all streams. From this data set, one may

attempt to define the normal characteristics of Piedmont stream

macroinvertebrate communities.

Table 3 gives average taxa richness values in the range of 43 to 56

(x = 48.5). There was often a remarkable constancy at the group level.

For example, Ephemeroptera varied only from 8.3 to 9.4. Plecoptera

values were generally in the range of 2 to 4, except at 4-Mile Creek. The

greater number of Plecoptera in this stream, which is at a higher eleva-

tion, may reflect colder water temperatures. The expected number of

Odonata is close to the average (1.4), except at Cates Creek. This is a

very narrow, slow stream and might be expected to show a high propor-

tion of bank associated (edge) species in quantitative samples. Odonata

are most frequently collected in bank areas. Variations in Coleoptera

and Trichoptera, especially the latter, are not easily explained. Various

regional differences in water chemistry, temperature, and gradient may

be responsible. The number of Diptera, especially chironomids, had a

Benthic Macroinvertebrates Cane Creek NC 65

wide range (14-21), but was always high. Even stressed areas may have a

diverse chironomid fauna (Penrose et al. 1980) although tolerant species

will become dominant. The "other" category includes miscellaneous

Insecta (Neuroptera, Lepidoptera), Turbellaria, Hirudinea, Nemertea,

Porifera, and Nematoda. Variation in this category is very unpredictable.

Table 4 presents average taxa richness values for streams in three

broad geographic areas in North Carolin: Mountains (data expanded

from Penrose et al. 1982); Piedmont (as in Table 5); and Coastal Plain.

These three physiographic regions have differing physical characteris-

tics. Going from the mountains to the coast one would expect increasing

water temperature, lower gradient, and increasing amonts of sand and

silt. The Coastal Plain data set is based on collections in Craven, Hert-

ford, and Northampton counties. These coastal streams are not entirely

unstressed, but the data are adequate to illustrate geographic trends.

The overall trend suggests a relatively constant number of species

in stream environments. Average taxa richness for Coastal Plain and

Mountain streams is well within the range expected for Piedmont

streams (Table 3). From this pattern one might advance the hypothesis

that unpolluted streams have a relatively constant number of niches.

Furthermore, these data imply that taxa richness is an excellent moni-

toring tool across a wide range of stream types. A similar hypothesis

was advanced by Patrick (1975:448). She examined species richness in

nine different rivers and concluded that "similar-sized areas of different

streams support similar numbers of species." This constancy of "a -

diversity" prevailed even when the number of species shared between

streams was low.

Table 4. Average taxa richness by group for Coastal Plain, Piedmont and

Mountain streams in North Carolina. See text for data sources.

Number of streams shown in parentheses.

66 David R. Lenat

At the group level, taxa richness varies considerably across the

three types of streams. Each stream type has a different assemblage of

invertebrate predators. Plecoptera and Trichoptera are most diverse in

Mountain streams, Megaloptera in Piedmont streams, and Odonata in

Coastal Plain streams. The "other" category, also most important in

Coastal Plain streams, includes many other predators: Hirudinea, Tur-

bellaria, and Prostoma graecens (Bohmig). Shifts may also be observed

in the collector-gatherer groups. Going from the mountains toward the

coast, Ephemeroptera-Trichoptera decline and are replaced by Coleop-

tera (Piedmont only), Oligochaeta, Crustacea, and Mollusca.

Table 5 shows density (expressed as a percentage of total density)

for Cane Creek and five other North Carolina Piedmont streams. The

density values show much greater between-stream variability than does

average taxa richness. These data may serve as controls only if used

with extreme caution. Variability is imposed by such factors as stream

size, substrate, and geographic locality. The data are further biased by

the selection of riffle areas and by the mesh size used in sample process-

ing. A smaller mesh size will increase the importance of Diptera and

Oligochaeta.

Table 5. Density by group (as percent of total density) for Cane Creek and

other unstressed North Carolina Piedmont streams. Data are the aver-

age of 3-4 collections, rounded to integer value.

ACKNOWLEDGMENTS.— I would particularly like to acknowl-

edge the help and support of two colleagues: Samuel Mozley, N.C.

State University, and David Penrose, N.C. Division of Environmental

Management. Taxonomic assistance was received from many sources:

Chironomidae — Samuel Mozley; Patrick Hudson, U.S. Fish and Wild-

life Service; Leonard Ferrington, University of Pittsburgh; Robert Bode,

Benthic Macroinvertebrates Cane Creek NC 67

N.Y. Department of Health; Broughton Caldwell, Georgia Department

of Natural Resources; N.W. Boesel, Miami University; Don Oliver, Bio-

systematic Research Institute, Canada; and David Smith, U.S. Envir-

onmental Protection Agency, Athens, GA. Ephemeroptera — Paul

Carlson, S.C. Department of Health and Environmental Control. Tri-

choptera — John Morse, Clemson University. Crustacea — John E.

Cooper, N.C. State Museum of Natural History. People assisting in col-

lections included Barbara Burchard, Ross Green, and Ken Eagleson.

Two anonymous reviewers and John E. Cooper provided valuable

comments on the manuscript.

LITERATURE CITED

Bednarik, A. F., and W. P. McCafferty. 1979. Biosystematic revision of the

genus Stenonema (Ephemeroptera: Heptageniidae). Can. Bull. Fish. Aquat.

Sci. 207:1-73.

Coffman, William P. 1973. Energy flow in a woodland stream ecosystem: II.

The taxonomic composition and phenology of the Chironomidae as deter-

mined by the collection of pupal exuviae. Arch. Hydrobiol. 77:281-322.

, K. W. Cummins and J. C. Wuycheck. 1971. Energy flow in a

woodland stream ecosystem: I. Tissue support trophic structure of the

autumnal community. Arch. Hydrobiol. 65:232-276.

Frost, S., A. Huni and W. E. Kershaw. 1971. Evaluation of a kicking technique

for sampling stream bottom fauna. Can. J. Zool. 49:167-173.

Fullner, Richard W. 1971. A comparison of macroinvertebrates collected by

baskets and modified multiple-plate samplers. J. Water Pollut. Control

Fed. 45:494-499.

Holeski, Paul M., and R. C. Graves. 1978. An analysis of shore beetle com-

munities of some channelized streams in northeast Ohio (Coleoptera).

Great Lakes Entomol. 77:23-26.

Lenat, David R., L. A. Smock and D. L. Penrose. 1980. Use of benthic

macroinvertebrates as indicators of environmental quality. Pp. 97-1 12 in D.

L. Worf (ed.). Biological Monitoring for Environmental Effects. Lexington

Books, D. C. Heath Co., Lexington. 227 pp.

Lewis, P. A. 1974. Taxonomy and ecology of Stenonema mayflies (Heptage-

niidae: Ephemeroptera). EPA Monitoring Series EPA-670/ 4-74-006.

Morse, John C, J. W. Chapin, D. D. Herlong and R. S. Harvey. 1980. Aquatic

insects of Upper Three Runs Creek, Savannah River Plant, South Caro-

lina. Part I. Orders other than Diptera. J. Ga. Entomol. Soc. 75:74-101.

Mozley, Samuel C. 1978. Preimpoundment survey of species of benthic

macroinvertebrates in Cane Creek. Unpubl. report Environmental Services

Corp., Chapel Hill. 40 pp.

N. C. Division Environmental Management. 1975. River Basin Plans.

Patrick, Ruth. 1975. Stream communities. Pp. 445-459 in M. S. Cody and J. M.

Diamond (eds.). Ecology and Evolution of Communities. Belknap Press,

Harvard Univ., Cambridge. 545 pp.

Pennak, Robert W. 1978. Freshwater Invertebrates of the United States. 2nd

edition. Ronald Press, New York. 780 pp.

68 David R. Lenat

Penrose, David L., D. R. Lenat and K. W. Eagleson. 1980. Biological evalua-

tion of water quality in North Carolina streams and rivers. N. C. Div.

Environ. Manage., Biological Series #103. 181 pp.

, and . 1982. Aquatic macroinvertebrates of the

Upper French Broad River basin. Brimleyana 8:27-50.

Simmons, Clyde E., and R. C. Heath. 1979. Water quality characteristics of

streams in forested and rural areas of North Carolina. U. S. Geol. Survey

Water Resour. Invest. 79-108. 49 pp.

Smock, L. A., and H. L. Hughes. 1975. Benthic collections. Haw and New

Hope rivers. May 1974-April 1975. Rep. to U.S. Army Corps. Engineers,

Wilmington District, NC. 52 pp.

Vannote, Robert L., and B. W. Sweeney. 1980. Geographic analysis of thermal

equilibria: a conceptual model for evaluating the effect of natural and

modified temperature regimes on aquatic insect communities. Am. Nat.

7/5:667-695.

White, Tina R., and R. C. Fox. 1980. Recolonization of streams by aquatic

insects following channelization. Clemson Univ. Water Resour. Inst. Tech.

Rep. 87, Vol. 1. 119 pp.

Wiggins, G. B. 1977. Larvae of North American caddisfly genera (Trichoptera).

Univ. Toronto Press, Toronto, Canada. 401 pp.

Accepted 30 September 1982

Pleistocene Mammals from the Rock Springs Local Fauna,

Central Florida

Kenneth T. Wilkins^

Florida State Museum,

University of Florida, Gainesville. Florida 3261 1

ABSTRACT. — Although several of the interesting fossil mammals

from Rock Springs, Orange County, Florida, have been mentioned in

scientific papers, this is the first comprehensive report of its mammal-

ian fauna. The fauna, which evidently accumulated during several

intervals of late Pleistocene deposition, consists of at least 27 species,

including both marine and terrestrial forms. The faunal composition,

coupled with modes of preservation (e.g., a barnacle-encrusted speci-

men of a terrestrial species), provides direct evidence of at least one

marine transgression (probably corresponding to the Pamlico shore-

line) into central peninsular Florida. Two extralimital taxa character-

ize the Rock Springs fauna: (1) current affinities of Thomomys sp.

(pocket gophers) are with western North America, and (2) Mormoops

sp. (leaf-chinned bats) now occur in western North America and the

Neotropics.

INTRODUCTION

The Rock Springs site in central Florida has yielded an interesting

sample of late Pleistocene vetebrate fossils. Although the first collec-

tions were made in the 1920s, the first mention of fossil vertebrates from

Rock Springs was a list of seven taxa (Gut 1939). Of the five vertebrate

classes represented in this fauna, only the avifauna has been comprehen-

sively reported (Woolfenden 1959). Auffenberg (1963) included two ser-

pent species — Drymarchon corais, an indigo snake, and Crotalus gigan-

teus, a large rattlesnake — from Rock Springs in his review of the fossil

snakes of Florida. Ray et al. (1963) discussed the presence of Mor-

moops megalophylla, a leaf-chinned bat, and Ray (1964) and Gillette

(1976) studied a species of small cat, Felis amnicola, from this site.

Webb (1974) listed some 17 mammalian species from the Rock Springs

site.

This paper presents the first comprehensive compilation of the

Rock Springs mammalian fauna. Re-examination of all available fossil

material, including a new collection made in 1982, reveals 27 mammal-

ian taxa, some 14 of which are extinct (Table 1). Ten species in the

Rock Springs local fauna presently occur in Florida. Because one spe-

cies was not identifiable to genus (i.e., the large felid), its current status

' Present Address: Department of Biology, Baylor University, Waco, Texas 76798.

Brimleyana No.9:69-82. June 1983. 69

70 Kenneth T. Wilkins

Table 1. Mammalian fauna of the late Pleistocene Rock Springs site, Orange

County, Florida.

is uncertain. The one extant species in the Rock Springs fauna not now

occurring in Florida is Mormoops megalophylla. The other extralimital

genus in the Rock Springs fauna is Thomomys, the smooth-toothed

pocket gophers. General affinities of Mormoops and Thomomys lie in

western North America and the Neotropics.

Rock Springs is located in Kelly Park about 10 km (6 mi.) north of

Apopka, Orange County, Florida (Sorrento quad. NEl/4, NEl/4,

NWl/4, Sec. 15, T20S, R28E). Its elevation is about 7 m above present

sealevel. The spring is one of many flowing into the Wekiva River, a

tributary of the St. Johns River. The spring discharges at a mean rate of

1.83 cubic m/ second (64.6 cubic ft. /second) from a partially-submerged

Pleistocene Mammals Central Florida 71

cavern in a 5 m high limestone bluff of the early Miocene Hawthorne

Formation (Rosenau et al. 1977). The mouth of the cavern is about 1.5

m in diameter. The Rock Springs fauna has been collected along the

initial 200 m of the spring run, and the underlying Crystal River Forma-

tion of the Ocala Group (late Eocene age) is exposed in places along the

run. Present habitats in the surrounding region are the longleaf pine-

turkey oak sandhill ecosystem and the live oak xeric hammock associa-

tion, as is common in the upland karst regions of peninsular Florida.

Mesic hardwoods and other floodplain and freshwater marsh species

occupy the adjacent riparian corridor.

The material comprising the Rock Springs mammal fauna repre-

sents the efforts of many amateurs and professionals, including J.

Bauer, R. and J. Franz, L. F. Lovell, Mrs. C. A. Meyer, G. S. Morgan,

J. W. Pierce, G. M. Ponton, A. E. Pratt, R. Savage, C. Simpson, J. R.

Todd, K. T. Wilkins and G. E. Woolfenden. Additional Rock Springs

fossils are contained in the private collection of H. James Gut, which is

now in the custody of his son, Robert M. Gut of Jacksonville, Florida.

Specimen numbers cited in the following annotated species list refer to

the catalogue of the Florida State Museum Vertebrate Paleontology

Collection at the University of Florida (UF). Catalogue numbers pre-

ceded by "V" are part of the Florida Geological Survey collection which

is also housed with the UF collection.

ANNOTATED SPECIES LIST

Order Insectivora

Family Soricidae

Blarina cf. carolinensis (Bachman) 1837

Material— Partial right mandible with iT, P4-M2, (UF 48997);

partial left mandible with Ml, (UF 48998).

Remarks. — Short-tailed shrews in Pleistocene deposits in Florida

have previously been referred to B. brevicauda, the name formerly app-

lied to most living populations of Blarina in eastern North America.

Recent morphometric and karyotypic study of modern populations

indicates that northern, southeastern and southwestern populations are

separate species, with southeastern populations referred to Blarina caro-

linensis (Genoways and Choate 1972; George et al. 1982). I apply caro-

linensis to the Rock Springs Blarina solely on geographic grounds. Sta-

tistical comparisons of later Pleistocene material from Florida with

modern B. brevicauda, B. hylophaga and B. carolinensis are necessary

to establish species identification.

72 Kenneth T. Wilkins

Order Chiroptera

Family Mormoopidae

Mormoops megalophylla (Peters) 1864

Material. — Cranial and post-cranial material (UF 3860-3866) as

described by Ray et al. (1963).

Remarks. — Identification was established by Ray et al. (1963),

who compared Rock Springs Mormoops with samples of the two living

species of the genus. Kurten and Anderson (1980) noted Rock Springs as

the only Pleistocene occurrence of the species in continental North

America.

Family Vespertilionidae

Myotis austroriparius (Rhoads) 1897

Material. — Skull with partial maxillary dentition (UF 8925); ros-

trum with partial maxillary dentition (UF 8926); 6 dentaries with partial

dentitions and 1 edentulous dentary (UF 8927); 20 humeri (UF 8928); 25

radii (UF 8929); 30 metacarpals (UF 8930); 4 femora (UF 8931).

Remarks. — This material is assigned to M. austroriparius rather

than to M. grisescens Howell 1909 because of its relatively small size.

M. austroriparius is a very common cave-dwelling bat of the karst

regions of present-day Florida (Rice 1957).

Order Edentata

Family Dasypodidae

Holmesina septentrionalis (Leidy) 1889

Material— 2 movable scutes (UF 24845); 4 fixed scutes (UF 24846).

Dasypus bellus (Simpson) 1930

Material. — 1 fixed scute (V-4455).

Family Mylodontidae

cf. Glossotherium (Owen) 1840

Material— Distal phalanx (V-4394).

Remarks. — Poorly ossified epiphyses indicate that this claw is that

of a juvenile. The small size (greatest length = 51 mm) and roundness of

the claw suggest Glossotherium. Measurements taken midway between

proximal and distal ends are: mediolateral width 15 mm; dorsoventral

depth 22 mm. Claws of other ground sloth genera are more mediolater-

ally compressed than in Glossotherium.

Pleistocene Mammals Central Florida 73

Order Lagomorpha

' Family Leporidae

Sylvilagus sp. Gray 1867

Material— Left dentary with P3-P4, M2-M3 (V-4467); proximal

femur (UF 49201); distal femur (UF 49202).

Remarks., — Two species of similar-sized cottontails, Sylvilagus

palustris and S.floridanus, presently range throughout most of Florida.

The distribution of a third, much larger species, S. aquaticus, closely

approaches the Florida panhandle in Alabama; quite possibly the spe-

cies has recently extended its range into extreme western Florida. Left

mandibular toothrow lengths were measured in five specimens each

(mixed sex) of modern S. palustris and S. floridanus from Alachua

County, Florida, in efforts to assign specific identity to the Rock

Springs rabbit material. The sample mean for toothrow length for S.

floridanus is 15.20 mm (s = 0.29 mm) and for S. palustris is 15.24 mm (s

= 0.45 mm). Toothrow length in the Rock Springs specimen is 14.9 mm.

Lack of mensural differences between these three samples dictates refer-

ral of the Rock Springs rabbit to Sylvilagus sp.

Order Rodentia

Family Geomyidae

Geomys pinetis Rafinesque 1817

Material— Isolated upper P4, (UF 49205).

Remarks. — Geomys pinetis is the only extant pocket gopher in the

eastern United States. The southeastern pocket gopher is presently

abundant in the vicinity of Rock Springs, and in other parts of the

southeastern United States characterized by the longleaf pine-turkey

oak sandhill ecosystem. Despite the limited referred material, its generic

identity is certain. Extant Geomys species possess an enamel band on

the posterior surface of the fourth upper premolar. This posterior

enamel investment on upper P4 is absent in Thomomys species and in

the early-to-middle Irvingtonian species of Geomys in Florida (Wilkins

1984).

Thomomys cf. orientalis Simpson 1928

Material — Six mandibles with partial to complete dentitions (UF

46571-46576).

Remarks. — Generic assignment of these specimens is indicated by

the presence of both anterior and posterior enamel bands on the lower

molars; anterior bands are absent in Geomys species. The genus Thom-

omys presently occurs in western North America (Hall 1981). The only

74 Kenneth T. Wilkins

other Thomomys material reported from the southeastern United States

is from Sabertooth Cave (Rancholabrean, Citrus Co., Florida). Simp-

son's (1928) description of the species referred only to skull material,

none of which is available in the Rock Springs deposit. Pending further

study, the Rock Springs Thomomys is tentatively referred to orientalis.

The Rock Springs site has yielded additional geomyid material for

which generic assignment cannot be made on qualitative grounds. Sim-

ilarity in size and preservation to the Thomomys material Hsted above

suggests that four mandibles lacking cheekteeth (UF 46577-46580) are

referable to Thomomys. No effort has been made to identify isolated

lower P4's (UF 46581) and molars (UF 46582) beyond the family level.

Family Castoridae

Castor canadensis ¥.\x\i\ \%20

Material. — Lower molar (V-4399).

Remarks. — The presence of beaver in the Rock Springs fauna was

previously noted by Johns (1958), who argued that its present distribu-

tion (restricted in Florida to the panhandle and northern peninsula) is

less extensive than it was in historic and Pleistocene times. The Rock

Springs specimen, coupled with two nearly complete mandibles with P4-

M3 (V-5403) from the shores of nearby Lake Monroe, Volusia County,

comprises the species' southernmost Pleistocene record in Florida. Post-

Wisconsinan records include specimens from middens in Seminole,

Volusia and Brevard counties (Furgeson 1951) and along the Indian

River (Allen 1942).

Family Cricetidae

cf. 5/gmo<io« Say and Ord 1825

Material. — Two isolated upper incisors (UF 48999).

Remarks. — Comparison of these incisors with modern Sigmodon

hispidus and other rodents from Florida suggests this generic assign-

ment. Martin (1974) recognized the occurrence of two species of cotton

rats, S. bakeri and S. hispidus, in the United States and Mexico during

the late Irvingtonian and Rancholabrean. Identification to species

requires molar teeth.

Pleistocene Mammals Central Florida 75

Order Cetacea

Family Delphinidae

cf. Tursiops

Material— Vertebra (UF 48976).

Remarks. — The specimen is a complete centrum (diameter = ca. 38

mm) with one complete transverse process (length = ca. 75 mm). The

paired portion of the neural arch ventral to its fusion into the spine is

present. Comparison with modern small odontocetes from Florida sug-

gests Tursiops. The relatively intact nature of this specimen, and the

lack of water-wear, suggests in situ deposition and comprises further

evidence of marine transgression above the Rock Springs elevation.

Order Carnivora

Family Canidae

Urocyon cinereoargenteus (Schreber) 1775

Material.— Distal humerus (UF 49000).

Remarks. — This specimen resembles humeri of modern individuals

of the gray fox from central Florida.

cf. Canis dirus Leidy 1858

Material. — Canine fragment (V-4397).

Remarks. — This identification follows Webb (1974).

Family Ursidae

Tremarctos floridanus (Gidley) 1928

Material— Molar fragment (UF 8946); 2 upper M3's (UF 8947-

8948).

Remarks. — The genus is extinct in North America, with the sole

living species restricted to the South American Andes. Tremarctos flori-

danus, the Florida spectacled bear, was a common member of many of

Florida's Pleistocene faunas; Ursus and Tremarctos remains are often

found in the same deposits. Ecological differences of the two species

apparently allowed coexistence (Kurten 1966). Adaptation of T. florida-

nus to a highly herbivorous lifestyle is evident from its relatively broad

molars, which allowed for increased occlusal surface areas. The rela-

tively narrow and elongate molars of U. americanus indicate more

omnivorous adaptation.

76 Kenneth T. Wilkins

Ursus americanus

Material— Two upper M3's (UF 8951-8952); femur (UF 8953).

Remarks. — The black bear presently occurs throughout much of

Florida. An unfossilized femur attests to its relatively recent occurrence

in the Rock Springs vicinity. Generic identity of other ursid material is

uncertain: distal humerus (UF 49203); canine fragment (UF 49204).

Family Felidae

Felis amnicola Gillette 1976

Material— Right dentary with P3-MT (UF 4522)

Remarks. — This specimen was originally recognized and illustrated

by Ray (1964) as a jaguarundi, Felis yagouaroundi. Subsequently,

Gillette (1976) described a new species of small river cat, F. amnicola, to

which he referred this Rock Springs dentary.

cf. Felidae

gen. et sp. indet.

Material— Phalanx (UF 8954)

Remarks. — Webb's (1974) tabulation of "Felis sp." at Rock Springs

probably refers to this specimen, which compares favorably in size with

the modern Florida panther, Felis concolor coryi Bangs 1899.

Order Proboscidea

Family Mammutidae

Mammut americanum (Kerr) 1791

Material. — Right dentary fragment (V-4378); molar fragments (UF

48986, V-4385, V-4464); molar enamel fragment (V-4465).

Family Elephantidae

Mammuthus sp. (Falconer) 1857

Material— Molar (V-4473); molar plates (UF 48987, V-4383); tusk

fragments (V-4384).

Remarks. — In accordance with Kurte'n and Anderson (1980),

mammoth material is provisionally referred to Mammuthus sp. pending

comprehensive study of the genus.

Order Sirenia

Trichechus manatus Linnaeus 1758

Material— Cheektooth (UF 48981); cheektooth fragment (UF

48982); edentulous maxillary fragment (V-4451); ear ossicle (UF 48983);

Pleistocene Mammals Central Florida 77

tympanic (UF 48984); vertebrae material (UF 48979-48980, V-4469);

ribs (UF 48977-48978, V-4380).

Remarks. — Various states of preservation are evident in this mana-

tee material. Some is well permineralized, whereas other specimens are

only slightly fossilized. The material was probably deposited during sev-

eral different intervals, including times when the site was marine or

estuarine, or when it comprised a freshwater system occupied by mana-

tees much as are many other river systems in Florida today.

Order Perissodactyla

Family Tapiridae

Tapirus veroensis Sellards 1918

Material— Palate with left U, 13, PJ_, M3 and right PJ_-M3 (UF

18702); left maxilla with MNM3 (V-4389);_upper molar fragment (UF

48970); left dentary and_symphysis with P4-M3 (V-4390/4391); partial

right dentary with_P4-M3 (UF 12485); left 12 (UF 48971); deciduous P2

(UF 8945); left P2 (V-4396); lower molar (UF 48969); 5 isolated teeth

(V-4454).

Remarks. — All referred material resembles T. veroensis rather

than the larger T. copei Simpson 1945.

Family Equidae

Equus sp.

Material— Upper cheektooth (UF 48975); lower molar (UF 48974);

incisors (UF 48972-48973); radius (V-4471).

Remarks. — The material represents several states of preservation.

The unfossilized radius, that of an immature individual, probably

represents the domestic Equus caballus. The dental material is fossilized

and is provisionally referred to Equus sp.

Order Artiodactyla

Family Tayassuidae

Mylohyus nasutus (Leidy) 1869

Material— Partial right mandible with P3-M3(UF 17720).

Remarks. — This specimen is particularly interesting because of its

preservation. Overall coloration of the specimen is black, in contrast to

buff or chalky colors of most other Rock Springs material. The ventral

surface of the dentary is broken away to expose the mandibular canal.

External surfaces of the jaw, as well as the internal surfaces of the man-

dibular canal, are encrusted with growths of various marine organisms,

including barnacles. Assuming in situ deposition, this specimen further

78 Kenneth T. Wilkins

documents marine transgression of the Rock Springs site. The most

recent occurrence of a transgression sufficient to cover the Rock Springs

site probably corresponds to the Pamlico marine terrace (Healy 1975).

Hence, the minimum age estimate for this specimen is about 125,000

years (see Discussion).

Family Camelidae

Paleolama mirifica (Simpson) 1929

Material— Deciduous P4 (V-4398); P4 (V-4453); M3 (UF 12491).

Remarks. — Webb (1974:183) recorded this stout-legged llama as a

member of the Rock Springs fauna in his study of Florida Pleistocene

Lamini.

Family Cervidae

Odocoileus virginianus (Zimmermann) 1780

Material — Right dentary with P3-M3 (V-4395); isolated cheek-

teeth (UF 48988-48989, V-4462); distal tibia (UF 48990); distal metatar-

sal (UF 48991).

Remarks. — The white-tailed deer presently occurs throughout

Florida. Because of differential preservation, deer material in the Rock

Springs fauna appears to have been deposited during several intervals.

Family Bovidae

Bison sp.

Material— Upper cheektooth (UF 10037); M3 (UF 52184); lower

cheektooth (UF 15065).

Remarks. — Robertson (1974) stated that unequivocal species

determination for the three species of Bison in Florida can be made only

from horn core material. He was unable to identify diagnostic dental

characters, although he noted that, by its smaller size. Bison bison (Lin-

naeus) 1758 could sometimes be distinguished from the two larger spe-

cies, Bison latifrons (Harlan) 1825 and Bison antiquus Leidy 1852.

Mesurements of the nearly unworn lower M3(UF 52184) are: greatest

alveolar length = 47.7 mm, greatest alveolar width =21.2 mm, and

greatest crown height = 69. 1 mm. The large size of the Rock Springs M3

suggests the presence of one of the larger Bison species rather than B.

bison.

DISCUSSION

Chronology

As with many late Pleistocene fluvial sites in Florida, the Rock

Springs local fauna accumulated during several intervals. Some material

Pleistocene Mammals Central Florida 79

is Recent or even unfossilized (e.g., some of the Equus and Odocoileus).

In addition, some Miocene fossils of marine invertebrates and sharks

have eroded from the limestone bedrock through which the stream

flows. The late Pleistocene fossilized materials represent accumulation

in two types of situations: one of higher sealevels and another of much

lower sealevels. The cetacean vertebra, the barnacle-covered Mylohyus

dentary, and the Trichechus material comprise evidence of marine intru-

sion. The last time seas could have covered the Rock Springs locale

(elevation 7 m above present sealevel) was some 125,000 years before

present (ybp). According to eustatic studies, 125,000 ybp was the most

recent occasion that seas have been higher than at present (Shackleton

and Opdyke 1973; Bloom et al. 1974; Chappell 1974). In Florida, this

late Sangamonian interglacial transgression corresponds to the Pamlico

shoreline at about 9 m (Healy 1975). It is possible, yet unHkely, that

some earlier transgressions of similar or greater extent introduced the

marine influence. Thus, the cetacean, the barnacles encrusting the

previously-deposited peccary jaw, and at least some of the sirenian

material are probably of Sangamonian age.

Most of the terrestrial vertebrates presumably were deposited dur-

ing one or more phases of reduced sealevel. During at least one such

interval, sealevel and the piezometric surface dropped sufficiently to

produce habitats suitable to vertebrates that do not now occur in east-

ern North America. Woolfenden (1959) noted a western contingent in

the Rock Springs avifauna. A portion of the Rock Springs mammal

fauna also exhibits noneastern United States affinities. Smooth-toothed

pocket gophers (genus Thomomys) are known from the Rock Springs

deposit, but the genus now occurs only in Mexico and western North

America (Hall 1981). The range of Mormoops megalophylla is now res-

tricted to the southwestern United States, southward through much of

Central America and into northern South America (Smith 1972). Mor-

moops blainvilli, the only other extant species of the genus, occurs in

the West Indies (Hall 1981).

Paleoenvironmental Interpretations

The genus Mormoops (as well as the species M. megalophylla

itself) is neotropical and temperate in distribution and occupies humid

to semiarid to arid situations at elevations generally less than 3000 m

(Smith 1972). Similarly, Thomomys pocket gophers occupy habitats

including deserts, prairies, montane meadows and forests ranging over

some 35 degrees of latitude and 3000 m of elevation. It is apparent that

no single temperature or rainfall regime characterizes the entire range of

either genus. Therefore, because of their broad habitat and environmen-

tal tolerances, the presence of M. megalophylla or Thomomys at Rock

Springs need not be directly indicative of any particular habitat types.

80 Kenneth T. Wilkins

Yet, their presence does offer clues in reconstructing certain aspects of

the paleoenvironment.

Bats. — The two species of bats recorded as fossils at Rock Springs

are usually considered obligate cave-dwellers, although alternate roosts

are also used (Barbour and Davis 1969). Their presence suggests a

reduced piezometric surface. No modern records of M. austroriparius

are known from the Rock Springs cavern, although the species is

reported from various northwestern Orange County caverns in the

immediate vicinity (Rice 1957). The present water level in the Rock

Springs cavern is probably too high to accomodate sizable colonies of

bats. Additionally, bat bones are quite delicate; that most of the fossil-

ized bat material is intact suggests the absence of flowing water in the

cavern during the depositional intervals. Yet, some water was probably

on the cave floor during occupation by Myotis austroriparius. This is

perhaps the most abundant extant bat species in Florida and it resides

in caves with either still or flowing water beneath roosting sites. Avian

remains from Rock Springs also showed Httle evidence of water-wear

(Woolfenden 1959). As with many other stream deposits, the context

and associations of fossil specimens is unknown and cannot be recon-

structed. However, the similarity of preservation of both Mormoops

and Myotis fossils indirectly suggests that occupation of the cave could

have been contemporaneous.

Pocket gophers. — The contemporaneous occurrence of two con-

trageneric pocket gopher species in a fossil deposit was previously

reported by Dalquest and Kilpatrick (1973). They interpreted their cave

deposit (Shulze Cave, early Holocene, Edwards County, Texas) as the

roost of barn owls that foraged over a broad area, including the differ-

ent habitats occupied by Thomomys bottae and Geomys bursarius,

which had mutually exclusive microgeographic distributions. It is quite

possible that the Rock Springs cavern served as an owl roost during

times of reduced water table levels. Fossil barred owls, Strix varia, are

known from Rock Springs (Woolfenden 1959). Lack of stratigraphic

context disallows determination of whether Geomys and Thomomys

occurred contemporaneously in the Rock Springs vicinity rather than

being members of faunas of different time intervals. Nevertheless, the

mere presence in the fauna of the two species of pocket gophers allows

inferences regarding soil characteristics. Miller (1964) examined soil

preferences and competitive interactions of three genera of pocket

gophers in Colorado. He found that Geomys species required deeper,

sandier and more friable soils than either Pappogeomys species (includ-

ing Cratogeomys species) or Thomomys species. Thomomys, in con-

trast, was well suited to shallow, gravelly, less friable soils, although

Thomomys could and would inhabit deeper soils where available.

Requirements of Pappogeomys were intermediate to the other genera.

Pleistocene Mammals Central Florida 81

Miller found in competition experiments that, in its preferred deeper

soils, Geomys excluded species of other genera, whereas Thomomys was

the superior competitor in shallow soils. Hence, presence of Geomys

and Thomomys in the Rock Springs deposit suggests local occurrence

of two markedly different substrates. The deep, sandy soils predominat-

ing the vicinity today favor Geomys and generally represent sediments

of marine terraces of interglacial periods. Shallower, gravelly soils often

occupied by Thomomys in western North America are uncommon in

Florida today. However, such soils could be developed via erosion of

deep sands overlying Umestone. Shallow sands mixed with gravel formed

of eroding limestone could form a substrate inhabitable by Thomomys

but not by Geomys.

ACKNOWLEDGMENTS.— I thank Richard Franz, Jeff Franz,

Gary Morgan and Ann Pratt for field assistance. Discussions with

Richard Franz, Richard Hulbert, Gary Morgan and David Webb aided

me in identifying specimens and in making paleoecological interpreta-

tions. The efforts of Mary Ellen Ahearn, Jon Becker, Richard Hulbert,

Bruce MacFadden, Gary Morgan, Ann Pratt and David Webb in

reviewing drafts of this manuscript are greatly appreciated. This paper is

University of Florida Contribution to Vertebrate Paleontology No. 223.

LITERATURE CITED

Allen, Glover M. 1942. Extinct and vanishing mammals of the western hemis-

phere with the marine species of all the oceans. Intelligence Printing Co.,

Lancaster. 620 pp.

Auffenberg, Walter. 1963. The fossil snakes of Florida. Tulane Stud. Zool.

70(3):131-216.

Barbour, Roger W., and W. H. Davis. 1969. Bats of America. Univ. Press

Kentucky, Lexington. 286 pp.

Bloom, Arthur L., W. S. Broecker, J. M. A. Chappell, R. K. Matthews and K.

J. Mesolella. 1974. Quaternary sea level fluctuations on a tectonic coast:

230 Th/234 U dates from the Huon Peninsula, New Guinea. Quat. Res.

(N.Y.) ¥.185-205.

Chappell, John. 1974. Geology of coral terraces, Huon Peninsula, New Guinea:

a study of Quaternary tectonic movement and sea-level changes. Bull. Geol.

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Dalquest, Walter W., and C. W. Ki;patrick. 1973. Dynamics of pocket gopher

distribution on the Edwards Plateau of Texas. Southwest. Nat. 75(1): 1-9.

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Genoways, Hugh H., and J. R. Choate. 1972. A multivariate analysis of syste-

matic relationships among populations of the short-tailed shrew (genus

Blarina) in Nebraska. Syst. Zool. 27.- 106-1 16.

George, Sarah B., H. H., Genoways, J. R. Choate and R. J. Baker. 1982. Kary-

otypic relationships within the short-tailed shrews, genus Blarina. J. Mam-

mal. 65(4):639-645.

82 Kenneth T. Wilkins

Gillette, David D. 1976. A new species of small cat from the late Quaternary of

southeastern United States. J. Mammal. 57(4):664-676.

Gut, H. James. 1939. Hitherto unrecorded vertebrate fossil localities in south-

central Florida. Q. J. Fla. Acad. Sci. 5:50-53.

Hall, E. Raymond. 1981. The Mammals of North America. John Wiley and

Sons, New York, 1 181 + 90 pp., 2 volumes.

Healy, Henry G. 1975. Terraces and shorelines of Florida. Fla. Dep. Nat. Re-

sour., Bur. Geol. Map Series No. 71.

Johns, Bette A. S. 1958. The distribution of the beaver in Florida and a study of

its ecololgy in the Flint-Chattahoochee-Apalachicola region. Unpubl. M.S.

thesis, Univ. Florida, Gainesville. 48 pp.

Kurt^n, Bjorn. 1966. Pleistocene bears of North America. I: Genus Tremarctos,

spectacled bears. Acta Zool. Fenn. 775:1-120.

, and E. Anderson. 1980. Pleistocene Mammals of North America.

Columbia Univ. Press, New York. 442 pp.

Martin, Robert A. 1974. Fossil mammals from the Coleman HA fauna, Sumter

County. Pp. 35-99 in S. D. Webb (ed.). Pleistocene Mammals of Florida.

Univ. Presses Fla., Gainesville. 270 pp.

Miller, R. S. 1964. Ecology and distribution of pocket gophers (Geomyidae) in

Colorado. Ecology 45:156-212.

Ray, Clayton E. 1964. The jaguarundi in the Quaternary of Florida. J. Mam-

mal. ¥5(2):330-332.

, S. J. Olsen and H. J. Gut. 1963. Three mammals new to the Pleis-

tocene fauna of Florida, and a reconsideration of five earlier records. J.

Mammal. ¥¥(3):373-395.

Rice, Dale W. 1957. Life history and ecology of Myotis austroriparius in Flor-

ida. J. Mammal. i5(l): 15-32.

Robertson, Jesse S., Jr. 1974. Fossil Bison of Florida. Pp. 214-246 in S. D.

Webb (ed.). Pleistocene Mammals of Florida. Univ. Presses Fla., Gaines-

ville. 270 pp.

Rosenau, Jack C, G. L. Faulkner, C. W. Hendry, Jr. and R. W. Hull. 1977.

Springs of Florida. Bull. Fla. Bur. Geol. 57.1-461.

Shackleton, Nicholas J., and N. D. Opdyke. 1973. Oxygen isotope and palaeo-

magnetic stratigraphy of equatorial Pacific core V28-238: Oxygen isotope

temperatures and ice volumes on a 10^ year and 10^ year scale. Quat. Res.

(N.Y.)5(l):39-55.

Simpson, George G. 1928. Pleistocene mammals from a cave in Citrus County,

Florida. Am. Mus. Novit. 525:1-16.

Smith, James D. 1972. Systematics of the chiropteran family Mormoopidae.

Univ. Kans. Mus. Nat. Misc. Publ. 56.1-132.

Webb, S. David (ed.). 1974. Pleistocene Mammals of Florida. Univ. Presses

Fla., Gainesville. 270 pp.

Wilkins, Kenneth T. 1984. Evolutionary trends in Florida Pleistocene pocket

gophers (genus Geomys), with description of a new species. J. Vertebr.

Paleontol. 5(3):166-181.

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Wilson Bull. 77(2): 183- 187.

Accepted 8 August 1983

Bird Density and Habitat Use in Forest Openings Created

by Herbicides and Clearcutting in

The Central Appalachians

William C. McComb and Robert L. Rumsey ^

Department of Forestry, University of Kentucky

Lexington, Kentucky 40546-0073

ABSTRACT. — Winter and breeding bird communities were sampled

on clearcuts, on plots receiving four rates of picloram herbicide appli-

cation, and on untreated plots. Four-year-old picloram treated plots

did not increase bird species diversity, but did increase bird density

over untreated plots. Positive response of some bird species to treat-

ment was due to creation of an edge. Bird density was affected by

physiographic site. Hairy woodpeckers, Picoides villosus; wintering

red-bellied woodpeckers, Melanerpes carolinus; and Carolina chicka-

dees, Parus carolinensis, used south-facing slopes more than north-

facing slopes, while red-eyed vireos, Vireo olivaceus, and ovenbirds,

Seiurus aurocapillus, used north-facing slopes more than ridgetop

sites. White-breasted nuthatches, Sitta carolinensis', red-bellied wood-

peckers; tufted titmice, Parus bicolor; and Carolina chickadees exhi-

bited changes in habitat use from winter to spring. A variety of piclo-

ram herbicide treatments (27-68 kg/ ha) and clearcutting in small (0.5-

1.0 ha) blocks, on both south-facing and north-facing slopes, is

recommended to increase bird density and provide more diverse habi-

tat than exists in undisturbed forests.

INTRODUCTION

Herbicides are used by foresters for timber stand improvement and

by wildlife managers to create forest openings (McCaffery et al. 1974;

Loftis 1978; Dewey 1980). Any herbicide use or timber harvesting

affects the structure and composition of forests as wildlife habitat. The

effects of forest cutting on bird communities have been reported by

McComb and Noble (1980), Strelke and Dickson (1980), and Crawford

et al. (1981), who found that tree thinning tended to increase bird spe-

cies diversity (BSD) and bird abundance, probably due to an increase in

foliage height diversity of the stand. In contrast, little information is

available on the effects of herbicide application on forest bird communi-

ties. Beaver (1976), Savidge (1978), and Best (1972) reported the effects

of 2,4,5-T and 2,4-D applications to shrubs and trees on birds and

found either no effects or reductions in bird density and /or diversity.

Shipman (1972) reported greater wildlife diversity for up to 10 years

after application of fenuron herbicide on 0.04-ha plots in Pennsylvania.

Present address: Department of Agriculture, McNeese State University,

Lake Charles, Louisiana 70609

Brimleyana No.9:83-95. June 1983. 83

84 William C. McComb and Robert L. Rumsey

Picloram-based herbicides are desirable for creating forest openings

because of their low toxicity to many vertebrates (Kenaga 1969), but we

could find no studies that compared bird use of picloram-created clear-

ings with use of clearcut or uncut areas. Our objectives were to: 1) com-

pare the relative abundance and diversity of winter birds and breeding

birds among 4-year-old herbicide-created forest clearings, 4-year-old

clearcuts, and mature untreated areas, 2) evaluate the effect of edge

creation on bird use of habitats, and 3) identify the characteristics of

habitats used by common species.

STUDY AREA AND METHODS

Snag Ridge Fork watershed, in the University of Kentucky's

Robinson Forest, Knott and Breathitt counties, Kentucky, contains a

second-growth mixed-mesophytic forest typical of much of the central

Appalachians (Carpenter and Rumsey 1976). Ridges are dominated by

shortleaf pine, Pinus echinata\ pitch pine, P. rigida; chestnut oak, Quer-

cus prinus; and scarlet oak, Q. coccinea. South-facing slopes are domi-

nated by hickories, Carya spp.; white oak, Q. alba; black oak, Q. velu-

tina; and sourwood, Oxydendrum arboreum. North-facing slopes are

dominated by northern red oak, Q. rubra; cucumbertree. Magnolia

acuminata; and yellow-poplar, Liriodendron tulipifera.

Fifteen of 18 0.4-ha square plots were treated in the watershed.

This included four plots on each of a north-facing slope, south-facing

slope, and ridge-top, which were randomly assigned one of the follow-

ing hand-broadcast herbicide treatments applied in May 1976: 23 kg/ ha

TORDON 10k; 45 kg/ha TORDON lOK; 68 kg/ha TORDON lOK; or

90 kg/ ha M-3864. TORDON lOK is a pelletized picloram-based (10%

4-amino-3,5,6-trichloropicolinic acid) herbicide and M-3864 is a 5% pic-

loram pellet. (Mention of trade names is for identification and does not

imply endorsement by the Kentucky Agricultural Experiment Station,

Lexington.) A fifth plot on each aspect was clearcut (all stems cut, no

residuals); fallen trees were not removed. A sixth plot on each aspect

was established as a control in the untreated forest at least 75 m from

any treated plot. Plots were located along the contour, and treated plots

were 15 m to 50 m apart.

Fifteen stations were established perpendicular to the contour

through the center of each plot. Thirty environmental characteristics

(Table 1) chosen based on Anderson and Shugart (1974), Stauffer and

Best (1980), and Crawford et al. (1981) were measured at each station.

Estimates of cover of rocks, logs, leaves, canopy, and midstory followed

methods described by James and Shugart (1971). Environmental char-

acteristics other than understory vegetation were measured in May

1980. Understory vegetation was quantified on one 4-m^ circular plot

(radius = 1.2 m), 2 m away from each station along the contour in

Bird Response to Clearings 85

Table 1. Habitat variables used in correlation analyses, bird habitat preference

on clearcut, herbicide-treated and uncut plots, January to June 1980

and 1981, Knott County, Kentucky.

Description

Number of trees>10 cm dbh, within 2 m of a station.

Diameter of nearest tree (cm).

Distance to nearest tree (m).

Basal area of living stems (m^/ha), wedge prism.

Number of snags> 10 cm dbh, > 1 .8 m tall, within 2 m of a station.

Diameter of nearest snag (cm).

Distance to the nearest snag (m).

Number of stumps>10 cm in diameter and<1.8 m tall, within 2 m of a station.

Distance to the nearest stump (m).

Number of logs>10 cm diameter,>1.8 m long, within 2 m of a station.

Maximum diameter of the nearest log (cm).

Length of the nearest log (m).

Distance to the nearest log (m).

Percent of ground covered by logs within 2 m of a station.

Distance of the nearest rock>5 cm above ground.

Percent of ground covered by rocks within 2 m of a station.

Percent crown cover above 6.1 m at a station.

Percent vegetation cover between 1.8 and 6.1 m at a station.

Percent of the ground covered by fallen leaves within 2 m of a station.

Percent understory cover<1.8 m tall per 4-m^ in Jan. and in Apr.

Number of understory stems per 4-m^ in Jan. and in Apr.

Number of understory taxa per 4-m^ in Jan. and in Apr.

Understory species diversity per 4-m^ in Jan. and in Apr.

Distance to water (m).

Slope (%).

Foliage height diversity.

January and April of 1980. A modified Aldous method, similar to that

described by Murphy and Noble (1972), was used to quantify vegeta-

tion. Percent cover and stem density were determined for each plant

taxon (at least to genus) on each 4-m^ plot. Total cover, total stem

density, plant species richness, and a Shannon-Weaver plant species

diversity index were calculated for each station for both sampling peri-

ods. April understory values were used in conjunction with May mid-

story and overstory values to calculate a foliage height diversity index

based on estimated cover of the three layers. Mean habitat characteris-

tics were calculated for each 0.4-ha plot and used to characterize the

plot. A more detailed discussion of habitat characteristics was given by

McComb and Rumsey (in press).

86 William C. McComb and Robert L. Rumsey

Birds were counted 15 times on each 0.4-ha plot from 15 January

to 5 March 1980 and 1981 (winter birds), and 13 times on each plot

from 20 March to 15 June 1980 and 1981 (breeding birds). Approxi-

mately 10 minutes were spent on the center of each plot during each

visit counting birds seen or heard on the plot. Locations of birds in the

plot were judged to be within 9 m of the plot edge or within the plot

center. Only winter birds observed foraging were assumed using a plot

while breeding birds which were either singing or foraging were tallied.

Birds were counted within three hours of sunrise or sunset. Eight morn-

ing and seven evening visits were made to each plot during winter, and

seven morning and six evening visits were made to each plot during the

breeding season. Shannon-Weaver BSD, equitability, richness, and

average density per plot per visit of each species were compared among

treatments blocking on aspects with analysis of variance and Duncan's

New Multiple Range Test. Tests among treatments are conservative due

to variability among aspects. Linear correlation was used to identify

plot characteristics potentially important in determining use by common

bird species. A t-test was used to compare weighted mean environmental

characteristics for plots used by a species vs. overall means and to com-

pare mean BSD, equitabiHty, richness, and density between seasons.

Chi-square analysis was used to compare occurrences of birds at the

edge vs. center of plots.

RESULTS AND DISCUSSION

Seasonal Effects. — Not unexpectedly, the breeding bird communi-

ties had higher diversity (x = 0.91 1), richness (x = 2.2), equitability (x =

0.238), and density (x = 2.4 birds/ 0.4 ha) than the winter bird commun-

ities (x = 0.265, 1.0, 0.069, and 1.0, respectively) (Table 2). Winter bird

communities were dominated by hairy woodpeckers, Picoides villosus,

and Carolina chickadees, Parus carolinensis (Table 2), and the breeding

bird communities were dominated by red-eyed vireos, Vireo olivaceus,

and hooded warblers, Wilsonia citrina (Table 3). Several resident spe-

cies changed their habitat usage from winter to spring. White-breasted

nuthatches, Sitta carolinensis, used areas in spring with a higher den-

sity (x = 1/43 m^) of larger diameter (x = 1.23 cm) snags and fewer trees

(x = 1/32 m^) than in winter (x = 1/171 m^, 7.9 cm and 1/27 m^ respec-

tively) (P<0.05). Red-bellied woodpeckers, Melanerpes carolinus, used

areas in spring with more snags (x = 1/50 m^), higher understory density

(x = 45.2 stems/ 4 m^), and lower foliage height diversity (x = 1.23) than

in winter (x = 1/171 m^ 34.7 stems/4 m^, and 1.40, respectively)

(P<0.05). Both white-breasted nuthatches and red-bellied woodpeckers

probably used areas with higher snag density in spring for nesting pur-

poses, since both will forage on dead or living trees in the winter

(Conner 1979).

Bird Response to Clearings

87

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Carolina chickadees used areas with greater (P<0.05) understory

species richness in the spring (x = 7.3 species/ 4 m^) than in the winter (x

= 6.6), and tufted titmice used areas with lower (P<0.05) leaf cover (a

more open canopy) (x = 46.4%) in the spring than in the winter (x =

63.5%).

Winter Birds. — Bird species diversity, equitability, and richness did

not vary significantly among treatments nor among aspects (P>0.05).

Winter bird density was higher on clearcuts, 23 kg/ ha, and 45 kg/ ha

plots than on the control plots (P<0.05) (Table 2), and bird density on

ridge-tops (x = 1.1 bird/ plot/ visit) (P<0.05) was higher than on north

facing slopes (x = 0.53 birds/ plot/ visit) (P<0.05). South-facing and rid-

getop areas are warmer than north-facing slopes throughout the day-

light hours and probably influenced bird activity (Shields and Grubb

1974).

Hairy woodpecker abundance was similar among treatments, but

they used south-facing slopes (17 individuals) more than north-facing

slopes (7 individuals), and they were observed using plot edges (20 indi-

viduals) more than plot centers (4 individuals) (P<0.05). Carolina

chickadee abundance was similar among treatments, but they used

ridge-tops (25 individuals) more than north-facing slopes (5 individuals)

(P<0.05). Strelke and Dickson (1980) found breeding Carolina chicka-

dees associated with pine-hardwood clearcut edges in Texas. Carolina

chickadee numbers were correlated with distance to the nearest stump

(r =+ 50.6), slope (r =- 50.5), and understory density (r =+ 49.4) (P< 0.05).

Distance to stumps on plots used by chickadees (23.2 m; SD = 7.1)

differed significantly (P<0.05) from overall stump distance (17.4 m; SD

= 7.9). Large, soft stumps may afford roosting sites for chickadees.

White-breasted nuthatch abundance was similar among aspects

(P>0.05), but more individuals were observed on 45 kg/ ha plots than

on any other treatment except 23 kg/ ha plots (P<0.05) (Table 2).

White-breasted nuthatch numbers were correlated with log length (r =

+ 51.1). Log presence indicates prior snag presence so nuthatches may be

using an area that recently had snags present. Plots used by wintering

white-breasted nuthatches had higher crown cover (x = 57.5%), lower

understory diversity (x = 4.73), and lower understory density (x = 26.3

stems/ 4 m^) than overall plot means (x = 31.6, 5.15, and 36.2, respec-

tively). Wintering white-breasted nuthatches occurred on plots with

open crowns, sparse understory, and many long logs on the ground.

Red-bellied woodpeckers used ridge-tops (13 individuals) more

than south- or north-facing slopes (2 individuals each) and they were

found more frequently at plot edges (12 individuals) than centers (5

individuals) (P<0.05). Red-bellied woodpecker numbers were negatively

correlated with slope (r = 45.7), a result of their use of ridge sites.

Bird Response to Clearings 91

Wintering northern cardinals, Cardinalis cardinalis, used clearcuts

more than other treatments except 23 kg/ ha plots (P<0.05) (Table 2).

Both treatments resulted in dense understory vegetation (>100%) and

low basal area (<17.5 m^ha). Cardinals showed no association with

edge. Northern cardinal abundance was correlated with snag and tree

distance (r =+ 62.4, + 57.7), and basal area (r =- 54.1). Snags provide

singing perches for cardinals.

Rufous-sided towhees, Pipilo erythrophthalmus, used clearcuts

more than any other treatment (P<0.05) (Table 2). Clearcuts had the

lowest basal area (9.3 m^/ha 4-year-old regrowth) and the highest

ground cover (logs, 3.5/ 12 m^; rocks, 6.7% cover; stumps, 0.8/ 12 m^) of

any of the plots. Towhee numbers were correlated with distance to the

nearest tree (r =+ 80.0) and basal area (r =- 70.3) (P<0.01). Trees near

young clearcuts probably provide singing perches for towhees. These

results concur with those of Crawford et al. (1981) and Conner and

Adkisson(1975).

Sample sizes for 13 other winter resident species were too small to

detect meaningful differences among treatments or aspects (Table 2).

Breeding Birds. — We found no differences in BSD or equitability

among treatments or aspects (P>0.05). Average breeding bird density

was higher on 23 kg/ ha plots than on other treatments except 45 and 68

kg/ ha plots (P<0.05) (Table 3). Density was higher on south- and

north-facing slopes (x = 1.6 birds/ plot/ visit) than on ridge-tops (x = 1.2

birds/ plot/ visit) (P<0.05). Smith (1977) reported that moist forests

were of more importance to most species of breeding birds in Arkansas

than extremely dry sites.

The red-eyed vireo was the most abundant species and it used

south- and north-facing slopes (31 and 26 individuals) more than ridge-

tops (13 individuals), and edges (54 individuals) more than centers (16

individuals) (P<0.05). Because edges of most plots had more complete

canopy development than plot centers (except control plots), it is not

surprising that red-eyed vireos used edges. Anderson and Shugart

(1974), Stauffer and Best (1980), and Crawford et al. (1981) reported

red-eyed vireos associated with closed canopies and sparse understory.

We also found that red-eyed vireo numbers were correlated with tree

density (r=+ 47.8).

Hooded warblers, Wilsonia citrina, were ubiquitous over treat-

ments and aspects. Plots used by hooded warblers had lower basal area

(x = 7.1 m^/ha) than plots not used (18.6 m^ha) (P<0.05). Occurrence

of hooded warblers was associated with understory diversity (r =+ 59.3)

and understory density (+ 50.1), similar to Anderson and Shugart 's

(1974) findings.

Black-throated green warblers, Dendroica virens, were ubiquitous

over treatments and aspects. Plots used by black-throated green

92 William C. McComb and Robert L. Rumsey

warblers had less leaf cover (x = 26.4%) (hence lower crown cover),

greater slope (x = 49.0%), and higher understory diversity (x = 5.49)

than overall plot averages (x - 56.0, 36.2, and 5.15, respectively).

Freedman et al. (1981) reported adverse effects of forest cutting on

black-throated green warblers in Nova Scotia, but we found this species

tolerant of forest disturbance in Kentucky, though densities were low on

clearcut, 45 kg/ ha and 90 kg/ ha plots.

Ovenbirds, Seiurus aurocapillus, used control and 90 kg/ ha plots

more than clearcut and 68 kg/ ha plots (P<0.05) (Table 3). The 90

kg/ ha of M-3864 resulted in dense understory (x = 105%), partial mid-

story (x = 13%), and a discontinuous overstory (x = 31.9%) (McComb

and Rumsey, in press). Further, ovenbirds used edges more frequently

(13 individuals) than plot centers (2 individuals) (P<0.05), and they

were found more often on north-facing slopes (10 individuals) than on

ridges (1 individual) (P<0.05). Allaire (1978), Robbins (1979), Stauffer

and Best (1980) and Crawford et al. (1981) indicated that ovenbirds are

a forest interior species and that they are adversely affected by forest

disturbance or edge presence, but our study and that of Freedman et al.

(1981) indicated that ovenbirds will occur on edges of some disturbed

areas surrounded by undisturbed forest. Plots where ovenbirds occurred

in our study had higher crown cover (x = 49.7%), leaf cover (x = 73.2%),

basal area (x = 41.7 m^/ha), foliage height diversity (x = 1.44), lower log

density (x = 0.8/ 12 m^), and a shorter distance to water (x - 189.8 m)

than overall plot means (x = 31.5, 56.0, 33.3, 1.30, 1.5, and 335.7,

respectively). Ovenbird numbers were correlated with basal area (r =

+ 54.6), crown cover (r -+ 48.0), and distance to water (r =- 47. 1).

Kentucky warblers, Oporornis formosus, used 68 kg/ ha plots more

than control and clearcut plots (P<0.05) (Table 3). Abundant (x = 1/31

m^) and large-diameter (x = 34.8 cm) snags, low basal area (17.2 m^/ha),

and low midstory cover (17.1%) characterized 68 kg/ ha plots. Kentucky

warblers were found on plots with higher snag density (x = 1/13 m^),

rock cover (x - 8.8%), and lower midstory cover (x = 15.5%), snag dis-

tance (x = 16.2 m), rock distance (x = 14.9 m), and log length (x = 19.7

m), than overall averages ( x = 1/20, 3.3, 28.0, 3.4, 32.8, and 28.4, respec-

tively). Kentucky warbler numbers were correlated with midstory cover

(r =- 52.3) and snag diameter (r =+ 48.6). Kentucky warbler habitat on

our plots was characterized by many logs, rocks and large snags and

little midstory on sites away from permanent water. Log and rock pres-

ence may be important as a nest location for this ground-nesting

species.

Tufted titmice had no association with aspect or edge, but they

were observed most frequently on 68 kg/ ha plots (P<0.05) (Table 3).

The abundance of large-diameter snags (x = 34.8 cm dbh) that occurred

on 68 kg/ ha plots may have influenced use of these plots by this cavity-

Bird Response to Clearings 93

nesting species. Stauffer and Best (1980) reported this species tolerant of

habitat disturbance and Crawford et al. (1981) found titmice using a

wide range of habitats.

Red-bellied woodpeckers used control and 45 kg/ ha plots more

frequently than clearcut or 23 kg/ ha plots (P<0.05) (Table 3). Control

and 45 kg/ ha plots were characterized by having higher basal area (x =

22.5 m^ha) than either clearcut (9.3 m^ha) or 23 kg/ ha plots (17.5

m^/ha). Red-bellied woodpeckers exhibited no association with aspect

or edge, but hairy woodpeckers used ridge-tops (9 individuals) more

(P<0.05) than north- or south-facing slopes (3 and 2 individuals, respec-

tively). Red-bellied woodpeckers used habitats with an open understory

(high crown cover) and high basal area.

Sample sizes were insufficient to identify treatment, aspect, or habi-

tat preferences of 22 other species of breeding birds encountered during

the course of our study (Table 3).

MANAGEMENT IMPLICATIONS

Long-term effects of picloram herbicide application in field situa-

tions on soil invertebrates, invertebrates inhabiting herbicide-created

snags, plant succession, and reproductive physiology of terrestrial verte-

brates have not been thoroughly investigated. If future studies indicate

minimal effects of picloram herbicides on these ecological processes,

then pelletized picloram application may be a more desirable and less

expensive non-game bird management tool in remote or rugged terrain

than manually cutting trees. Four years after application of picloram

herbicide or clearcutting, increases in BSD through changes in species

composition and increased density of some species in the forest may be

expected, but no single treatment will accomplish this goal. Although

we found no differences in BSD among treatments, some species pre-

ferred or occurred exclusively on one or several treatments.

Total bird density was higher on treated plots than control plots in

both winter and spring. This may have been due to the presence of edge

for some species (wintering hairy and red-bellied woodpeckers, red-eyed

vireos, and ovenbirds) and/ or to the changes in the biotic and abiotic

habitat characteristics produced by the treatment. For instance, changes

in habitat characteristics due to clearcutting produced predictable

occurrences of rufous-sided towhees and cardinals. Changes in habitat

structure brought about by herbicide application allowed predictable

occurences of some species (e.g., Kentucky warblers and ovenbirds) but

not others (black-throated green warblers and tufted titmice).

We recommend that a range of pelletized picloram rates from 27 to

68 kg/ ha be used on small plots (0.5 - 1.0 ha) in conjunction with small

clearcuts to provide desirable habitat requirements for as many species

of birds as possible, and to increase the density of many of these species

94 William C. McComb and Robert L. Rumsey

over what would occur in an undisturbed forest. Ninety kg/ ha of

M-3864 was not a desirable treatment since none of the species used

plots with this rate over one of the other treatments. If ten plots per

1,000 ha were treated each year, habitat would be provided on a sus-

tained basis while allowing 100 years for recovery on each site.

Herbicide-treated plots and clearcuts placed on both north-facing and

south-facing slopes would increase the advantages to breeding migrants

and wintering residents or breeding Picids, respectively.

ACKNOWLEDGMENTS.— We wish to thank R. L. Anderson,

D. M. Allen, M. J. Immel, and C. Rowell for advice on statistical ana-

lyses; Dow Chemical Corp. for supplying herbicides; J. B. Davis and the

Robinson Forest staff for assistance with field work; and R. N. Conner,

P. D. Doerr, B. A. Thielges, P. N. Allaire, R. E. Noble, R. B. Hamilton,

R. D. Shipman, R. N. Muller, and C. J. Liu for reviewing an early draft

of the manuscript. The investigation reported in this manuscript (No.

82-8-40) is in connection with Kentucky Agricultural Experiment Sta-

tion Project No. 620 and is published with the approval of the Director.

LITERATURE CITED

Allaire, Pierre N. 1978. Effects on avian populations adjacent to an active

stripmine site. Pp. 232-240 in D. E. Samuel, J. R. Stauffer and W. T.

Mason (eds.). Surface mining and fish/ wildlife needs in the eastern United

States. U. S. Dep. Inter., Fish Wildl. Serv. Program FWS - OBS 78/81.

Anderson, Stanley H., and H. H. Shugart, Jr. 1974. Habitat selection of breed-

ing birds in an east Tennessee deciduous forest. Ecology 55:828-837.

Beaver, Donald L. 1976. Avian populations in herbicide treated brush fields.

Auk 95:543-553.

Best, Louis B. 1972. First-year effects of sagebrush control on two sparrows. J.

Wildl. Manage. 36:534-544.

Carpenter, Stanley B., and R. L. Rumsey. 1976. Trees and shrubs of Robinson

Forest, Breathitt County, Kentucky. Castanea ^^7:277-282.

Conner, Richard N. 1979. Snag management for cavity-nesting birds. Pp. 120-

128 in R. M. DeGraaf (Tech. Coord.). Proceedings of the Workshop on

Management of Southern Forests for Nongame Birds. U. S. Dep. Agric.

For. Serv. Gen. Tech. Rep. SE-14.

, and C. S. Adkisson. 1975. Effects of clearcutting on the diversity of

breeding birds. J. For. 7i:78 1-785.

Crawford, Hewlette S., R. G. Hooper and R. W. Titterington. 1981. Songbird

response to silvicultural practices in central Appalachian hardwoods. J.

Wildl. Manage. 45:680-692.

Bird Response to Clearings 95

Dewey, John B. 1980. "Gridball Pellets" — a new tool for brush control in

pines. For. Farmer ^0(2): 14-15, 34.

Freedman, B., C. Beauchamp, I. A. McLaren and S. I. Tingley. 1981. Forestry

management practices and populations of breeding birds in a hardwood

forest in Nova Scotia. Can. Field-Nat. 95:307-31 1.

James, Francis C, and H. H. Shugart, Jr. 1971. A quantitative method of habi-

tat description. Audubon Field Notes 2'/:727-736.

Kenaga, Eugene E. 1969. Tordon herbicide — evaluation of safety to fish and

birds. Down-to-Earth 25:5-9.

Loftis, David L. 1978. Preharvest herbicide control of undesirable vegetation in

southern Appalachian hardwoods. South. J. Appl. For. 2:51-54.

McCaffery, Kenneth R., F. L. Johnson and L. D. Martoglio. 1974. Maintaining

wildlife openings with pellets containing picloram. Wildl. Soc. Bull. 2:40-45.

McComb, William C, and R. E. Noble. 1980. Small mammal and bird use of

some managed and unmanaged forest stands in the mid-south. Proc. Annu.

Conf. Southeast. Assoc. Fish Wildl. Agencies 5^:482-491.

, and R. L. Rumsey. In press. Habitat characteristics of forest clear-

ings created by picloram herbicides and clearcutting. Proc. Annu. Conf.

Southeast. Assoc. Fish Wildl. Agencies 55.

Murphy, Patrick K., and R. E. Noble. 1972. The monthly availability and use of

browse plants by deer on a bottomland hardwood area in Tensas Parish,

Louisiana. Proc. Annu. Conf. Southeast. Assoc. Fish Wildl. Agencies

2(5:39-57.

Robbins, Chandler S. 1979. Effect of forest fragmentation on bird populations.

Pp. 198-212 in R. M. DeGraaf (Tech. Coord.). Proceedings of the Work-

shop on Management of North Central and Northeastern Forests for Non-

game Birds. U. S. Dep. Agric. For. Serv. Gen. Tech. Rep. NC-51.

Savidge, Julie A. 1978. Wildlife in a herbicide-treated Jeffrey pine plantation in

eastern California. J. For. 7(5:476-478.

Shields, William M., and T. C. Grubb, Jr. 1974. Winter bird densities on north

and south slopes. Wilson Bull. 5(5:125-130.

Shipman, Robert D. 1972. Converting low-grade hardwood forests to Japanese

larch with fenuron herbicides. Tree-Planters' Notes 2^:1-3.

Smith, Kimberly G. 1977. Distribution of summer birds along a forest moisture

gradient in an Ozark watershed. Ecology 55:810-819.

Stauffer, Dean F., and L. B. Best. 1980. Habitat selection by birds of riparian

communities: evaluating effects of habitat alterations. J. Wildl. Manage.

^^:1-15.

Strelke, William K., and J. G. Dickson. 1980. Effect of forest clearcut edge on

breeding birds in east Texas. J. Wildl. Manage. 44:559-561.

Accepted 29 April 1983

Drainage Records and Conservation Status Evaluations

for Thirteen Kentucky Fishes

Melvin L. Warren, Jr. ' and Ronald R. Cicerello

Kentucky Nature Preserves Commission,

Frankfort, Kentucky 40601

ABSTRACT.— Recent ichthyofaunal surveys in Kentucky have

resulted in drainage records, refinement of distributional patterns, and

evaluation of conservation status for 13 fish species. The following

species are recorded for the first time from the Kentucky drainages

indicated in parentheses: Lampetra appendix (Cumberland River),

Umbra limi (West Fork Clarks River), Hybognathus hayi (Tennessee

River), Hybopsis insignis (Red River), Notropis ariommus (Kinnicon-

ick Creek), Notropis telescopus (Green River), Lepomis marginatus

(Tennessee River), Etheostoma camurum (Licking River), Percina

phoxocephala (Red River), and Percina shumardi (Little Sandy River).

In addition, the continued existence of Percina macrocephala in the

Barren River system and Kinniconick Creek, and of Fundulus chryso-

tus in extreme southwestern Kentucky, are confirmed. Evaluation of

recently acquired distributional data necessitates recommended changes

in the conservation status (established by the Kentucky Academy of

Science) of four species: (1) Lepisosteus oculatus should be removed

from the threatened category and reassigned to special concern status;

(2) Hybopsis insignis and Fundulus chrysotus should be elevated from

special concern to threatened and endangered status, respectively; and

(3) Percina phoxocephala does not warrant conservation status recog-

nition.

INTRODUCTION

The freshwater ichthyofauna of Kentucky is one of the most speci-

ose in North America, ranking third behind Tennessee and Alabama

(Burr 1980). Nevertheless, attempts to thoroughly document the distri-

bution and conservation status of this fauna have only recently been

reaHzed through publication of Clay*s (1975) book on Kentucky fishes,

an updated distributional checkHst (Burr 1980), and a Hst of endangered

and threatened Kentucky fishes endorsed by the Kentucky Academy of

Science (Branson et al. 1981b). These efforts stimulated renewed interest

in the Kentucky fish fauna, with emphasis on taxonomic status, refine-

ment of distributions, and re-evaluation of conservation status as exem-

plified by Starnes and Starnes (1978, 1979), Bauer and Branson (1979),

Burr and Mayden (1979), Burr et al. (1980), Starnes (1981), Warren

(1981), Page and Burr (1982), Warren and Cicerello (1982), and others.

' Present address: Department of Zoology, Southern Illinois University at Car-

bondale, Carbondale, Illinois 62901.

Brimleyana No.9:97-109. June 1983. 97

98 Melvin L. Warren, Jr. and Ronald R. Cicerello

Recent ichthyofaunal surveys conducted under the auspices of the

Kentucky Nature Preserves Commission have further contributed to the

refinement of distributional patterns and evaluation of the Kentucky

Academy of Science conservation status (Branson et al. 1981b) of sev-

eral fish species. It is the primary purpose of this report to elucidate and

summarize these findings for a better understanding of Kentucky's speci-

ose ichthyofauna.

SPECIES ACCOUNTS

Several new drainage records, significant distributional informa-

tion, and conservation status evaluations for 13 Kentucky fishes are

presented in the following species accounts. Each account includes col-

lection numbers followed in parentheses by the number of specimens,

standard or total length (TL) range in millimeters, stream and major

drainage, locality, county, and date of collection. All scientific and

common names follow Robins et al. (1980). Collecting materials and

methods were the same as those presented by Harker et al. (1980). All

collections are housed at the Kentucky Nature Preserves Commission

(KNP), pending deposition at Southern Illinois University at Carbon-

dale (SIUC). A number of specimens, as noted, are deposited at the

Kentucky Department of Fish and WildHfe Resources (KFW), the Ken-

tucky Department of Transportation (KDOT), or SIUC.

Lampetra appendix (DeKay). American brook lamprey. KNP uncat.

(1, 182 TL), Cumberland R. (Ohio R. dr.), 30 m below the mouth of

Sulphur Cr., Monroe Co., 22 October 1982.

Burr (1980) considered Lampetra appendix (as L. lamottei) to be

occasional in the upper Barren, Green, Kentucky, and Big Sandy rivers

of Kentucky. The specimen reported here is the first published record

for the Cumberland River of Kentucky, and one of three from the entire

drainage (Rohde 1980). Based on development of dentition, eyes, and

fins, the specimen was judged a sub-adult in the latter stages of trans-

formation; myomeres numbered approximately 69-70. Seagle and Nagel

(1982) noted that metamorphosis of this species in streams of eastern

Tennessee occurred from mid-August through October; this is sup-

ported by capture of a metamorphosing specimen in Kentucky in late

October. The specimen was collected from shallow (15 cm), clear water

with moderate current over a gravel riffle near the shore. When col-

lected the specimen was sluggish, relatively easy to capture, and hemor-

rhaging at the base of the dorsal fins. It should be noted that the Cum-

berland River was at unusually low flow and high water temperatures

due to temporary curtailment of hypolimnetic discharge from upstream

Wolf Creek Dam. Extensive seining in the area produced only a few

specimens of Coitus carolinae (Gill) and a single juvenile Etheostoma

Kentucky Fishes 99

rufilineatum (Cope). We speculate that the lamprey was a waif from

nearby Sulphur Creek, which more closely approximates the preferred

small river and stream habitat (Rohde 1980). Lampetra appendix is

regarded as a species of special concern in Kentucky (Branson et al.

1981b).

Lepisosteus oculatus (Winchell). Spotted gar. KNP Tn04GRV (1,

464 TL; 1 not retained), unnamed wetland (W. Fk. Clarks R. dr.), 0.9

km ENE Clear Springs, Graves Co., 4 May 1982; KNP OOl MCC (1,380

TL; 1 not retained). Metropolis Lake (Ohio R. dr.), 2.3 km N KY 1420

and KY 996 jet, McCracken Co., 6 June 1982; KNP M02CAE (1, 603

TL), Fish Lake (Mississippi R. dr.), at Burkley, Carlisle Co., 20 June

1982; KNP M05HIC (1, 557 TL; 1 not retained), Obion Cr. (Mississippi

R. dr.), 0.1 km downstream from KY 307 crossing, Hickman Co., 22

June 1982.

In Kentucky, the spotted gar was thought to be restricted to the

mainstem and tributaries of the Mississippi, lower Ohio (exclusive of

the lower Tennessee and Clarks rivers), and lower Cumberland rivers

(Burr 1980), until specimens from the Tradewater and Green river

drainages were reported by Warren and Cicerello (1982). More recently.

Rice et al. (1983) pubUshed the first substantiated Kentucky record for

L. oculatus from the lower Tennessee River drainage and, in additon,

six records for locaHties in CarUsle, Fuhon, and Hickman counties. In

light of these records and those presented herein, we believe a reapprai-

sal of the threatened status assigned to L. oculatus by Branson et al.

(1981b) is warranted. The spotted gar is more widespread in Kentucky

than previously thought despite the continuing loss of wetland habitat

favored by the species (Trautman 1981; Warren and Cicerello 1982).

The apparent rarity of the species is probably a result of the difficulty of

sampling the preferred vegetated, wetland habitat which has only recently

begun to be surveyed in Kentucky. We recommend that the status of L.

oculatus be changed from threatened to special concern, so that the

impact of habitat modification can be monitored.

Umbra limi (Kirtland). Central mudminnow. KNP M02GRV (1,

74), unnamed wetland (Terrapin Cr. dr.), 1.45 km S KY 97 and KY

1485 jet at Bell City, Graves Co., 27 April 1982; KNP Tn03GRV (1, 64),

old channel (W. Fk. Clarks R. dr.), at KY 131 crossing. Graves Co., 4

May 1982; KNP Tn04GRV (2, 46-53), unnamed wetland (W. Fk. Clarks

R. dr.), 0.9 km ENE Clear Springs, Graves Co., 4 May 1982; KNP

Tn08GRV (4, 18-21), unnamed wetland (W. Fk. Clarks R. dr.), 1.1 km

SSW mouth Spring Cr., Graves Co., 5 May 1982; KNP M04FUL (4,

40-66), unnamed wetland (Reelfoot Lake dr.), 0.4 km NE Tyler on E

side KY 94, Fulton Co., 24 June 1982; KDOT Q18 (2, -), slough along

KY 94 (Reelfoot Lake dr.), NW of KY 94 and 1500 m NE of Tennessee-

100 Melvin L. Warren, Jr. and Ronald R. Cicerello

Kentucky line, Fulton Co., 12 June 1979; KDOT Q20 (2, -), tributary to

Blue Pond (Reelfoot Lake dr.), KY 311 bridge, Fulton Co., 17 June

1979; KDOT Q27 (-, -), Rittenhouse Slough (Reelfoot Lake dr.), 1700 m

S Bondurant, Fulton Co., 24 August 1978; KDOT Q32 (2, -), unnamed

trib. Running Slough (Reelfoot Lake dr.), N Illinois Central Railroad

and 170 m SE Ledford, Fulton Co., 22 August 1978.

This species was previously known from only three Kentucky local-

ities (Sisk 1978; Burr 1980). Burr (1980) regarded it as rare and Branson

et al. (1981b) listed it as threatened. Umbra limi is apparently firmly

established in Terrapin Creek (Brooks M. Burr, pers. comm.) and the

Reelfoot Lake drainage of extreme southwestern Fulton County, Ken-

tucky, and is sporadically distributed in Clarks River, being most pre-

valent in the West Fork. Although the populations in Clarks River

represent the only published localities of the species in the Tennessee

River drainage, U. limi has also been collected in the Big Sandy River

of Tennessee (David A. Etnier, pers. comm.). These populations are

near the southern periphery of the range (Gilbert 1980a). The apparent

absence of the fish in streams of western Kentucky draining directly into

the Mississippi River (e.g.. Bayou du Chien, Obion and Mayfield

creeks) is zoogeographically puzzling; however, further intensive sam-

pling of wetland habitats in these drainages will probably reveal its

presence. The future existence of U. limi in Terrapin Creek (Obion R.

dr.) and West Fork Clarks River may be jeopardized by drainage of

remaining wetlands as witnessed at two of our collection sites (i.e., KNP

M02GRV, KNP Tn04GRV). Likewise, rapid erosion of the Mississippi

loess bluffs and expansion of agriculture in the floodplain threaten this

and other species inhabiting the Reelfoot Lake drainage in both Tennes-

see (Starnes and Etnier 1980) and Kentucky.

Hybognathus hayi Jordan. Cypress minnow. KNP Tn04GRV (1,

74), unnamed wetland (W. Fk. Clarks R. dr.), 0.9 km ENE Clear

Springs, Graves Co., 4 May 1982.

Recent works addressing the distribution of H. hayi in Kentucky

have revealed records for direct Mississippi and Ohio river tributaries

and floodplain lakes in the extreme western part of the state (Burr et al.

1980) and a relictual population in lower Green River (Warren and

Cicerello 1982). The discovery of the species in the West Fork Clarks

River system represents the first record for the Tennessee River drain-

age in Kentucky, although records are available for the drainage in

Tennessee (David A. Etnier, pers. comm.) and Alabama (Gilbert 1980b).

The specimen was secured from a shallow (<0.6 m) pothole in a recently

cleared and drained wetland. Hybognathus hayi is considered threa-

tened in Kentucky (Branson et al. 1981b) and in consideration of the

elimination and destruction of the preferred wetland habitat by oil

Kentucky Fishes 101

exploration and coal mining (Warren and Cicerello 1982), channeliza-

tion (Burr et al. 1980), and drainage as witnessed herein, the future of

the species in Kentucky is increasingly tenuous.

Hybopsis insignis Hubbs and Crowe. Blotched chub. KNP C04LOG

(1, 89), S. Fk. (Red R. dr.), 1.7 km NE Smith Grove Church, Logan

Co., 13 July 1982; KNP C05LOG (15, 47-79), Red R. (Cumberland R.

dr.), at Dot, Logan Co., 13 July 1982.

In Kentucky, H. insignis was known to persist only in the Little

South Fork Cumberland River of southeastern Kentucky (Marker et al.

1979) and was formerly known to occur in the mainstem of Cumberland

River (Harris 1980) and the lower Tennessee River (Hubbs and Crowe

1956) before impoundment. In the Cumberland River drainage of Ten-

nessee, the species is known from four localities (Harris 1980). The dis-

covery of a substantial population in the Red River represents a new

record for that drainage in Kentucky and adds hope for the continued

existence of the species in the state. Branson et al. (1981b) considered

the species of special concern in Kentucky. In light of strip-mine and oil

field related water pollution in Little South Fork (Harker et al. 1979,

1980) and heavy siltation and pesticide pollution in Red River (pers.

observ.) the species should be considered at least threatened within

Kentucky.

Notropis ariommus (Cope). Popeye shiner. KFW uncat. (2, 45),

Kinniconick Cr. (Ohio R. dr.), near mouth Pipe Lick Cr., Lewis Co., 7

May 1981; KFW uncat. (3, 45-48), Kinniconick Cr. (Ohio R. dr.),

downstream Laurel Fk. mouth, Lewis Co., 7 May 1981.

The distribution of the popeye shiner in Kentucky was previously

defined as the upper Cumberland, Green, Barren, RoUing Fork, and

Kentucky river drainages (Gilbert 1969, 1980c; Burr 1980). Despite this

rather wide distribution, the popeye shiner is sporadic in occurrence and

seldom common and was thus listed as of undetermined status in Ken-

tucky (Branson et al. 1981b). In middle and upper Ohio River tributar-

ies other than those aforementioned, A^. ariommus is known from six

widely separated populations in Indiana, Pennsylvania, and West Virgi-

nia (Gilbert 1969, 1980c). Several of these represent old records for

populations apparently extirpated (Gilbert 1969). The collections

reported herein are a significant eastward range extension in the Ohio

River valley of Kentucky and also close the hiatus between the widely

separated middle and upper Ohio River populations. It is increasingly

apparent from data presented by Gilbert (1969, 1980c) and that of this

report that A^. ariommus once occupied much of the Ohio River valley,

but is now extirpated or reduced in the northern and upper regions of

the valley to widely disjunct, sporadically distributed localities. Kinnic-

onick Creek is a high quality stream with a predominantly forested

102 Melvin L. Warren, Jr. and Ronald R. Cicerello

watershed that has fortuitously escaped degradation from development.

However, proposed plans to develop extensive Devonian age oil shale

deposits in the watershed (Harker et al. 1980) may alter these conditions

and threaten the future existence of A^. ariommus in the drainage.

Notropis telescopus (Cope). Telescope shiner. KNP uncat. (1, 47),

E. Fk. (Barren R. dr.), at mouth of Isenburg Cr., Monroe Co., 12 June

1979; SIUC 3931 (17, 39-57), E. Fk. (Barren R. dr.), at KY 63 crossing,

Monroe Co., 24 September 1981.

Notropis telescopus was known in Kentucky only from the Cum-

berland River (below the Falls) in the southeastern part of the state

(Burr 1980) where it is common to abundant in small to medium-sized,

high quality streams and rivers. Collection of the telescope shiner in

East Fork Barren River represents the first substantiated report of the

species from the Green River system. Woolman (1892) noted the species

as rare in the Little Barren River; however, Gilbert (1969) regarded the

record as erroneous; the specimen(s) was apparently not available for

examination. These records are of particular interest because several

other taxa typical of the Cumberland River of Kentucky and Tennessee,

such as Nocomis effusus Lachner and Jenkins, Notropis leuciodus

(Cope), Fundulus catenatus (Storer), and closely related members of the

subgenera Catonotus, Nanostoma, and Nothonotus of the genus

Etheostoma, are also represented in the ichthyofauna of the Barren and

Green rivers. Furthermore, Lachner and Jenkins (1971) and Zorach

(1972) cited evidence that stream capture has been responsible for ich-

thyofaunal exchange between the Cumberland and Green rivers. The

ichthyofauna of the upper Barren and Green rivers is relatively well

known (Burr 1980), and the failure of workers subsequent to Woolman

to capture A^. telescopus in the drainage presents an enigma in interpret-

ing its native or non-native status. Since Woolman was apparently quite

familiar with A^. telescopus and differentiated it from the similar A^. ari-

ommus (Cope), the likeUhood of misidentification of the Little Barren

River specimen(s) is reduced. Also of interest is the fact that both East

Fork Barren River and Little Barren River headwaters lie in close prox-

imity to Cumberland River streams (Meshack and Marrowbone creeks,

respectively) which harbor large populations of A^. telescopus (pers.

observ.). The possibility of headwater capture is heightened, especially

along the Meshack Creek-East Fork Barren River divide, by the pres-

ence of karst development including numerous sinkholes and subterra-

nean drainage. This is best illustrated on the southeast corner of the U.

S. Geological Survey 7.5 minute Sulphur Lick quadrangle map (Harris

1964). The current extent of dispersal of A^. telescopus in the Barren

River is unknown, but the capture of the species only in East Fork

implies a localized distribution. Based on the available evidence, the

origin of A^. telescopus in the Green River is unclear; however, as noted

Kentucky Fishes 103

by Jenkins et al. (1971) even the most suggestive evidence concerning

stream capture may lead to invalid conclusions, and limited distribu-

tions may result from introduction rather than natural factors. Further

collecting aimed at probable theatres of stream capture should further

elucidate the native or non-native status of A^. telescopus as well as rela-

tionships among other shared taxa.

Fundulus chrysotus (Giinther). Golden topminnow. KNP M03FUL

(9, 42-51), Running Slough (Reelfoot Lake dr.), at Ledford, Fulton Co.,

24 June 1982.

According to Burr (1980), the golden topminnow was known in

Kentucky only from Open Pond, Fulton County, where two collections

were made by Sisk (1973). Field observations made during the summer

of 1982 revealed that Open Pond and surrounding wetlands had been

drained, cleared, and converted to agricultural land. Recent collecting

efforts in appropriate habitat throughout the general drainage area in

Kentucky revealed that the golden topminnow is now known from a

single extant population in Running Slough. Although F. chrysotus is

listed as of special concern in Kentucky by the Kentucky Academy of

Science (Branson et al. 1981b), the species should be placed in the

endangered category in light of its extremely limited distribution in

Tennessee (Starnes and Etnier 1980) and the potential for rapid loss of

habitat as witnessed for Open Pond.

Lepomis marginatus (Holbrook). Dollar sunfish. KNP Tn04GRV

(1, 49), unnamed wetland (W. Fk. Clarks R. dr.), 0.9 km ENE Clear

Springs, Graves Co., 4 May 1982; KNP Tn08GRV (1, 72), unnamed

wetland (W. Fk. Clarks R. dr.), 1.1 km SSW mouth Spring Cr., Graves

Co., 5 May 1982.

Lepomis marginatus, previously known from only two Kentucky

localities, neither of them in the Tennessee River drainage (Burr 1980),

was discovered by Rice et al. (1983) at a third site in West Fork Clarks

River. These records represent an addition to the ichthyofauna of the

Tennessee River in Kentucky. The species is apparently widely distrib-

uted in West Fork Clarks River and at some localities occurs in good

numbers (Rice et al. 1983). The populations in West Fork, although

threatened by wetland drainage as witnessed by the authors (e.g., at

KNP Tn04GRV), may prove to be critical in preserving the species as a

viable member of the native Kentucky ichthyofauna. Branson et al.

(1981b) listed the species as threatened.

Etheostoma camurum (Cope). Bluebreast darter. KNP LOIBAT

(26, 34-54), Licking River (Ohio R. dr.), at mouth Slate Cr., Bath Co.,

16 September 1982.

Etheostoma camurum was previously known in Kentucky from the

upper Cumberland (below the Falls) and upper Kentucky river drain-

ages (Burr 1980; Zorach 1972). The collection reported herein represents

104 Melvin L. Warren, Jr. and Ronald R. Cicerello

an addition to the ichthyofauna of the speciose Licking River system

and reduces the distributional hiatus among known populations in the

upper and middle Ohio River drainage (Zorach 1972). Ahhough pre-

vious surveys of the Licking River have not revealed E. camurum

(Woolman 1892; Welter 1938; Clark 1941a, b; Jones 1970), other

members of the subgenus Nothonotus (i.e., E. maculatum Kirtland and

E. tippecanoe Jordan and Evermann) have been reported from the

drainage (Woolman 1892; Clark 1941a, b; Burr 1980). However, E.

maculatum, often confused with E. camurum (Zorach and Raney 1967),

is known in the drainage only from a collection made by J. A. Henshall

in South Fork (Woolman 1892). Henshall apparently recognized both

species, according to details presented in Trautman (1981), and the E.

maculatum record is considered valid by some authors (Zorach and

Raney 1967; Burr 1980), although Etnier (1980) did not include the

Licking River in its distribution. We speculate that E. camurum was

missed in the Licking River by previous investigators because popula-

tions are often localized (Stauffer 1980). Moreover, large stream or river

habitat often occupied by members of the subgenus Nothonotus is diffi-

cult to collect and has elsewhere recently yielded species missed during

many years of collecting (Williams and Etnier 1978). Our specimens

were collected from a 0.3-0.6 m deep riffle habitat with moderate to

swift current. Substrate consisted of bedrock overlain with slab boulder

where the current was swiftest, and cobble/ gravel in areas of moderate

flow. Additional collecting in the poorly sampled mainstem of the Lick-

ing and South Fork Licking rivers (Burr 1980) is necessary to determine

the extent of distribution of E. camurum and to verify the existence of

E. maculatum in the drainage.

Percina macrocephala (Cope). Longhead darter. KNP uncat. (1,

72), Kinniconick Cr. (Ohio R. dr.), 2.1 km upstream from mouth Pine

Br., Lewis Co., 5 May 1981; KNP uncat. (4, 45-52), Kinniconick Cr.

(Ohio R. dr.), near mouth Pipe Lick Cr., Lewis Co., 7 May 1981; KNP

uncat. (2, 51-75), Kinniconick Cr. (Ohio R. dr.), between Mill and Lea-

therbelly branches, Lewis Co., 14 May 1981; KNP GOl ALL (15, 67-90),

Trammel Fk. (Barren R. dr.), at old state rd. ford 1.55 km N of Red

Hill, Allen Co., 14 July 1982; KNP G05WAR (6, 36-72), Trammel Fk.

(Barren R. dr.), at ford 0.2 km upstream from mouth Drakes Cr., 16

July 1982.

According to Page (1978) and Burr (1980), the longhead darter

occurs sporadically in Kentucky in the upper Barren, upper Green, Ken-

tucky, Licking, and Big Sandy river systems and Kinniconick Creek and

has apparently been extirpated from the Cumberland River. Burr (1980)

noted that the species was once common in the Barren River prior to

impoundment of Barren River Reservoir as indicated in pre-

Kentucky Fishes 105

impoundment surveys conducted by the Kentucky Department of Fish

and Wildlife Resources. Our collections in Trammel Fork (Barren R.

dr.) indicate the species continues to persist in good numbers in tributar-

ies unaffected by impoundment. The only previous collection of P.

macrocephala in Kinniconick Creek is based on a single specimen col-

lected in the 1930s and housed at the University of Louisville. Our

observations indicate that the species is moderately common in approp-

riate habitat along the Kinniconick Creek mainstem. In spite of these

relatively healthy populations in the Barren River drainage and Kinnic-

onick Creek, the conservation status of the species in Kentucky should

remain threatened (Branson et al. 1981b) because of pollution and habi-

tat destruction associated with coal mining in the upper Kentucky, Lick-

ing, and Big Sandy rivers (Harker et al. 1979), the extirpation of the

species from the Cumberland River of Kentucky (Page 1978), and the

threat of oil shale development in the Kinniconick Creek drainage

(Harker etal. 1981).

Percina phoxocephala (Nelson). Slenderhead darter. KNP CO 1 LOG

(1, 49), Whippoorwill Cr. (Red R. dr.), 0.7 km W Millertown Church,

Logan Co., 8 July 1982; KNP C03LOG (3, 58-63), S. Fk. (Red R. dr.),

Kentucky-Tennessee line, Logan Co., 13 July 1982; KNP C04LOG (5,

65-69), S. Fk. (Red R. dr.), 1.7 km NE Smith Grove Church, Logan

Co., 13 July 1982; KNP C05LOG (3, 61-84), Red R. (Cumberland R.

dr.), at Dot, Logan Co., 13 July 1982; KNP C02TRI (5, 55-67), Little R.

(Cumberland R. dr.), 1.3 km downstream from KY 1253 crossing, Trigg

Co., 6 July 1982.

Percina phoxocephala was previously unknown from the Red

River of Kentucky and Tennessee (Starnes and Etnier 1980; Thompson

1980), and Burr (1980) noted its former occurrence in the lower Cum-

berland River. The collections noted herein are new records for the Red

River and substantiate the persistence of the species in a lower Cumber-

land River tributary. In Kentucky, the species was regarded as of special

concern (Branson et al. 1981b) and was noted by Burr (1980) as occa-

sional within several drainages. Recent collections made by the authors

and fresh material examined by us indicate good populations of P.

phoxocephala in Tygarts Creek of eastern Kentucky (Warren 1981, and

unpublished), and upper and lower Green, Barren, and Rough rivers

(Retzer et al. 1983; Warren and Cicerello 1982, and unpubhshed). The

slenderhead darter has been taken by others in recent years from several

stations each in Eagle Creek (lower Kentucky River) (Horseman and

Branson 1973) and Salt River and tributaries (Hoyt et al. 1979). It

therefore appears that retention of the species on the Kentucky Academy

of Science list of rare fishes is unwarranted.

106 Melvin L. Warren, Jr. and Ronald R. Cicerello

Percina shumardi (Girard). River darter. KNP SOI GUP (2, 40-44),

Little Sandy R. (Ohio R. dr.), 0.5 km W Argillite, Greenup Co., 14

September 1982.

According to Burr (1980), P. shumardi is sporadic and uncommon

in every major river of the state except the Salt and Big Sandy rivers

and the direct Ohio River tributaries in extreme northeastern Kentucky.

Collection of the species in the Little Sandy River is a new record for

this system and suggests the stream is in need of further study despite

recent collections (Marker et al. 1979; Branson et al. 1981a). It further

suggests that the species will eventually be taken in adjacent rivers and

streams (e.g.. Big Sandy River, Kinniconick and Tygarts creeks). The

rarity of the species is partially attributable to difficulty in collecting the

preferred habitat (Trautman 1981). The specimens reported herein were

taken from a deep (1.3 m), swift chute over a substrate of large cobble.

Apparently this is the last shoal present on the Little Sandy River

before its waters are embayed by the Greenup Lock and Dam on the

Ohio River. The river darter is considered threatened in Kentucky

(Branson et al. 1981b); however, we believe that further collecting in

appropriate habitat will reveal new populations and result in its removal

from the state threatened category.

ACKNOWLEDGMENTS.— Our appreciation is extended to Richard

R. Hannan, Director of the Kentucky Nature Preserves Commission,

for support throughout this effort. Special thanks go to Brian D. And-

erson, Keith E. Camburn, and Dan E. VanNorman for assistance with

field work. The following individuals and their agencies or institutions

kindly confirmed identifications, shared collecting information, or pro-

vided specimens and other courtesies: Brooks M. Burr, Southern Illinois

University at Carbondale; David A. Etnier, University of Tennessee;

John L. Harris, Arkansas Department of Transportation; Lewis E.

Kornman, Kentucky Department of Fish and Wildlife Resources; and

Stephen P. Rice, Kentucky Department of Transportation. Our grati-

tude is also expressed to Branley A. Branson, Eastern Kentucky Univer-

sity; Brooks M. Burr; Keith E. Camburn, Kentucky Nature Preserves

Commission; and David A. Etnier for critical review of the manuscript.

LITERATURE CITED

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Kentucky Fishes 107

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Clark, Minor E. 1941a. A list of the fishes in northeastern Kentucky. Ky. Dep.

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of Fishes from the United States and Canada. 4th ed. Am. Fish. Soc. Spec.

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Accepted 16 April 1983

Winter Food Habits of Bobcats in North Carolina

Anne M. King

Department of Zoology,

Richard A. Lancia

Department of Forestry,

AND

S. Douglas Miller, iDavid K. Woodward and Jay D. Hair '

Department of Zoology

North Carolina State University, Raleigh, North Carolina 27650

ABSTRACT.— Carcasses of 505 bobcats, Felis rufus (229 9, 276 5),

were collected from October through March in 1978-79 and 1979-80.

Stomach contents were analyzed and the results summarized by physi-

ographic regions, sex, and age. The top eight prey groups, ranked by

frequency of occurrence, were rabbits, Sylvilagus spp.; birds; cotton

rats, Sigmodon hispidus; white-tailed deer, Odocoileus virginianus;

rodents; gray squirrels, Sciurus carolinensis; raccoons, Procyon lotor,

and opossums, Didelphis virginianus. Ranked by frequency of occur-

rence, rabbits were first in the Coastal Plain and cotton rats first in the

Piedmont Plateau. Adult male bobcats consumed larger prey than did

adult females or kittens. Kittens tended to exploit smaller and a wider

variety of prey items than did adults.

INTRODUCTION

Because of restrictions placed on harvesting many species of spot-

ted cats, the value of bobcat, Felis rufus, pelts increased greatly on the

international fur market. In 1977 concern about the national status of

bobcats prompted the Council on International Trade of Endangered

Species of Flora and Fauna (CITES) to list them as a species that could

become threatened with extinction unless trade was subject to regula-

tion. As part of a program to learn more about the status and ecology

of bobcat populations in North Carolina, we initiated a study of bobcat

food habits based on analyses of trapper-harvested carcasses. Other

aspects of our research program are reported elsewhere (Lancia et al., in

press).

Previous food habits studies in the southeastern United States

reported rabbits, Sylvilagus spp., as the major food item (Progulske

1952; Davis 1955; Fritts 1973; Buttrey 1974; Fox and Fox, in press). In

addition, cotton rats, Sigmodon hispidus; white-tailed deer, Odocoileus

' Present address: National Wildlife Federation, 1412 Sixteenth Street, N.W.,

Washington, D.C. 20036

Paper number 7035 of the Journal Series of the North Carolina Agricultural

Research Service, North Carolina.

BrimleyanaNo.9:lll-122. June 1983. Ill

112 Anne M. King, et al.

virginianus; squirrels, Sciurus carolinensis; voles, Microtus spp.; and

birds were frequently identified food items (Progulske 1952; Davis 1955;

Kight 1962; Fritts 1973; Buttrey 1974; Miller and Speake 1978).

MATERIALS AND METHODS

Skinned bobcat carcasses were provided by fur dealers and trappers

during the 1978-79 and 1979-80 trapping seasons. The carcasses were

weighed, sexed, and necropsied. Adult and kitten (< 8 months) age

classes were assigned using lacteal tooth replacement criteria (Crowe

1975).

Stomach contents, prepared following Korschgen (1980), were

identified to order and, when possible, to species. Data were recorded

by percent frequency of occurrence because not all material in each

stomach could be identified, hence percent volume could not be meas-

ured precisely. Food items were identified macroscopically or, when

necessary, mammals were identified by microscopic examination (at

lOOx and 400x) of sample hairs using hair keys (Spiers 1973; Moore et

al. 1974).

Statistical Analyses and Interpretation

The state was divided into three physiographic regions (Stuckey

1965), and the data were examined at statewide and regional levels.

Comparisons of carcass weights were made using least-squares fit to a

fixed-effect linear model using the Scientific Analysis System (SAS;

Barr et al. 1979). Weights of kittens and adults were analyzed sep-

arately. Food items were ranked by frequency of occurrence, and differ-

ences in ranks were evaluated using Wilcoxon's Rank Test (Wilcoxon

1945). Chi-square tests were used to evaluate differences in prey selec-

tion. Calculation of expected frequencies of prey items for chi-square

tests were based on an assumed equal probability of occurrence in a

particular sex and age group.

Caution must be observed when making inferences about the pro-

portion of food items present in the diet based solely on stomach anal-

yses because: 1) smaller prey species are frequently under represented in

stomach analyses due to differences in digestibility (Weaver and Hof-

fman 1979; Merriwether and Johnson 1980); 2) some food items may

have been eaten incidentally as trap bait, garbage or carrion; and 3)

frequency of occurrence data cannot be used to evaluate food preferen-

ces without estimates of both prey abundance and availability.

RESULTS AND DISCUSSION

From October through March 1978-79 and 1979-80, 505 bobcats

(229 $, 276 5) were collected. Weights between samples collected each

year were not different (p>0.05); therefore, the samples were pooled for

subsequent analyses. Among the three physiographic regions, weights

Bobcat Winter Food Habits

113

were different (p<0.01) for both age groups, but no clear biological sig-

nificance in the trends was apparent (Tables 1, 2). Weights of females

were less than males (p<0.01) for both adults and kittens. Sex ratios of

adults and kittens were not different among regions or between yearly

samples (p>0.05).

Table 1. Analysis of variance for weights of skinned carcasses of 505 bobcats

collected in 1978-79 and 1979-80 in North Carolina.

Adults (N=355)

Source DF F value

Kittens (N= 150)

Source DF F value

Sample

Region (Sample)

Sex (Region)

0.02

4.13**

134.74**

Sample

Region (Sample)

Sex (Region)

1.75

5.24**

3.84**

** Significant at the P<0.01 level.

Table 2. Skinned weights (kg) of 505 bobcats collected in the three physiograph-

ic regions of North CaroHna.

Of 473 stomachs suitable for examination, 398 (179 $, 219 $) con-

tained food. This sample comprised 257 adults and 141 kittens. Stom-

achs containing food were collected from 50 of the 100 counties in the

state, with the largest samples in the Coastal Plain (307) and Piedmont

Plateau (73) regions and the smallest sample (18) from the Mountains

(Fig. 1).

Some disproportionate sampling was evident in these regions, with

one or two counties contributing more than a third of the total number

114

Anne M. King, et al.

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Bobcat Winter Food Habits

115

of stomachs for the region. To determine if these county samples biased

the total regional sample, we ranked the top eight food items in these

counties and compared them to the top eight food items in the remain-

ing counties of the region. The differences in ranks were not statistically

significant (p>0.05).

Mammals were the most frequently represented food group, with

20 species identified statewide (Table 3). The top eight food items

ranked by frequency of occurrence were rabbits; birds; cotton rats;

white-tailed deer; small rodents; gray squirrels; raccoons, Procyon lotor;

and opossums, Didelphis virginianus. Small numbers of reptiles, am-

phibians, fish, and insects also were identified.

Rabbits were the most frequently identified species in bobcat stom-

achs (Table 3, Fig. 2). The percent occurrence of rabbits decreased from

the Coastal Plain to the Mountain region (p<0.05), probably in response

to differences in abundance of rabbits in the regions rather than to shifts

in prey preference. The low frequency (5.6%) of rabbits in stomachs

from the Mountain region contrasted the findings of Progulske (1952)

ADULT MALES (156 STOMACHS)

MISC. PREY (1,8%)

OTHER MAMMALS

(4 4%)

SMALL RODENTS

( 13%)

SQUIRREL (4.8%)

<DS

(16 7%)

RACCOON (7 5%)

DEER (172%)

COTTON RAT (10 6%)

ADULT FEMALES (134 STOMACHS)

-MISC PREY (18%)

UNKNOWN MAMMALS

(77%)

SQUIRREL (4 7%)

RACCOON ( I % )

DEER (5.7%)

COTTON RAT (12%)

KITTENS (108 STOMACHS)

MISC PREY (3.7%)

SQUIRREL (4.3%)

DEER (1.8%)

COTTONRAT (15.8%)

Fig. 2. Relative percent frequency of occurrence of food items identified in adult

male, adult female, and kitten bobcat stomachs. Small rodents include voles,

marsh rice rat, and mice; other mammals include opossum, muskrat, pig, rats,

chipmunk, flying squirrel, and beaver; miscellaneous prey include reptiles,

amphibians, fish, and arthropods.

116

Anne M. King, et al.

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Bobcat Winter Food Habits

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118 Anne M. King, et al.

who examined 25 stomachs from the Pisgah and Nantahala National

Forests and reported 24% contained rabbits.

Birds were found in a greater percentage of the stomachs in 1978-79

than in 1979-80 (p<0.05). In both years the percent occurrence of birds

was greater in stomachs from the Mountains than from the Piedmont

region (p<0.05). Birds were found in proportionately more adult female

and kitten stomachs than in adult male stomachs (Fig. 2). Although this

relationship was not statistically significant in our study, Fritts and Sea-

lander (1978) also observed a similar trend in Arkansas.

Statewide, cotton rats were the second ranking mammal by fre-

quency of occurrence. A high occurrence of cotton rats in kitten stom-

achs, although not significantly greater than in adults (p>0.05), reflects

a trend in prey size similar to that observed for birds, i.e., smaller prey

occurred more frequently in adult females and kittens than in adult

males.

The percent occurrence of cotton rats was highest in the Piedmont

region where this species was the major food item (Table 3). Beasom

and Moore (1977) and Miller and Speake (1978) have shown that in

habitats where cotton rats were the most abundant small mammal, they

were the major prey item. Cotton rats ranked second in frequency in the

Coastal Plain region, but were found less than one-third as frequently as

rabbits, the major food item (Table 3). Cotton rats were not found in

stomachs examined from the Mountain region.

Deer were identified in stomachs from all regions and ranked third

statewide among mammals in frequency of occurrence (Table 3). Al-

though a major food item in the Mountain region, the percent occur-

rence of deer was not significantly different (p>0.05) from that found in

other regions.

Progulske (1952) reported deer as a winter food of bobcats in west-

ern North Carolina and Virginia and suggested they were eaten as car-

rion. Based on 124 scats from Virginia, he concluded deer were con-

sumed primarily during the fall and winter rather than in spring or

summer, and suggested hunting season casualties on deer might be par-

tially responsible. Managers of North Carolina wildlife areas reported

16 deer killed by bobcats in 1968 (Barrick 1969).

Deer were found more frequently in stomachs of adult males than

in adult females or kittens (Fig. 2). In the Coastal Plain, observed fre-

quencies of deer were greater for adult males and less for adult females

and kittens than expected frequencies (p<0.05), possibly indicating a

selective preference for deer by adult males. Observed versus expected

frequencies of deer were not different (p>0.05) between the Piedmont or

Mountain regions, probably due to the small number of stomachs

examined from these regions.

Bobcat Winter Food Habits 119

Greater use of deer by adult male bobcats may be related to differ-

ences in average weight and/ or size of home ranges of males and

females. Weights of males were larger than those of females, and a tele-

metry study in the Coastal Plain region indicated the average size of

adult male home ranges (N=5) was about 41% larger than home ranges

of adult females (N=3) (Lancia et al., in press). Similar ratios of male to

female ranges were observed by Bailey (1974) in Idaho, Hall and New-

som (1976) in Louisiana, and Miller (1980) in Alabama.

Gray squirrels ranked fourth in overall frequency of occurrence

among mammals (Table 3). The relative percent frequencies of squirrels

in this study were similar for all bobcat sex and age groups (Fig. 2).

Squirrels were found in two stomachs from the Mountain region and

one stomach from the Piedmont region. Similarly, Progulske (1952)

found squirrels in two of 25 stomachs he examined from western North

Carolina. In the Coastal Plain region, squirrels ranked fourth in fre-

quency of occurrence.

Raccoons ranked fifth among mammals in the statewide sample

and occurred only in stomachs from the Coastal Plain and Piedmont

regions (Table 3). Raccoons occurred primarily in adult male stomachs

and were not found in any kitten stomachs (Fig. 2). The greater occur-

rence of raccoons in adult male stomachs again may reflect the greater

weight and larger home range of male bobcats.

Small rodents were infrequent food items. Voles, Microtus penn-

sylvanicus and M. pinetorum, were the most frequently identified small

rodents and occurred in stomachs from all regions. Voles ranked first

(with deer) in percent occurrence in the Mountain region (Table 3) and

were found primarily in stomachs of kittens and adult females. In the

Coastal Plain, adult females and kittens had greater observed than

expected frequencies of voles (p<0.05). Other small rodents were prey

of adult females and kittens but were seldom taken by adult males (Fig.

2). Fritts and Sealander (1978) also reported that occurrences of rats

and mice were greater in the stomachs of adult females and kittens than

in adult male bobcats.

We examined 141 kitten (59% males) stomachs containing food.

Like adults, kittens relied primarily on rabbits and cotton rats for food

(Fig. 2); however, the occurrence of some items in kitten stomachs was

notably different from adults. Deer occurred in only three (2.1%) kitten

stomachs, compared to 51 (19.8%) adult stomachs. Two intermediate-

sized prey species, raccoons and opossums, occasionally were eaten by

adults but were absent or rarely found in the kitten sample (Fig. 2). The

low occurrence of large and intermediate-sized prey species in kitten

stomachs probably reflected an inability of kittens to capture larger-sized

prey. The infrequent occurrences of these prey species in kitten stom-

120

Anne M. King, et al.

achs may have been the result of food provided by the aduh female, or

evidence of feeding on carrion.

A greater variety of food items was eaten by kittens than by adults.

For example, kitten stomachs represented 28% of the total sample, but

accounted for 71% of the occurrences of Peromyscus spp. and 57% of

the voles. Fritts and Sealander (1978) also reported a greater use of

cricetid rodents by kittens than by adults.

Although prey species in the cotton rat-rabbit size range (0.1-2.0

kg) were major components in the diet of all bobcat sex and age classes,

some differential selection was evident. Kittens and adult females were

the principal consumers of the smallest prey (<0. 1 kg), while the largest

prey species (>2.0 kg) were consumed most frequently by adult males

(Fig 3). The greatest occurrence of prey in the 0. 1 - 2.0 kg range may

represent a specialization of bobcats for a selected prey size (Miller and

Speake 1978).

o

3

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o

or

LU

40 r

30

20

10

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ADULT cT

oL

0.0 - 0. 1

0.1-0.4

0.4-2.0 2.0-16.0 16.0-120.0

PREY SPECIES WEIGHT CLASSES (kg)

Fig. 3. Relative percent occurrences of different weight classes of mammalian

prey items identified in adult male, adult female, and kitten bobcat stomachs.

Species included in the weight (kg) classes are: 0.0-0.1 (mice); 0.1-0.4 (chipmunk,

cotton rat, woodrat, and Norway rat); 0.4-2.0 (squirrel, rabbit, muskrat); 2.0-

16.0 (opossum, raccoon); and 16.0-120.0 (deer).

Bobcat Winter Food Habits 121

CONCLUSIONS

The results of our study are similar to previous winter food habits

studies in the southeastern United States and indicate rabbits, birds,

cotton rats, and deer were the top four prey items by percent occurrence

in stomachs of bobcats collected statewide in North Carolina during

winter. Furthermore, adult males appeared to select the larger prey

items (>2.0 kg), and adult females and kittens the smaller prey items

(<0. 1 kg). All sex and age classes apparently preferred intermediate-

sized prey (0.1 - 2.0 kg).

The identification of food habits has management imphcations.

The prey species most used by bobcats as food items were species

adapted to early successional stages (e.g., rabbits, cotton rat, and deer).

An interpretation based on prey selection suggests bobcats are not a

climax species, but rather a subclimax species, benefitting from land

management practices that maintain early successional habitats.

ACKNOWLEDGMENTS— The U. S. Fish and Wildlife Service, the

N. C. Wildlife Resources Commission (NCWRC), the N. C. Agricul-

tural Research Service, and North Carolina State University provided

funding. The NCWRC collected carcasses from fur dealers and made

them available to us. Numerous trappers cooperated by providing car-

casses. D. Hazel, L. Petrovick, D. Stewart, J. Wooding, J. Henegar, S.

Habel, and K. Lewis assisted in the necropsy of the carcasses. A. Bras-

well of the N. C. State Museum of Natural History identified the herpe-

tological material.

LITERATURE CITED

Bailey, Theodore N. 1974. Social organization in a bobcat population. J. Wildl.

Manage. 55:435-446.

Barr, Anthony J., J. H. Goodnight, J. P. Sail and J. T. Helwig. 1979. User's

guide to SAS 1979. SAS Institute, Inc., Raleigh. 474 pp.

Barrick, Frank B. 1969. Deer predation in North Carolina and other southeast-

ern states, pp. 25-31 in L. K. Halls (ed.). White-tailed deer in the southern

forest habitat. South. For. Exp. Sta., USDA, Nacogdoches, TX. 130 pp.

Beasom, Samuel L., and R. A. Moore. 1977. Bobcat food habit response to a

change in prey abundance. Southwest. Nat. 27:451-457.

Buttrey, George W. 1974. Food habits and distribution of the bobcat, Lynx

rufus rufus (Schreber), on the Catoosa Wildlife Management Area. Tenn.

Wildl. Res. Agency Tech. Rep. No. 75-12. Nashville. 64 pp.

Crowe, Douglas M. 1975. Aspects of aging, growth and reproduction of bobcats

from Wyoming. J. Mammal. 55:177-198.

Davis, James R. 1955. Food habits of the bobcat in Alabama. Unpubl. M. S.

Thesis, Alabama Polytech. Inst., Auburn. 79 pp.

122 Anne M. King, et al.

Fox, Lloyd B., and J. Fox. In press. Population characteristics and food habits

of bobcats in West Virginia. Proc. Annu. Conf. Southeast. Assoc. Fish

Wildl. Agencies.

Fritts, Steven H. 1973. Age, food habits and reproduction of the bobcat {Lynx

rufus) in Arkansas. Unpubl. M. S. Thesis, Univ. Arkansas, Fayetteville. 80 pp.

, and J. A. Sealander. 1978. Diets of bobcats in Arkansas with special

reference to age and sex differences. J. Wildl. Manage. 2^:226-228.

Hall, Harlan T., and J. D. Newsom. 1976. Summer home ranges and move-

ments of bobcats in bottomland hardwoods of southern Louisiana. Proc.

Annu. Conf. Southeast. Assoc. Fish Wildl. Agencies i0:427-436.

Kight, James. 1962. An ecological study of the bobcat {Lynx rufus Schreber), in

west-central South Carolina. Unpubl. M. S. Thesis, Univ. Georgia, Athens.

52 pp.

Korschgen, Leroy J. 1980. Procedures for food habits analysis, pp. 1 13-127 in S.

D. Schemnitz (ed.). Wildlife Management Techniques, 4th ed. The Wildlife

Society, Washington. 686 pp.

Lancia, Richard A., D. K. Woodward and S. D. Miller. In press. Summer

movement patterns and habitat use by bobcats on Croatan National Forest,

North Carolina. International Cat Symposium, Texas A&I Univ., Kings-

ville, Oct. 4-6, 1982.

Merriwether, David, and M. K. Johnson. 1980. Mammalian prey digestibility

by coyotes. J. Mammal. 67:774-775.

Miller, Stuart D., and D. W. Speake. 1978. Prey utilization by bobcats on quail

plantations in southern Alabama. Proc. Annu. Conf. Southeast. Assoc.

Fish Wildl. Agencies 52:100-1 1 1.

1980. Ecology of the bobcat in south Alabama. Unpubl. Ph.D. Dis-

sert., Auburn Univ., Auburn. 168 pp.

Moore, Tommy D., L. W. Spence and C. E. Dugonolle. 1974. Identification of

the dorsal guard hairs of some mammals of Wyoming. Wyoming Game

FishDep. Bull. 14. 177 pp.

Progulske, Donald R. 1952. The bobcat and its relation to prey species in Virgin-

ia. Unpubl. M. S. Thesis, Virginia Polytech. Inst., Blacksburg. 135 pp.

Spiers, James E. 1973. A microscopic key to the hairs of Virginia land mam-

mals. Unpubl. M. S. Thesis, Virginia Polytech. Inst., Blacksburg. 106 pp.

Stuckey, Jasper L. 1965. North Carolina: Its Geology and Mineral Resources.

N. C. Dep. Conserv. Develop., Raleigh. 550 pp.

Weaver, John L., and S. W. Hoffman. 1979. Differential detectability of rodents

in coyote scats. J. Wildl. Manage. -^3:783-786.

Wilcoxon, Frank. 1945. Individual comparisons by ranking methods. Biomet-

rics 1:80-83.

Accepted 17 September 1983

Foods and Feeding Behavior of Sauger,

Stizostedion canadense (Smith) (Pisces: Percidae),

from GaUipoUs Locks and Dam, Ohio River

Steven I. McBride and Donald Tarter

Department of Biological Sciences,

Marshall University, Huntington, West Virginia 25701

ABSTRACT. — The food habits of 151 sauger, Stizostedion canadense

(Smith), from the Ohio River at Gallipolis Locks and Dam, were stu-

died from March 1981 through April 1982. Seventy percent of the

stomachs contained food, which was analyzed for identification. Fishes

were the primary food, both by weight (99.7%) and frequency of

occurrence (100%). Emerald shiners, Notropis atherinoides Rafinesque,

and gizzard shad, Dorosoma cepedianum (Lesueur), comprised 96% of

all identifiable items. Emerald shiners were numerically and by weight

the most important item; they occurred in 49% of stomachs that con-

^' tained food, and made up 32% of food biomass. Terrestrial and aquat-

ic invertebrates were of minor dietary importance. Seasonal variation

in sauger foods was minimal. Size had no influence on the kind of

food contained in the stomachs of adult sauger. Sauger appeared to

feed most intensively in the fall. Regression analysis showed a signifi-

cant negative correlation (r = -0.87) between sauger caught by anglers

and water temperature.

INTRODUCTION

The sauger, Stizostedion canadense (Smith), is an important sport

and commercial fish (Scott and Grossman 1973). Information on its

foods and feeding behavior are essential to our understanding of percid

communities and to the effective management of these resources. Sev-

eral investigators, including Forbes (1878, 1888) and Richardson (1920),

Pearse (1921), Dendy (1946), and Priegel (1969), provided brief notes on

sauger food habits, but only one detailed study was concerned with a

population in a riverine habitat (Vanicek 1964). Our study was initiated

in March 1981 and continued through April 1982. Its objectives were to

determine: (1) what foods are most important to the sauger population,

(2) if food habits varied with season, (3) if food habits changed as

sauger increased in size, and (4) how water temperature correlated with

sauger catch.

METHODS AND MATERIALS

Study Area

Gallipolis Locks and Dam is one in a series of flood and naviga-

tional control dams situated on the Ohio River. It is located at river

mile (RM) point 279.2 (38°40'53'', 82°ir22"), between Galia County,

BrimleyanaNo.9:123-134. June 1983. 123

124 Steven I. McBride and Donald Tarter

Ohio, and Mason County, West Virginia (U. S. Army Corps of Engi-

neers 1980a). The Gallipolis pool extends 67.6 km (42 mi.) upriver to

Racine Locks and Dam and 49.4 km (31 mi.) up the Kanawha River to

Winfield Locks and Dam. The Gallipolis Dam marks the northern

boundary of the Greenup navigational pool, which extends 75.8 km (62

mi.) downriver to Greenup Locks and Dam. The floodplain of the river

at the Gallipolis Dam is slightly more than 1.61 km (1 mi.) wide. The

river channel has an average width of 366 m, and its depth varies from

approximately 7 to 10 m, with a navigable depth of 2.7 m (U. S. Army

Corps of Engineers 1980b).

Specimens were collected from several locations in the Greenup

pool and once from inside Belleville Locks and Dam. However, the

main collecting site was situated just below Gallipolis Locks and Dam.

This area is steeply sloped and becomes very rocky, then levels toward

the shoreline. Approximately 2 m from shore a submerged concrete wall

runs parallel with the shoreline for nearly 100 m. This wall provides an

excellent habitat for many types of aquatic life and fish are attracted to

the area.

Methods

Three methods were used to collect sauger. A weekly creel survey

of Gallipolis Locks and Dam conducted by the authors from 15 March

1981 through April 1982. Surveys resulted in 99 sauger stomachs,

donated by cooperative anglers. At various times during the study, gill

netting was conducted in the Gallipolis pool as a supplement to the creel

survey. Twelve days of netting yielded 18 sauger. Rotenone sampling

was done at Belleville Locks on 30 September 1981 in conjunction with

the Ohio River Valley Water Sanitation Commission (ORSANCO) and

the West Virginia Department of Natural Resources (WVDNR). Thirty-

four sauger were obtained,

Sauger and/ or sauger stomachs were transported on ice to the

laboratory, then frozen. Specimens were later thawed, and total length

of each fish measured to the nearest mm. Stomachs were excised and

their contents washed into a watch glass for sorting into various taxo-

nomic categories with the aid of a dissecting microscope. The number of

individuals in each taxon was recorded for each stomach. Standard tax-

onomic keys were used for identifications (list of references is available

from authors).

The following calculations were made for each monthly collection:

(1) percentage of stomachs in which a particular taxon occurred (per-

centage frequency of occurrence), (2) percentage of the total number of

food items that one taxon comprises, and (3) percentage of the total

volume (by weight) of all taxa made up by the combined assigned

weight of each particular taxon.

Foods and Feeding of Sauger 125

In order to overcome bias due to differences in stages of digestion

of food items, the original weight of individuals can be estimated

(Lagler 1956). This is done by averaging weight of 30 undigested indi-

viduals of the species. The figure obtained is an approximate "weight

when alive" value.

To determine if seasonal variations occur in the diet of the sauger,

151 specimens were arbitrarily grouped according to season: spring

(March, April, May), summer (June, July, August), and fall (Sep-

tember). The number of empty stomachs from each season also was

recorded, to determine if there was any seasonality in the percentage

frequency of their occurrence.

Specimens were divided into two total-length classes — 150 mm

and less, and 151 mm-400 mm — to determine if food varied with

length.

Our study was concurrent with a WVDNR creel survey, conducted

from 1 April through 31 November 1981.

The relationship between water temperature and sauger catch (from

the WVDNR creel survey) was determined by regression analysis and

tested with a coefficient of correlation. Temperatures used for the

regression analysis consisted of means calculated from monthly average

values from the Gallipolis Locks and Dam, and single day values from

other dams included in the investigation. Temperature data were

obtained from an electronic water quality monitoring system and a

manual sampling system, both maintained by ORSANCO.

RESULTS AND DISCUSSION

Of the 151 sauger stomachs examined, 105 (69.5%) contained food.

The number with food was highest in September (89.7%) and lowest in

July (50%) (Fig. 1). Thus, sauger appeared to feed most intensively in

the fall.

Food Items

Fishes were the primary food of sauger, both by weight (99.7%)

and by frequency of occurrence (100%) (Table 1). Emerald shiners,

Notropis atherinoides Rafinesque, and gizzard shad, Dorosoma cepedi-

anum (Lesueur), accounted for 56.6% of the total volume of food con-

sumed (Table 1). Together these two forage fish comprised 96% of all

identifiable food items (Table 1).

Emerald shiners were the most important food item by number,

occurring in 49% of stomachs that contained food, and by weight mak-

ing up 32% of food biomass (Table 1). Gizzard shad were second in

importance appearing in 11% of stomachs that contained food and

comprising 25% of the total volume of food consumed (Table 1). Other

identifiable food fishes were striped shiners, A^. chrysocephalus (Rafi-

126

Steven I. McBride and Donald Tarter

100

CO

40

Z 30

liJ

O

S 20

Q.

10

M A M J J A S

MONTHS

Fig. 1. Monthly fluctuations in percent of sauger stomachs containing food,

Ohio River at Gallipolis Locks and Dam.

Foods and Feeding of Sauger

127

Table 1. Food organisms consumed by sauger from the Ohio River at Gallipolis

Locks and Dam, March 1982-April 1982. N = total number of

organisms, % N = percentage of all organisms combined, % FO = percent

frequency of occurrence of each organism, and % Wt = percentage of

the total weight of food items.

nesque), and freshwater drum, Aplodinotus grunniens (Rafinesque).

Fishes digested beyond identification were found in 43% of the stom-

achs (Table 1).

Terrestrial and aquatic invertebrates occurred in 3% of stomachs

containing food and comprised 0.3% of the total volume of food con-

sumed (Table 1).

Forbes and Richardson (1920) examined the stomach contents of

14 sauger and concluded that this species feeds entirely on fishes. Pearse

(1921) listed the food of sauger as 75% fish, and 25% insects and plank-

ton. Dendy (1946) found food in 1,980 of 3,807 sauger stomachs col-

lected in 1943 and 1944 from Norris Reservoir. The stomach contents

consisted chiefly of shad, with crappie (Pomoxis) second in importance.

Priegel (1969) found Lake Winnebago sauger utilizing troutperch, Per-

copsis omiscomaycus (Walbaum), as their major food item. Young and

128 Steven I. McBride and Donald Tarter

yearling freshwater drum were also an important item in Priegel's study.

Vanicek (1964) examined 133 sauger collected from Lewis and Clark

Lake and the Missouri River. He found young gizzard shad to be the

most important food for both reservoir and river sauger. Although the

emerald shiner was abundant in Lewis and Clark Lake, it appeared in

only 5% of the stomachs. Swenson (1977) reported that troutperch

were the primary source of food for sauger in Lake of the Woods, Min-

nesota.

Seasonal Food Habits

Seasonal variation in sauger diet was minimal (Figs. 2 and 3).

Fishes were the dominant food item by number (100% FO) and weight

(99.7%) throughout the year (Table 1). Emerald shiners were most

important in both numerical frequency and frequency of occurrence at

all seasons. Gizzard shad ranked second in importance by number

throughout the year.

Volumetric proportions of food items generally coincided with per-

cent numbers in the diet. Emerald shiners were the most important iden-

tifiable food item volumetrically in both spring (38%) and summer

(29%). Gizzard shad comprised the largest proportion of food volume

(36%) during the fall (Fig. 3).

The greatest variation in seasonal feeding was an increase in the

number of stomachs containing food from spring (61%) and summer

(64%) to fall (89.7%).

There have been relatively few food habit studies documenting sea-

sonal variation in diet. Dendy (1946) reported that it was not uncom-

mon to find as many as 30 small shad in an individual stomach. The

shad, tending to "hibernate," are undoubtedly easy prey for sauger in

winter. Priegel (1969) reported that in the winter. Lake Winnebago

sauger consumed equal amounts of troutperch and emerald shiners.

When forage fishes were scarce, however, sauger consumed more chiro-

nomid larvae.

Food Habits by Length Class '

150 mm and less. — Our methods enabled us to collect no data

concerning food of sauger of this length class. Priegel (1969) found the

food of young sauger to vary with size, changing from zooplankton to

chironomid larvae, to immature and adult mayflies. He considered

sauger less than 50 mm to be plankton feeders; however, at 50 mm they

would cease consuming plankton if forage fishes were extremely abun-

dant. Of the invertebrates utilized by sauger in the 12-50 mm size class,

Daphnia sp. was the most important item consumed. The major forage

fishes consumed by young sauger were troutperch, freshwater drum,

and white bass, Morone chrysops (Rafinesque).

Foods and Feeding of Sauger

129

ES

GS

D

SS

Al

Tl

Spring (54)

ES

GS

D

SS

Al

Tl

Summer (58)

Fall (39)

ES

GS

D

SS

Al

Tl

0 10 20 30 40 50 60 70 80 90 100

PERCENT OCCURRENCE

Fig. 2. Seasonal frequency of occurrence of food items in sauger stomachs, Ohio

River at Gallipolis Locks and Dam. ES = emerald shiners, GS = gizzard shad, D

= freshwater drum, SS = striped shiners, Al = aquatic invertebrates, Tl = terres-

trial invertebrates.

130

Steven I. McBride and Donald Tarter

LO

-I

LO

ttC </) H--

oo

LO

« 2

c/>

Foods and Feeding of Sauger 131

151-400 mm. — Fishes were the number one food item by weight

(99.7%) and by frequency of occurrence (100%) of sauger in this length

class. Emerald shiners were the most important forage fish, with gizzard

shad second in importance. Dendy (1946) found that for sauger whose

standard lengths ranged from 200 to 480 mm, size had no influence on

the kind of food contained in the stomach. We came to a similar con-

clusion for fish in that size range. However, examination of 21 fish in

the 150-200 mm size range showed that none had consumed gizzard

shad, but all stomachs with identifiable food items contained emerald

shiners. This supports Vanicek (1964), who found that young-of-the-

year and yearling sauger in July and August consumed mostly Notropis,

despite the fact that gizzard shad were the most abundant forage fish

available.

Catch Related to Temperature

The WVDNR creel survey of the Ohio River tallied 6,499 sauger

harvested from the tailwaters of six dams on the river: New Cumberland

(RM 54.4), Pike Island (RM 84.2), Hannible Island (RM 126.4), Willow

Island (RM 161.8), Belleville (RM 203.9), and Gallipolis (RM 279.2)

dams.

Regression analysis of the relationship between sauger catch and

water temperature showed a significant negative correlation (r = -0.87)

(Fig. 4). Sauger catch was greatest in November (2,268) with the mean

water temperature at 54° F, and smallest in August (44) with the mean

water temperature at 78.8° F.

Factors other than temperature may relate to sauger catch in the

Ohio River. Doan (1941) found a high correlation (r = =0.m9) between

catch and turbidity for Lake Erie sauger.

CONCLUSIONS

Populations of emerald shiners and gizzard shad dominated the

supply of forage fish in the Ohio River during the study period. Clay

and Carter (1962) reported that the emerald shiner was by far the most

abundant species in the Ohio River, making up 57.8% of 741,438 fish

collected in a three-year period. In the same study, gizzard shad were

second in importance by number (14.5%) and most important by

weight, comprising 44.6% of 16 tons of fish. The future of sauger popu-

lations in the Ohio River is apparently dependent on the existence of

large numbers of gizzard shad and emerald shiners. Whether or not

sauger would become more opportunistic feeders in the absence of these

two forage species remains questionable. However, in all likelihood

sauger, as well as other piscivorous fishes, would suffer to some degree

if either of these food sources becomes depleted.

132

Steven I. McBride and Donald Tarter

2200-1

2000

1800H

1600

1400

5 1200

« 1000

C/3

800-

600

400

200

y=4698.63 + 58.12

r=-0.87

~T — I — I — I — I — r~

0 20 40 60

80 100

TEMPERATURE

Fig. 4. Relationship of water temperature (F) and sauger catch from tailwaters

of New Cumberland, Pike Island, Hannible, Willow Island, Belleville, and Gal-

lipolis dams, Ohio River.

Foods and Feeding of Sauger 133

If sauger catch can be interpreted as active feeding behavior, then

the negative coefficient of correlation (r= -0.87), obtained from examining

the relation of sauger catch to water temperatures, may indicate that

sauger feed most intensively during periods of cool water temperatures.

This is supported by the highest percentage of stomachs with food

(89.7%) occurring in the fall.

ACKNOWLEDGMENTS.— The authors thank Mr. Glenn Moore,

ORSANCO, for supplying the water quality data for this study. We also

express our gratitude to all the members of ORSANCO and WVDNR

who were helpful in collecting fish. Special thanks go to Bernie Dowler,

Assistant Chief of Warmwater Fisheries, WVDNR, for supplying creel

survey data.

Several persons assisted in the field work during this study. We are

particularly grateful to Curtis Hardman, George Bakewell and Mark

Sheridan. Also, we express our appreciation to Ms. Vickie Crager for

typing the manuscript.

LITERATURE CITED

Clay, William M., and B. T. Carter. 1962. Aquatic-life resources of the Ohio

River. Ohio River Valley Water Sanitation Commission, Cincinnati. 218 pp.

Dendy, Jack S. 1946. Food of several species of fish, Norris Reservoir, Ten-

nessee. J. Tenn. Acad. Sci. 27(1):105-127.

Doan, Kenneth H. 1941. Relation of sauger catch to turbidity in Lake Erie.

Ohio J. Sci. ¥/(6):449-452.

Forbes, Stephen A. 1878. The food of Illinois fishes. Bull. 111. State Lab. Nat.

Hist. l(2):71-89.

1888. On the food relations of freshwater fishes. Bull. 111. State Lab.

Nat. Hist. 2:475-538.

, and R. E. Richardson. 1920. The fishes of Illinois. 111. Nat. Hist.

Survey, Vol. 3, Ichthyology. 278 pp.

Lagler, Karl F. 1956. Freshwater Fishery Biology. W. C. Brown Co., Dubuque,

Iowa. 421 pp.

Pearse, A. S. 1921. The distribution and food of the fishes in three Wisconsin

lakes in summer. Univ. Wise. Stud. Sci. No. 3. 61 pp.

Priegel, Gordon R. 1969. The Lake Winnebago Sauger. Wis. Dep. Nat. Resour.

Tech. Bull. 43. 63 pp.

Scott, W. B., and E. J. Crossman. 1973. Freshwater Fishes of Canada. Bull.

Fish. Res. Board Can. 184. 966 pp.

Swenson, William A. 1977. Food consumption of walleye {Stizostedion vitreum

vitreum) and sauger {Stizostedion canadense) in relation to food availability

and physical conditions in Lake of the Woods, Minnesota, Shagawa Lake, and

western Lake Superior. J. Fish. Res. Board Can. i^( 10): 1643-1654.

, and L. L. J. Smith. 1973. Gastric digestion, food consumption,

feeding periodicity, and food conversion efficiency in walleye {Stizostedion

vitreum vitreum). J. Fish. Res. Board Can. i0(9): 1327- 1336.

134 Steven I. McBride and Donald Tarter

and 1976. Influence of food competition, predation.

and cannibalism on walleye {Stizostedion vitreum vitreum) and sauger {S.

canadense) populations in Lake of the Woods, Minnesota. J. Fish. Res.

Board Can. ii(9): 1946- 1954.

S. Army Corps of Engineers. 1980a. Gallipolis Locks and Dam Replacement,

Ohio River. Phase 1, Advanced Engineering and Design Study. General

Design Memorandum. Appendix J, Volume 2. 202 pp.

1980b. GalHpolis Locks and Dam Replacement, Ohio River. Phase

1, Advanced Engineering and Design Study. General Design Memorandum.

Main Report. 89 pp.

Vanicek, C. D. 1964, Age and growth of sauger, Stizostedion canadense

(Smith), in Lewis and Clark Lake. Iowa State J. Sci. 55(4):48 1-502.

Accepted 3 February 1983

Notes on Breeding Period, Incubation Period, and Egg

Masses oi Amby stoma jeffersonianum (Green) (Amphibia:

Caudata) from the Southern Limits of Its Range

Charles K. Smith ^

Department of Biology,

University of Kentucky, Lexington, Kentucky 40502

ABSTRACT. — Observations on breeding and incubation periods of

Ambystoma jeffersonianum were made at three central Kentucky

ponds during 1980 and 1981. Breeding began in early January, 1980

and in early February, 1981. There were also substantial differences

between years in duration of the breeding period. During both years

hatching occurred over a two-week period between late March and

early April. Mean number of eggs per mass was 23.4 ± 13.5, and mean

mass dimensions (length x diameter, mm) were 43.1 ± 11.9 x 39.0 ±

8.6. There are indications that the number of eggs per mass for Ken-

tucky populations is greater than for more northern populations.

INTRODUCTION

Relatively little life history data exist for Ambystoma jeffersonia-

num (Green). Data published prior to Uzzell's (1964) definitive work on

the A. jeffersonianum complex may not, in fact, deal with A. jeffersoni-

anum. The species ranges throughout most of the northeastern United

States and reaches its southern limits in central Kentucky. Life history

data from Kentucky populations, since they are peripheral, should shed

light on factors that limit the species' distribution. Unfortunately, very

few published studies concern Kentucky populations. This note provides

observations on breeding, incubation periods, and characteristics of egg

masses of A. jeffersonianum in central Kentucky. To investigate annual

and local variation in breeding and incubation periods, observations

were made in three local ponds over a two-year period.

STUDY SITES AND METHODS

Data were collected during 1980 and 1981 from three abandoned,

semi-permanent farm ponds in Jessamine County. Each pond had a sur-

face area of less than 288 square meters, and an average depth of 21 cm.

Pond 1 is located 0.4 km north of Pond 2, and Pond 3 is approximately

1.5 km northwest of Pond 1. Ambystoma jeffersonianum is the only

ambystomatid salamander that regularly uses any of the ponds, although

over a four-year period I have occasionally observed eggs and larvae of

A. texanum in Ponds 1 and 2.

• Present Address: Department of Biology, University of North Carolina, Chapel Hill,

North Carolina 27514.

Brimleyana No.9: 135-140. June 1983. 135

136 Charles K. Smith

The ponds were visited during the day once a week, and on every

rainy night beginning with the first rains of late fall (19 December 1979

and 1 November 1980). During each visit the ponds were searched for

adults and egg masses. Developmental stages of the embryos were used

to determine lengths of the breeding, incubation, and hatching periods.

Masses in early stages of development (preblastula) were assumed to be

recently deposited. In addition, data on the number of eggs per mass,

mass size (greatest length and diameter), and mass placement were col-

lected from Pond 3 on 23 March 1980 and 8 March 1981 while transects

were waded across the pond.

RESULTS AND DISCUSSION

A summary of breeding and incubation periods oi Amby stoma jef-

fersonianum is given in Table 1 . Substantial differences occurred between

years in the initiation and duration of the breeding season. Hatching,

however, occurred over a 1 to 2 week period at approximately the same

time during both years.

Since the migration oi Amby stoma to breeding ponds is correlated

with precipitation and temperature (Baldauf 1952; Whitford and Vin-

egar 1966; Douglass and Monroe 1981), variable weather conditions

Table 1. Summary of dates and duration of breeding and incubation periods for

Ambystoma jeffersonianum in three central Kentucky Ponds. '

Ambystoma jeffersonianum Reproduction 137

offer the most probable explanation for the differenes between years in

the date that breeding was initiated. Noticeably less precipitation oc-

curred at the study site between November 1980 and January 1981 than

for the same period during the previous year. Whereas the ponds filled

by January in 1980, they were not filled until February in 1981.

Differences between years in local variation in the duration of the

breeding periods, and, hence, incubation periods can best be explained

if migration to the ponds is both a function of favorable weather condi-

tions and an individual's physiological readiness to respond to those

conditions. During 1980, by the time weather conditions were favora-

ble for migration to the ponds, a relatively large group of individuals

may have been ready to migrate. Thus, a more synchronous mass

migration to the ponds resulted, and breeding and egg deposition

occurred over a shorter time period.

Synchrony of hatching is of particular interest, given the variation

that occurred in initiation and duration of the breeding season. Tem-

poral variation in oviposition was buffered and did not produce equal

variation in initiation and duration of the hatching period. Temperature

and developmental rates show positive correlation (Moore 1939; Has-

singer et al. 1970). Because ambient temperatures tend to progressively

increase during the incubation period, embryos of eggs deposited earlier

in the breeding season take longer to develop, and, as a result, the tem-

poral variation of the hatching period is much less in comparison to the

breeding period.

Worthington (1968, 1969), and Douglas and Monroe (1981) pro-

posed that temporal staggering of breeding is adaptive because it results

in a partitioning of resources among larvae of different sizes. Since the

hatching period was much shorter than the breeding period, my study

suggests that the efficacy of staggered breeding in partitioning pond

resources may be greatly reduced.

At a breeding pond approximately 135 km west of my study site,

Douglas and Monroe (1981) reported that adult A. jeffersonianum were

present from late November until early March. They did not note if

oviposition occurred over the entire period. They found a peak occur-

rence of adults during early January, as did Creusere (1972) for popula-

tions in western and central Kentucky. Seibert and Brandon (1960)

found eggs in southern Ohio as early as February, while Smith (1911)

and Bishop (1941) reported that breeding begins in late March or early

April in New York. Brandon (1961) and Creusere (1972) observed hatch-

lings in western Kentucky on dates similar to those reported here.

Bishop (1941) reported hatching to occur during April and May in New

138 Charles K. Smith

York populations. Geographic differences in initiation of breeding are

likely due to differences in temporal occurrence of weather conditions

that stimulate adult migration to the ponds. The later hatching time for

northern populations corresponds with a later breeding season.

During 1981 my study ponds filled slowly, and water levels fluctu-

ated greatly during the breeding and incubation periods. Ponds 2 and 3

did not dry until late summer, and pond drying was not a source of

embryo mortality. Pond 1, however, dried completely by 22 March, des-

troying all egg masses. Although pond 1 partially refilled before April,

no additional egg masses were found.

Egg mass data are presented in Table 2. The mean number of eggs

per mass (23.4) is similar to that reported by Brandon (1961) for popu-

lations in southern Ohio (x = 22), but greater than that reported for New

York populations by Bishop (1941; x = 16), Smith (1911; x = 14), and

Uzzell (1967; x = 14). Further study of geographic variation in the

number of eggs per mass is required in order to determine if this appar-

ent north-to-south cline is real.

Table 2. Selected data, egg masses oi Amby stoma jeffersonianum in a central

Kentucky pond (pond 3). R = range, N = sample size.

Ambystoma jeffersonianum Reproduction 139

Masses were found attached both singly and in series to blades of

herbaceous vegetation, petioles of leaves, and twigs. The descriptions of

attachment sites and mass dimensions agree with those of Uzzell (1967)

and Bishop (1941).

ACKNOWLEDGMENTS.— I thank E. K. Smith for her assistance

in the field, and J. W. Petranka for his critical review of the manuscript

and discussion of the problem. This note was taken from a larger study

of larval ecology of A. jeffersonianum which was completed for the M.

S. degree under the direction of R. W. Barbour, and was supported in

part by two Graduate Student Grants from the University of Kentucky.

LITERATURE CITED

Bishop, Sherman C. 1941. The Salamanders of New York. N. Y. State Mus.

Bull. 324. 365 pp.

Baldauf, Richard J. 1952. Climatic factors influencing the breeding migration of

the spotted salamander, Ambystoma maculatum (Shaw). Copeia 1952 (3):

178-181.

Brandon, Ronald A. 1961. A comparison of the larvae of five northeastern spe-

cies oi Ambystoma (Amphibia: Caudata). Copeia 1961 (4): 377-383.

Creusere, F. Michael. 1972. Behavioral and ecological aspects of five ambysto-

matid species. Unpubl. M. S. Thesis, Eastern Kentucky Univ., Richmond.

140 pp.

Douglas, Michael E., and B.L. Monroe Jr. 1981. A comparative study of topo-

graphical orientation in Ambystoma (Amphibia: Caudata). Copeia 1981

(2): 460-463.

Hassinger, Dawn D., J. Anderson and G. H. Dalrymple. 1970. The early life

history and ecology oi Ambystoma tigrinum and A. opacum in New Jersey.

Am. Midi. Nat. 84: 474-495.

Moore, John A. 1939. Temperature tolerance and rates of development in the

eggs of amphibia. Ecology 20: 459-478.

Smith, Bertram G. 1911. Notes on the natural history oi Ambystoma jeffersoni-

anum. A. opacum, and A. tigrinum. Bull. Wisconsin Nat. Hist. Soc. 9:

14-27.

Seibert, Henri C, and R.A. Brandon. 1960. The salamanders of southeastern

Ohio. Ohio J. Sci. 60\ 291-303.

Uzzell, Thomas M. 1964. Relations of the diploid and triploid species of

Ambystoma jeffersonianum complex (Amphibia: Caudata). Copeia 1964

(2): 257-300.

Whitford, Walter G., and A. Vinegar. 1966. Homing, survivorship, and over-

wintering of larvae oi Ambystoma maculatum. Copeia 1966 (3): 515-518.

140 Charles K. Smith

Worthington, Richard D. 1968. Observations on the relative size of three species

of salamanders in a Maryland pond. Herpetologica 24: lAl-lAd.

1969. Additional observations on sympatric species of salamander

larvae in a Maryland pond. Herpetologica 25: 227-229.

Accepted 10 March 1984

Occurrence and Habitat Preference of

Fundulus luciae (Baird) (Pisces: Cyprinodontidae) on

a Southeastern North Carolina Salt Marsh

Mark A. Shields and Carol H. Mayes

Department of Biological Sciences,

University of North Carolina at Wilmington,

Wilmington, North Carolina 28403

ABSTRACT.— Thirty-two Fundulus luciae were collected in pit traps

on a salt marsh in southeastern North Carolina during October and

November 1983. This species remained on the marsh at low tide,

exhibiting a strong preference for high marsh areas dominated by Jun-

cus roemerianus while tending to avoid low marsh habitats dominated

by Spartina alterniflora. The reported rarity of F. luciae probably

reflects inadequate sampling of its preferred high marsh habitat by

conventional collecting techniques.

INTRODUCTION

Fundulus luciae (Baird), the spotfin killifish, is found sporadically

in brackish coastal waters from Long Island, New York (Butner and

Brattstrom 1960) to Georgia (Jorgenson 1969), with most populations

apparently concentrated in the Chesapeake Bay area (Lee et al. 1980).

In North Carolina, Hildebrand (1941) reported F. luciae common only

in marsh pools on Shackleford Banks and along the Newport River,

Carteret County. More recently, Kneib (1978) found F. luciae to be

common on the high salt marsh at Tar Landing Bay on Bogue Sound,

also in Carteret County. Few records of F. luciae from other areas of

coastal North Carolina have been published. In this note we document

the occurrence and describe the habitat preference of F. luciae on a salt

marsh in southeastern North Carolina.

STUDY AREA AND METHODS

Our study site was a salt marsh bordering Bradley Creek, a small

tidal river in New Hanover County. Three marsh habitats were defined

within the study area: tall-form Spartina alterniflora (TS), short-form

S. alterniflora (SS), and Juncus roemerianus (JU). TS formed a band 1

to 3 m wide along the levees of the creek and smaller rivulets in the

marsh, while SS vegetated the flat areas behind the creekbank levees.

The higher areas of marsh farthest from the creek were dominated by JU.

One 25 X 25 X 25 cm pit trap was dug in each marsh habitat. Pits

in TS, SS, and JU were placed 1.5, 18.5, and 51.5 m, respectively, from

Bradley Creek. Each pit was lined with 1 mm mesh cloth to facilitate

removal of trapped fish. Traps were visited during daylight at low tide

Brimleyana No.9: 141-144. June 1983. 141

142 Mark A. Shields and Carol H. Mayes

when the marsh was completely drained and all fish were removed and

preserved in 10% formalin. Standard length (SL) of all specimens was

measured to the nearest mm. Specimens were deposited in the Univer-

sity of North Carolina at Wilmington (UNCW) Fish Collection.

Eleven collections were made between 19 October and 20 November

1983. Salinity of water in the pit traps ranged from 23 ppt to 33 ppt

during the study, with salinities among the three pits varying no more

than 3 ppt on any given day. Water temperature ranged from 7° C to

19° C.

The percentage of each of the three habitats in a 0.5 ha area cen-

tered over the pit traps was estimated visually from an aerial photo-

graph. TS comprised 14.5% of the available marsh habitat, SS

accounted for 60.9%, and JU comprised 24.6%. A Chi-square test,

based on the null hypothesis that fish moved at random over the marsh

at high tide and used each habitat in proportion to its availability, was

employed to measure habitat preference by F. luciae.

RESULTS AND DISCUSSION

A total of 32 F. luciae (mean SL 19.4 mm, range 5-33 mm) was

collected. The largest number trapped in one day was 12 on 11 November.

No F. luciae were caught on 14 November when mean water tempera-

ture in the pit traps was 7° C, the coldest temperature recorded during

our study. Since F. luciae becomes relatively inactive and therefore

more difficult to catch during cold weather (Byrne 1978; Kneib 1978), it

may have been even more abundant on Bradley Creek Marsh than our

limited autumn collections indicated.

Distribution of F. luciae among the three marsh habitats was not

random {x^ =8.70, df=2, P<0.025). Figure 1 shows the distinct prefer-

ence of this species for the high Juncus marsh. Preference for high

marsh habitats was also reported by Byrne (1978) and Kneib (1978).

Avoidance of strong tidal currents and predators of the low marsh and

tidal creek habitats may account for this small species' preference for

high marsh areas (Byrne 1978).

We concur with Byrne (1978) and Kneib (1978) that the purported

rarity of F. luciae is due mainly to inadequate sampling of its preferred

high marsh habitat. As demonstrated by Kneib and this study, the use

of pit traps appears to be an effective method of collecting F. luciae,

which tends to remain on the marsh at low tide. We beUeve the species

is more common and widespread than is indicated by the few published

reports based on the results of seining and trawling.

Fundulus luciae on N.C. Salt Marsh

143

Dhabitat availability

o

u

0.

Fig. 1 . Availability and use by Fundulus luciae of salt marsh habitats on Brad-

ley Creek Marsh, North Carolina, October-November, 1983. TS = tall-form

Spartina alterniflora, SS - short-form S. alterniflora, JU = Juncus roemerianus.

ACKNOWLEDGMENTS.— This study was prompted by discus-

sions with David G. Lindquist. Robin D. Bjork and Jennifer E. Slack

provided helpful field assistance. Courtney T. Hackney and Paul E.

Hosier constructively criticized the manuscript.

LITERATURE CITED

Butner, Alfred, and B. H. Brattstrom. 1960. Local movement in Meni-

dia and Fundulus. Copeia 1960 (2): 139-141.

Byrne, Donald M. 1978. Life history of the spotfin killifish, Fundulus

luciae (PiscesiCyprinodontidae), in Fox Creek Marsh, Virginia.

Estuaries 7(4):21 1-227.

Hildebrand, Samuel F. 1941. An annotated list of salt and brackish

water fishes, with a new name for menhaden, found in North

Carolina since the publication of "The fishes of North Carolina" by

Hugh M. Smith in 1907. Copeia 1941(4):220-232.

144 Mark A. Shields and Carol H. Mayes

Jorgenson, Sherrell C. 1969. A Georgia record for the cyprinodontid

fish, Fundulus luciae. Chesapeake Sci. 70(1): 65.

Kneib, R. T. 1978. Habitat, diet, reproduction and growth of the spotfin

killifish, Fundulus luciae, from a North Carolina salt marsh. Copeia

1978(1):164-168.

Lee, David S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E.

McAllister and J. R. Stauffer, Jr. 1980. Atlas of North American

Freshwater Fishes. N. C. State Mus. Nat. Hist., Raleigh. 854 pp.

Accepted 12 April 1984

145

SUBSCRIPTIONS AND EXCHANGES

The editors anticipate two issues of approximately 150 pages each annually.

Rates for subscriptions for all issues appearing within the calendar year:

Individual— United States $ 9.00

Individual — Foreign $ 1 3.00

Institution $15.00

All subscriptions must be paid in advance.

Issues will be available on an exchange basis to organizations and institu-

tions publishing general natural history and ecology journals or papers on a

fairly regular schedule. Publications received on exchange will be placed in the

State Museum's H. H. Brimley Memorial Library.

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 2761 1. Back issues are available.

DATE OF MAILING

Brimleyana No. 8 was mailed on 12 April 1983.

PUBLICATION SCHEDULE

We apologize to all subscribers and contributors who have been inconven-

ienced by our publishing delay. Brimleyana Nos. 10 and 11 are being processed

simultaneously and should appear at about the same time this autumn (No. 10 is

devoted to various aspects of the biology of the salamander, Necturus lewisi).

We are trying to assure, also, that No. 12 will appear within a reasonable pro-

duction time.

FORTHCOMING . . .

* Fishes, of Buck Creek, Kentucky;

* An update of Sympotthastia (Diptera);

* Fossil bats from Florida;

* Seasonal weight changes in raccoons;

* A new species of Paracricotopus (Diptera);

* Aspects of the biology of Etheostoma rufilineatum (Pisces);

* Aquatic distributional patterns in the Interior Low Plateaus;

* Seabird core temperatures;

* Genetic variation in Agkistrodon piscivorus (Reptilia);

* Virginia in Virginia;

* The rete in seabirds;

. . . AND OTHERS.

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CONTENTS

A New Species of Woodland Salamander of the Plethodon glutinosus

Group from the Southern Appalachian Mountains. Richard

Highton 1

Lower Wilson Creek, Caldwell County, North CaroUna: A Thermal

Refugium for Reptiles? Robert Wayne Van Devender and Paul F.

Nicoletto 21

Comparative Food Studies of Yellowfin Tuna, Thunnus albacares, and

Blackfin Tuna, Thunnus atlanticus (Pisces: Scombridae) from the

Southeastern and Gulf Coasts of the United States. Charles S.

Manooch III and Diane L. Mason 33

Benthic Macroinvertebrates of Cane Creek, North Carolina, and

Comparisions with Other Southeastern Streams. David R.

Lenat 53

Pleistocene Mammals from the Rock Springs Local Fauna, Central

Florida. Kenneth T. Wilkins 69

Bird Density and Habitat Use in Forest Openings Created by Herbi-

cides and Clearcutting in the Central Appalachians. William C.

McComb and Robert L. Rumsey 83

Drainage Records and Conservation Status Evaluations for Thirteen

Kentucky Fishes. Melvin L. Warren, Jr. and Ronald R.

Cicerello 97

Winter Food Habits of Bobcats in North Carolina. Anne M. King,

Richard A. Lancia, S. Douglas Miller, David K Woodward and

Jay D. Hair Ill

Foods and Feeding Behavior of Sauger, Stizostedion canadense (Smith)

(Pisces: Percidae), from Gallipolis Locks and Dam, Ohio River.

Steven I. McBride and Donald Tarter 123

Notes on Breeding Period, Incubation Period, and Egg Masses of

Ambystoma jeffersonianum (Green) (Amphibia: Caudata) from the

Southern Limits of its Range. Charles K Smith 135

Occurrence and Habitat Preference of Fundulus luciae (Baird) (Pisces:

Cyprinodontidae) on a Southeastern North Carolina Salt Marsh.

Mark A. Shields and Carol H. Mayes 141

Miscellany 145