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N.C. DOCUMENTS

CLEARINGHOUSE

AUG 10 1993

N.C. STATE LBRARY

RALOGH

number 18 June 1993

EDITORIAL STAFF

Richard A. Lancia, Editor

Suzanne A. Fischer, Assistant Editor

Eloise F. Potter, Production Manager

EDITORIAL BOARD

James W. Hardin Rowland M. Shelley

Professor of Botany Curator of Invertebrates

North Carolina State University North Carolina State Museum

of Natural Sciences

William M. Palmer Robert G. Wolk

Director of Research and Collections Director of Programs

North Carolina State Museum North Carolina State Museum

of Natural Sciences of Natural Sciences

Brimleyana, the Zoological Journal of the North Carolina State Museum

of Natural Sciences, appears twice yearly in consecutively numbered issues.

Subject matter focuses on systematics, evolution, zoogeography, ecology,

behavior, and paleozoology in the southeastern United States. Papers stress

the results of original empirical field studies, but synthesizing reviews and

papers of significant historical interest to southeastern zoology are also

included. Brief communications are accepted.

All manuscripts are peer reviewed by specialists in the Southeast and

elsewhere; final acceptability is determined by the Editor. Address manuscripts

and related correspondence to Editor, Brimleyana, North Carolina State

Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. Information

for contributors appears in the inside back cover.

Address correspondence pertaining to subscriptions, back issues, and

exchanges to Brimleyana Secretary, North Carolina State Museum of Natural

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

In citations please use the full name - Brimleyana.

North Carolina State Museum of Natural Sciences

Betsy Bennett, Director

North Carolina Department of Agriculture

James A. Graham, Commissioner

CODN BRIMD 7

ISSN 0193-4406

The Myriapod Types of Oscar Harger

(Arthropoda: Diplopoda, Chilopoda)

Rowland M. Shelley

North Carolina State Museum of Natural Sciences,

P.O. Box 27647, Raleigh, North Carolina 27611

ABSTRACT — The type specimens of all five milliped species —

Trichopetalum lunatum, T. glomeratum, T. iuloides, lulus furcifer,

and Polydesmus armatus — and one of the two centipedes, Lithobius

pinetorum, authored by Oscar Harger in his only paper on myri-

apods and previously thought to be lost, are housed at the Peabody

Museum of Natural History, Yale University, New Haven, Connecti-

cut. From our knowledge of the itinerary of the Yale paleontologi-

cal expedition of 1871, we know the type locality of T. glomeratum,

I. furcifer, P. armatus, and L. pinetorum, previously stated as the

"John Day River Valley, Oregon" is restricted to the vicinity of

Canyon City, Grant County, on the western slope of the Blue Mount-

ains. The female holotype confirms that T. glomeratum is a repre-

sentative of the chordeumatoid family Conotylidae, and the name is

assigned provisionally to Taiyutyla pending collection of a male

topotype. Unidentifiable female conotylids are also reported from

another area in eastern Oregon and the Snake Mountains in eastern

Nevada, which suggests that the family is widespread in montane

forests at high elevations in the generally arid Columbia Plateau

and Basin and Range Physiographic Provinces. To facilitate future

studies, I provide gonopod drawings of male syntypes for /. furcifer

and P. armatus.

One of the more obscure authors of North American myriapods

is Oscar Harger (1843-87), whose sole publication on these arthropods

(Harger 1872) described the milliped genus Trichopetalum and seven

species, two centipedes {Lithobius pinetorum and Geophilus gracilis)

and five millipeds {Trichopetalum lunatum, T. glomeratum, T. iuloides,

lulus furcifer, and Polydesmus armatus).

Born at Oxford, Connecticut, Harger attended the Connecticut

Literary Institute at Suffield and Yale College, graduating from the

latter with honors in 1868 (Schuchert and LeVene 1940). After briefly

studying zoology under Professor A. E. Verrill, Harger became the

first assistant to the vertebrate paleontologist, O. C. Marsh, partici-

pating on the latter's expeditions into the American West in 1871 and

1873. From July to September 1872, Harger dredged marine organisms

on a Coast Survey steamer with Professors Verrill and Sydney I.

Smith, Yale's first professor of Comparative Anatomy, who earlier

had been naturalist to the U.S. Lake Survey and collected the types

Brimleyana 18:1-13, June 1993 1

2 Rowland M. Shelley

of T. iuloides. Harger was studious and an active reader, and Marsh

valued his scientific opinions in paleontology. However, Marsh would

not allow Harger to publish on vertebrate fossils, either alone or

jointly with him, so Harger 's only papers are on invertebrates — that

of 1872 on myriapods, two on isopods, and one on a fossil spider.

From 1870 to 1873, Marsh led four vertebrate paleontological

expeditions of Yale students and recent graduates into the West

(Schuchert and LeVene 1940). The idea of such efforts arose from

preliminary explorations he made on a trip to the end of the trans-

continental railroad in Wyoming in August 1868 after attending a

scientific meeting in Chicago. The 1871 expedition traveled to regions

of Kansas, Wyoming, and Utah, where Harger collected 10 fossil

species. The group then rested a few days in Salt Lake City with

Brigham Young while Marsh prepared to explore a new area, the

John Day River Basin in central Oregon. After traveling 12 days by

rail and stage, the party crossed the Blue Mountains and arrived at

Canyon City, Oregon, on the John Day River on 17 October 1871,

where it waited several days for a military escort from Fort Harney,

75 mi (120 km) to the south. The group collected fossils from 31

October to 8 November in the John Day region before traveling down

the Columbia River to Portland; it then traveled to San Francisco and

returned east directly by rail or by boat via Panama.

While the expedition was in the John Day River area, Harger,

or Harger and Professor G. H. Collier, collected four Oregon myriapods

that he described in 1872 — L. pinetorum, T. glomeratum, I. furcifer,

and P. armatus. Both the publication and labels in the vials give the

locality as just the "John Day River Valley," but knowledge of the

group's activities enabled me to infer a more precise site. The John

Day River arises on the western slope of the Blue Mountains in

Grant and Umatilla counties, flows westward into Wheeler County,

then heads northward to the Columbia River forming the boundaries

between Wheeler/Wasco and Sherman/Gilliam counties. It is not to be

confused with Days Creek, Douglas County, in the Umpqua River

drainage of southwestern Oregon, the probable type locality for Zantona

douglasia Chamberlin and Bollmanella oregona Chamberlin (Shear 1974,

Gardner and Shelley 1989), which Chamberlin (1941a) misnamed as

"John Day Creek." Because most millipeds require moist leaf litter

and much of the John Day Basin is in the arid rain shadow of the

Cascade Mountains, I (Shelley 1990) speculated that the site was probably

near the confluence of the John Day and Columbia rivers in either

Sherman or Gilliam county. However, as all the myriapods were collected

in October 1871, and the expedition reached Canyon City on 17

October and only collected fossils from 31 October to 8 November

Myriapod Types of Oscar Harger 3

after waiting for the military escort, it is evident that during most of

the part of October that the group was in the John Day Valley,

it was resting in Canyon City. Consequently, there was ample time

for relaxed explorations in the vicinity of Canyon City, and I, therefore,

believe that Harger 's myriapods were collected near this town. Because

Harger's paper specifies that L. pinetorum, I. furcifer, and P. armatus

were collected by Professor Collier and himself, and T. glomeratum

was taken by Harger alone, collecting probably occurred on at least

two different dates, as one day Harger went out alone and the other

he was accompanied by Collier. There could be as few as one site

and as many as four, but further specification is not possible with

what we know now. Consequently, the type locality for all of Harger's

Oregon species is restricted to the vicinity of Canyon City, Grant

County, on the western slope of the Blue Mountains.

Harger's centipedes have received little attention since their

description. They were included in the catalog of North American

myriapods by Bollman (1893), who noted that G. gracilis Harger,

1872, was preoccupied by G. gracilis Meinert, 1870, proposed for a

European geophilomorph. Cook and Collins (1891) remarked that

Harger's description of G. gracilis conformed very closely to Schendyla

nemorensis (C. L. Koch, 1837), and the former is now regarded as a

junior synonym (Crabill 1953, 1961). Stuxberg (1875) included L.

pinetorum in his list of North American lithobiids, but he had no

personal knowledge of the species. Kevan (1983a) listed both species

as potential inhabitants of Canada, recognizing the synonymy of G.

gracilis under S. nemorensis.

In contrast to the centipeds, Harger's millipeds have been cited

in a number of publications, but the type specimens were thought to

be lost. Chamberlin and Hoffman (1958) stated that their "present

location [was] unknown" or that they were "not known to exist," and

similarly, Shear (1971, 1972) said that the holotypes of T. glomeratum

and T. iuloides were lost and that the whereabouts of that of T.

lunatum was unknown. Causey (1967) guessed right when she stated

that the holotype of T. lunatum was at the "Peabody Museum of

Natural History, Yale University, if extant," but evidently she made

no inquiries to confirm this supposition. While recently visiting the

Peabody's Museum's collection, I unexpectedly discovered these types

in the myriapod cabinet, where they have languished in obscurity for

120 years. A few vials were still capped with wax and had not been

touched for decades. The types of P. armatus were in the general

collection and not labeled as such, but those of the other millipeds

were clearly marked as types and grouped in a clamp-top jar. A

concerted search failed to reveal the types of G. gracilis, which ap-

4 Rowland M. Shelley

parently are lost, but those of L. pinetorum were in an individual

vial and clearly labeled. The sample consists of 12 nearly legless

syntypes, seven males and five females, and is number 2175; according

to the label it was collected by Harger alone, whereas the published

account states that it was collected by him and Professor Collier.

All the millipeds are listed in the continental checklist

(Chamberlin and Hoffman 1958), and detailed accounts of those Harger

assigned to Trichopetalum have recently appeared (Palmen 1952;

Shear 1971, 1982; Shelley 1988, In Press).

In the following accounts I update these reports by providing

information on the type specimens, a brief historical review of each

species, and pertinent anatomical observations. Complete synonymies

are presented, and each species is placed in its proper order and

family.

Chordeumatida: Trichopetalidae

Trichopetalum lunatum Harger

Trichopetalum lunatum Harger, 1872:3, pi. II, figs. 1-4. Ryder,

1881:527. Packard, 1883:192. McNeill, 1888:8. Cook and Collins,

1895:63-64, pi. Ill, figs. 52-54. Williams and Hefner, 1928:115,

fig. 12d. Causey, 1951:119, figs. 6-8; 1967:80, fig. 1. Palmen,

1952:8-11, figs. 10-17. Chamberlin and Hoffman, 1958:102-103.

Shear, 1972:277, figs. 497-499. Kevan, 19836:2967. Shelley,

1988:1650.

Trichopetalum album Cook and Collins, 1895:64-66, pis. II-III, figs.

22-29, 36-45. Chamberlin and Hoffman, 1958:102.

Type Specimens — Five male and nine female syntypes (nos.

2208-2209) collected by O. Harger in May 1872 at New Haven,

New Haven County, Connecticut; one male and one female syntype

(no. 2125) taken by an unknown collector on an unknown date at

Mt. Carmel, ca. 7 mi (11.2 km) north of New Haven, New Haven

County.

Remarks — Harger assigned three new species to his genus

Trichopetalum but did not specify the type species, so Cook and

Collins (1895) subsequently designated T lunatum. It is the only one

of Harger's five milliped species to retain its original combination.

The identity of T lunatum has been well established by Cook and

Collins (1895), Palmen (1952), Causey (1967), and Shear (1972); a

male syntype from New Haven that I dissected conformed to these

Myriapod Types of Oscar Harger 5

accounts. For details of the genitalia, refer to the illustrations in Palmen

(1952) and Shear (1972).

Chordeumatida: Conotylidae

Taiyutyla glomerata (Harger), new combination

Trichopetalum glomeratum Harger, 1872:118, pi. II, fig. 5. Ryder,

1881:527 Packard, 1883:192. McNeill, 1888:8. Chamberlin and

Hoffman, 1958:105. Shear, 1971:63.

Craspedosoma glomeratum: Bollman, 1893:120.

Conotyla glomerata: Cook and Collins, 1895:78. Cook, 1904:69.

Type Specimen — Female holotype (No. 2173) collected by O.

Harger in October 1871 from the vicinity of Canyon City, in the

John Day River Valley, Grant County, Oregon.

Remarks — The holotype is somewhat deformed, and its genitalia

have been dissected and are lost.

Cook and Collins (1895) stated that the original description was

too brief to allow accurate generic placement but that the segment

number, short fifth antennomere, and triangular eye patch resembled

the condition in Conotyla. Shear (1971) agreed that accurate generic

placement was impossible but perceived a similarity to Taiyutyla; he

did not think the name could be referred to either Trichopetalum or

Conotyla and considered it a nomen dubium. The holotype is about 8

mm long and has 30 post cephalic segments with obvious lateral

tergal knobs that give rise to two prominent setae, so it is clearly a

conotylid. Generic placement is impossible to determine with certainty

until a male topotype is obtained, but the milliped is smaller and its

lateral setae are much longer than those of comparative specimens of

Conotyla atrolineata (Bollman), the western-most known representative

of this genus, occurring in central British Columbia, northeastern

Washington, and northern Idaho, over 200 mi (320 km) north northeast

of Canyon City. These considerations tend to exclude Conotyla, but

the type locality is also well removed from most of the known distribu-

tions of the other northwestern conotylid genera Bollmanella and

Taiyutyla, which are from southern coastal Oregon to Mason County,

Washington, and in the Coast Ranges from San Francisco Bay to the

Columbia River, respectively (Shear 1974, 1986). However, one species

in each of these genera occurs east of the above ranges, B. bifurcata

Shear, in the Wallowa Mountains, Wallowa County, Oregon, and T

curvata Loomis and Schmitt, in Lincoln County, Montana, so either

genus could occur in the Blue Mountains, which occupy an intermediate

6 Rowland M. Shelley

geographical position between the Coast Range and both the Wallowa

Mountains and Montana. Furthermore, Canyon City is only about 110

mi (176 km) southwest of the type locality of B. bifurcata. Therefore,

I borrowed the types of both B. bifurcata and T. curvata

for direct comparisons with that of glomerata. Few setae remain on

the types of B. bifurcata, and those that do exist, on the caudal end

of the male holotype, seem shorter and are not nearly as prominent

as are those on glomerata. However, the setae on glomerata agree

closely in length and prominence with those on the holotype of T.

curvata. There is reasonable agreement in body dimensions between

glomerata and both other conotylids, but because of the similarity in

the setae, I provisionally assign glomerata to Taiyutyla, pending

collection of a male topotype. This change, which formalizes Shear's

(1971) perception of similarity to Taiyutyla, also necessitates the feminine

suffix of the specific name. Fieldwork is needed in the Blue Mountains

to collect a male conotylid to determine the identity and generic

position of glomerata and to confirm or disprove this decision.

Present evidence shows that the Conotylidae is much more wide-

spread in the West than currently known. There is a female in the

Florida State Collection of Arthropods from 12.5 mi (20 km) south

of Baker City, Baker County, Oregon, that might be conspecific

with glomerata, although this site is east of the Blue Mountains and

presumably is drier than Canyon City. I also recently received two

female conotylids that are superficially very similar to glomerata from

the Snake Mountains, White Pine County, Nevada, in the eastern part

of that state and hundreds of kilometers from any known site for the

family. These two records plus glomerata suggest that conotylids could

be scattered across the arid Columbia Plateau and Basin and Range

Physiographic Provinces, where they are undoubtedly restricted to cool-

er, forested regions at high elevations. The Ruby Mountains near Elko,

Nevada, is another plausible area for conotylids, as are ranges in the

central part of that state. Because only a few millipeds of any family

have ever been collected from the "inselberg" mountains of these

provinces, a concerted field effort is needed to both clarify the systema-

tic positions of these conotylids and document the total diplopod fauna.

Chordeumatida: Caseyidae

Underwoodia iuloides (Harger)

Trichopetalum iuliodes Harger, 1872:118. pi II, fig. 6.

Trichopetalum juloides: Ryder, 1881:527.

Trichopetalum iulioides: Packard, 1883:192.

Trichopetalum iuloides: McNeill, 1888:8.

Chordeuma iuloides: Bollman, 1893:121.

Myriapod Types of Oscar Harger 7

Underwoodia polygama Cook and Collins, 1895:80-82, pi. X,

figs. 180-190. Paleman, 1952:2-8, figs. l-9a. Chamberlin and

Hoffman, 1958:107. Kevan, 19836:2968.

Underwoodia iuloides: Cook and Collins, 1895:83-84, pi. X,

figs. 177-178. Chamberlin and Hoffman, 1958:107. Kevan,

19836:2968. Shelley, 1988:1648-1649; In Press:

Type Specimens — Eight female syntypes (No. 2207) collected by

S. I. Smith in 1871 at Simon's Harbor (misspelled as Simmon's) on

the north shore of Lake Superior, Ontario, Canada. This site is now

in Pukaskwa National Park.

Remarks — A review of Underwoodia with a description, discussion,

and illustrations of U. iuloides is in press. For details on this species,

see Shelley (1988).

Fig. 1-3. Bollmaniulus furcifer, male syntype. 1, anterior gonopods, anterior

view. 2, the same, posterior view. 3, posterior gonopods, anterior view.

Scale line = 2.2 mm for figs. 1-2, 1.6 mm for fig. 3.

Julida: Parajulidae

Bollmaniulus furcifer (Harger)

Figs. 1-3

lulus furcifer Harger, 1872:119, pi. II, fig. 7.

Parajulus furcifer: Bollman, 1887:44. Cook, 1904:70-71, pi. V, figs.

5a-e. Chamberlin, 1920:35.

8 Rowland M. Shelley

Paraiulus furcifer. Brolemann, 1895:69, pi. 7, figs. 21-23.

Bollmaniulus furcifer: Verhoeff, 1926:65. Chamberlin and Hoffman,

1958:133. Buckett, 1964:18. Kevin, 19836:2964.

Taijulus furcifer. Chamberlin, 1938:205.

Caliulus furcifer. Chamberlin, 1940:15; 1944:80.

Type Specimens — Three male and 13 female syntypes (No.

2172), most highly fragmented, collected by O. Harger and G. H.

Collier in October 1871 from the vicinity of Canyon City, in the

John Day River Valley, Grant County, Oregon.

Remarks — Bollman (1887) transferred this species into Parajulus,

misspelled as Paraiulus by Brolemann (1895), and Cook (1904) recorded

it from Corvallis, Oregon. Chamberlin (1920) reported it from Clare-

mont, Los Angeles County, California, surely a misidentification of

another, possible congeneric parajulid. Verhoeff (1926) listed furcifer

as the only component of his new genus Bollmaniulus, thereby mak-

ing it the type species by monotypy as reported by Jeekel (1971).

He did not specifically designate furcifer as the generotype, so this

status does not result from original designation, as stated by Chamber-

lin and Hoffman (1958). Chamberlin (1938, 1940) evidently was unaware

of Verhoeff s action when he transferred furcifer into his new genera

Taijulus and Caliulus, respectively, both of which have subsequently

been placed in synonymy under Bollmaniulus (Chamberlin and Hoff-

man 1958, Hoffman 1979). Chamberlin (1944) repeated the combination

C. furcifer for a form from McCloud, Siskyou County, California,

and added that the species was common over much of Oregon and

California. Buckett (1964) recognized the combination Bollmaniulus

furcifer and stated that it ranged from British Columbia into California.

As noted by Hoffman (1979, 1992), the Parajulidae is one of

the two most dominant Nearctic diplopod families in terms of com-

ponent genera and species, the other being the Xystodesmidae

(Polydesmida). It was studied from 1948 to about 1974 by Dr. Nell

B. Causey, who amassed a large collection and examined most type

specimens while conducting a detailed family revision. Unfortunately,

she never completed the project and published only a few brief papers

before her death in 1979. Consequently, knowledge of the Parajulidae

is not nearly as advanced as those of the other major Nearctic diplopod

families. Work on the taxon must essentially begin anew, a daunting

task because of the diversity of the family and the enormous amount

of preserved material in nearly every major and minor milliped repository

on the continent. The types of /. furcifer will be crucial to an investigation

of Pacific parajulids, because as the eighth oldest generic name in

the family, Bollmaniulus has priority over such other nominal Pacific

Myriapod Types of Oscar Harger

Figs. 4-5 Chonaphe armata, male syntype. 4, telopodite of left gonopod,

medial view. 5, the same, lateral view. Scale line = 1.14 mm for fig. 4,

1.0 mm for fig. 5.

genera as Saiulus, Spathiulus, Sophiulus, Codiulus, and Simiulus, all

authored by Chamberlin (1940), Tuniulus (Chamberlin 19415), and

Mulaikiulus (Chamberlin 1941a), so additional generic synonymies could

result from a study of these western forms. For the benefit of future

students, I have included drawings of the gonopods of a male syntype

(Figs. 1-3).

Polydesmida: Xystodesmidae

Chonaphe armata (Harger)

Fig. 4-5

Polydesmus armatus Harger, 1872:119-120, pi. II, fig. 8.

Leptodesmus armatus: Bollman, 1893:122. Chamberlin, 1911:264.

Chonaphe armata: Cook, 1904:56-57, pi. Ill, figs. 2a-c. Attems, 1931:65-

67, figs. 100-101; 1938:156, fig. 177. Chamberlin, 1949:125.

Chamberlin and Hoffman, 1958:27. Kevan, 19835:2968. Shelley,

1990:2314.

Type Specimens — One male and two female syntypes, all highly

fragmented, collected by O. Harger and G. H. Collier in October

10 Rowland M. Shelley

1871 from the vicinity of Canyon City, in the John Day River Valley,

Grant County Oregon. This sample was discovered in the general

milliped collection and is unnumbered.

Remarks — Harger's single gonopod illustration enabled Cook (1904)

to recognize that a male sent to him from an unknown locality in

Washington was referrable to armatus. Bollman (1893) had earlier

transferred armatus to Leptodesmus, a combination repeated by

Chamberlin (1911), but Cook (1904) assigned it to his new genus,

Chonaphe, a combination that subsequently has been recognized by

Attems (1931, 1938), Chamberlin (1949), Chamberlin and Hoffman

(1958), and Shelley (1990). Cook (1904) provided three additional

genitalia drawings, and I include here medial and lateral views of the

gonopod of a male syntype (Figs. 4—5). Five nominal species comprise

Chonaphe, but Hoffman (1979) thought these might be subspecies. I

(Shelley 1990) found few significant differences between these forms

and concluded that the genus might be monotypic with C. armata

being the oldest name. I am preparing a generic revision.

ACKNOWLEDGMENTS— \ thank C. L. Remington and R. J. Pupedis

for providing access to the Peabody Museum holdings and subsequently

loaning Harger's types. The holotype of Taiyutyla curvata, housed at

the National Museum of Natural History, Smithsonian Institution,

Washington, D.C., was loaned by J. A. Coddington; the types of

Bollmanella bifurcata, housed at the Museum of Comparative Zoology,

Harvard University, Cambridge, Massachusetts, were loaned by H. W.

Levi. The conotylid from Baker County, Oregon, was discovered in

material loaned by G. B. Edwards, Florida State Collection of Arthropods,

Gainesville. Cathy Wood typed and retyped numerous drafts of the

manuscript, and figures 1-5 were prepared by R. G. Kuhler.

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Myriapod Types of Oscar Harger 11

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Kentucky, new records and a key to the genus. Proceedings of the

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12 Rowland M. Shelley

Harger, O. 1872. New North American myriapods. American Journal of

Science and Arts 4:116-121.

Hoffman, R. L. 1979. Classification of the Diplopoda. Museum d'Histoire

Naturelle, Geneva, Switzerland.

Hoffman, R. L. 1992. On the taxonomy of the milliped genera Pseudojulus

Bollman, 1887, and Georgiulus, gen. nov., of southeastern United States

(Julida: Parajulidae). Jeffersoniana 1:1-19.

Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum:

A list of the genus and family-group names in the class Diplopoda

from the 10th edition of Linnaeus, 1758, to the end of 1957. Mono-

grafieen van de Nederlandse Entomologische Vereniging, Number 5.

Kevan, D. K. McE. 1983a. A preliminary survey of known and potentially

Canadian and Alaskan centipedes (Chilopoda). Canadian Journal of Zoo-

logy 61:2938-2955.

Kevan, D. K. McE. 1983b. A preliminary survey of known and potentially

Canadian millipedes (Diplopoda). Canadian Journal of Zoology 61:2956-

2975.

McNeill, J. 1888. A list, with brief descriptions, of all the species, including

one new to science, of Myriapoda of Franklin County, Indiana. Bulletin

of the Brookville Society of Natural History, Number 3.

Meinert, F. 1870. Myriopoda Musaei Hauniensis I. Geophili. Naturhistorisk

Tidsskrift 3:241-268.

Packard, A. S. 1883. A revision of the Lysiopetalidae, a family of Chilognath

Myriopoda, with a notice of the genus Cambala. Proceedings of the

American Philosophical Society 21:177-209.

Palmen, E. 1952. Survey of the Diplopoda of Newfoundland. Annales

Zoologici Societatis Zoologicae Botanicae Fennicae 'Vanamo' 15:1-31.

Ryder, J. A. 1881. List of the North American species of myriapods

belonging to the family of the Lysiopetalidae, with a description of a

blind form from Luray Cave, Virginia. Proceedings of the United States

National Museum 3:524-529.

Schuchert, C, and C. M. Levene. 1940. O. C. Marsh, Pioneer in Paleontology.

Yale University Press, New Haven, Connecticut.

Shear, W. A. 1971. The milliped family Conotylidae in North America,

with a description of the new family Adritylidae (Diplopoda:

Chordeumida). Bulletin of the Museum of Comparative Zoology

141:55-98.

Shear, W. A. 1972. Studies in the milliped order Chordeumida (Diplopoda):

A revision of the family Cleidogonidae and a reclassification of the

order Chordeumida in the New World. Bulletin of the Museum of

Comparative Zoology 144:151-352.

Shear, W. A. 1974. The milliped genus Bollmanella (Diplopoda, Chordeumida,

Conotylidae). Psyche 81:134-146.

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Chordeumida). Revision of the genus Taiyutyla with notes on recently

proposed taxa. American Museum Novitates Number 2600.

Myriapod Types of Oscar Harger 13

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Diplopoda). Canadian Journal of Zoology 66:1638-1663.

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Accepted 9 September 1992

14

THE SEASIDE SPARROW,

ITS BIOLOGY AND MANAGEMENT

Edited by

Thomas L. Quay, John B. Funderburg, Jr., David S. Lee,

Eloise F. Potter, and Chandler S. Robbins

The proceedings of a symposium held at Raleigh, North Carolina,

in October 1981, this book presents the keynote address of F. Eugene

Hester, Deputy Director of the U.S. Fish and Wildlife Service, a

bibliography of publications on the Seaside Sparrow, and 16 major

papers on the species. Authors include Arthur W. Cooper, Oliver L.

Austin, Jr., Herbert W. Kale, II, William Post, Harold W. Werner,

Glen E. Woolfenden, Mary Victoria McDonald, Jon S. Greenlaw,

Michael F. Delany, James A. Mosher, Thomas L. Merriam, James A.

Kushlan, Oron L. Bass, Jr., Dale L. Taylor, Thomas A. Webber, and

George F. Gee. A full-color frontispiece by John Henry Dick illustrates

the nine races of the Seaside Sparrow, and a recording prepared by

J. W. Hardy supplements two papers on vocalizations.

"The Seaside Sparrow, with its extensive but exceedingly narrow

breeding range in the coastal salt marshes, is a fascinating species. All

the authors emphasize that the salt marsh habitat is at peril. . . . The

collection is well worth reading." — George A. Hall, Wilson Bulletin.

1983 174 pages Softbound ISBN 0-917134-05-2

Price: $15 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: SEASIDE SPARROW, N.C. State Museum of Natural Sciences,

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

A Late Pleistocene Vertebrate Assemblage

from the St. Marks River, Wakulla County, Florida

Timothy S. Young1 and Joshua Laerm

The University of Georgia Museum of Natural History and

Department of Zoology,

University of Georgia, Athens, Georgia 30602

ABSTRACT— The St. Marks River in the central panhandle of Florida

contains a well known, apparently late Pleistocene vertebrate as-

semblage that has been only superficially examined and reported.

Previous collections are reviewed, and we report on new fossil

materials recently obtained. Included are 37 species of mammals,

26 birds, 13 reptiles, 2 amphibians, and 9 fish. Of these, 14

species of mammals and 2 reptiles are limited solely to the Pleis-

tocene. The fauna is mixed and reflects heterochronous deposition

over time beginning at least in the late Pleistocene (Wisconsinan)

and extending through the Recent. The species present reflect mixed

woodland and grassland terrestrial communities as well as mixed

estuarine and freshwater aquatic communities. The St. Marks River

assemblage compares well to other contemporaneous late Pleisto-

cene Florida panhandle sites. One extralimital taxa is reported,

Pylodictic cf. P. olivaris, the flathead catfish, whose natural range

has not been reported east of the Mobile Bay drainage basin.

Florida is characterized by a number of rich and well

documented Pleistocene vertebrate assemblages (Webb 1974a,

Lundelius et al. 1983, Webb and Wilkins 1984) that contain a mixture

of extant and extinct South American immigrant and North

American endemic species. The majority of these sites are

distributed throughout the peninsular portion of the State (Webb

1974a, Webb and Wilkins 1984). However, with the exceptions of

Wakulla Springs (Brodkorb 1963, Webb 1974a), Chipola River

(Martin 1969, Webb 1974a), and Aucilla River (Olson 1972, Webb

1914a, Gillette 1976a) very little attention has been devoted to sites

on the panhandle of Florida. The St. Marks River, located 32 km

south of Tallahassee in Wakulla County, is a particularly rich, late

Pleistocene panhandle site that has only been superficially

investigated (Gillette 19766, Steadman 1980).

Leidy (1870), who reported on the occurrence of Mammuthus

columbi (now M. jeffersonii), provided the first record of vertebrate

fossil remains recovered from the St. Marks River, though the exact

locality was not given. Subsequently, the St. Marks River has attracted

'Present address: Department of Zoology, University of Florida, Gainesville, FL 32611.

Brimleyana 18:15-57, June 1993 15

16 Timothy S. Young and Joshua Laerm

numerous amateur collectors but a limited number of professionals.

Storrs Olson (National Museum of Natural History) collected from a

broad, shallow water area in Wakulla County near the Leon County

line several times between 1968 and 1970. Tall Timbers Research

Station sponsored collecting parties in the same area during 1974.

Published accounts of the fauna are those of Gillette (1976b), who

reported the mammals, and Steadman (1980), who discussed two

specimens of Meleagris gallopavo. Storrs Olson (personal communication)

examined the avian assemblage from the previous collections; however,

he did not publish his findings.

With the exception of Mammuthus sp. Mammut americanum,

Synaptomys australis, and possibly Equus, the majority of species from

Gillette's (1916b) report are extant. Gillette (19766) suggested that

the assemblage was important because it represented a restricted

temporal interval of the latest Pleistocene through the Holocene. Olsen

(personal communication) felt the avian assemblage was very similar

to that of today. Steadman (1980) characterized the site as a late

Pleistocene deposit. The St. Marks River has also been reported by

Lundelius et al. (1983) as being a naturally accumulating, fluvial

Rancholabrean deposit.

The purpose of our study is to review previous collections

and to report on new fossil materials recently obtained from the

St. Marks River. We provide information regarding the paleoenvironment

of the depositional area and compare the St. Marks River fauna to

other late Pleistocene faunas in the region.

GEOLOGICAL AND GEOGRAPHIC SETTING

Florida consists of five naturally occurring topographical

divisions (Cooke 1939:14). The St. Marks River drainage basin is in

the coastal lowlands division. Although the panhandle of Florida

shows a topographical record of the relict shorelines, no ages have

been securely assigned to these formations (Winker and Howard

1977a, 6). The coast line of the panhandle during the late

Pleistocene is reported to be similar to that of today (Winker and

Howard 19776).

The St. Marks River is considered part of the Gulf Hammock

region; it is underlain by the Upper Oligocene Suwannee Limestone

(Harper 1914:302). The early Miocene St. Marks Formation overlies

the Suwannee Formation in almost all of Wakulla County (Puri and

Vernon 1964). The St. Marks Formation was revised to include the

calcareous downdip facies of the Tampa Formation (Puri 1953). These

formations can be found in many areas as outcroppings in springs

and rivers (Spencer and Rupert 1987). The surface is mostly loamy

St. Marks River Fauna

17

sand, probably Pleistocene in origin. The soil surrounding the river's

edge is classified as Tooles-Nutall fine sand that is frequently flooded

(Spencer and Rupert 1987). Topographically the region is nearly level,

except for a few hilly areas (Harper 1914:302). The whole area east

of the Apalachicola River in Wakulla County is called the Woodville

Karst Plain (Hendry and Sproul 1966, Yon 1966), characterized by

sand dunes overlying limestone (Hendry and Sproul 1966:154).

The St. Marks River is fed by the St. Marks Spring located

just inside Leon County (Fig. 1). Limestone lines the perimeter of

the spring. The vent is located about 26 m below the water surface

and has an average base flow of 14.7 m3/sec (Rosenau et al. 1977).

This measurement was taken approximately 800 m down stream

Fig. 1. St. Marks River in the central panhandle of Florida. Solid circles

indicate 1987 collection sites.

18 Timothy S. Young and Joshua Laerm

from the main vent. The pH and temperature as measured 16 July

1974 were 7.6 and 21. OC, respectively. Newport Spring also feeds

the river about 800 m north of the U.S. Highway 98 bridge. The

discharge of the spring as measured 2 March 1972 was 0.23 m3/sec

with a pH of 7.8 and water temperature of 19C (Rosenau et al.

1977).

Primary depositional site(s) were not located. The fossils are

probably eroding out of the banks along much of the length of the

river and washed down river by the current. Dense accumulations of

fossils may be found in sand deposits, around submerged debris, and

in deep holes along the entire length of the river.

The St. Marks River with its shallow, relatively clear waters

with abundant fossil and archaeological materials has been a popular

recreational S.C.U.B.A. diving are for the past 30 years. Local divers

report huge quantities of fossils have been recovered by amateur collectors.

One of us (J.L.) observed an entire pick-up truck load of fossils

being removed in 1978. Local divers report that have collected "tons

of it." Although several large private collections of St. Marks material

exist, unfortunately they have been mixed with fossils from other

regional aquatic systems which makes their inclusion here

inappropriate.

METHODS

We made extensive new collections and reviewed previously

collected materials housed at the Florida Museum of Natural History,

University of Florida (UF), and the National Museum of Natural

History (USNM), Washington, D. C. Our collections are housed at

the University of Georgia Museum of Natural History (UGAMNH).

Collection efforts were concentrated in six separate locations

approximately 3.2 km in either direction from the U.S. Highway 98

bridge that crosses the St. Marks River (Fig. 1). The fossils were

collected from 16 to 19 July 1987 by a team of six people from

the University of Georgia using S.C.U.B.A. gear. The majority of

the fossil materials was collected by hand from these locations along

the river. In addition, extensive sand samples were taken at each

site for subsequent screening.

To preliminarily identify recovered materials we used the

Comparative Reference Skeletal Collection of the Zooarchaeology

Laboratory, the University of Georgia Museum of Natural History.

Reference sources were also used in preliminary identifications.

All materials were subsequently taken to the Paleontology

Laboratory, the Florida Museum of Natural History, University of

Florida, to confirm identifications. Notes were made on the

element identified, side, and fusion of bones where possible.

St. Marks River Fauna 19

SYSTEMATIC PALEONTOLOGY

Standardized common and current scientific names follow Robins

et al. (1991) for fishes; Collins (1990) for amphibians and reptiles;

American Ornithologists' Union (1983) for birds, and Kurten and Anderson

(1980) and Jones et al. (1992) for mammals. Museum acronyms are

indicated in the introduction. A complete faunal listing of the species

recovered from the St. Marks site is provided in Table 1.

CLASS MAMMALIA

Order Didelphimorphia

Family Didelphidae

Didelphis virginiana Kerr

Virginia Opossum

Material— A single left dentary, UGAMNH1735.

Remarks — The single element is identical to that of modern

Didelphis virginiana. This was the only marsupial species present in

North America during the Pleistocene. It is known from numerous

fossil sites in Florida (Webb 1974a). Its stratigraphic range includes

Middle Blancan to Recent (Kurten and Anderson 1980). It occurs in

a variety of habitats, but it is usually found in forests and woodlands

near water (Gardner 1973). We follow Marshall et al. (1990) in the

use of the ordinal name Didelphimorphia as do Jones et al. (1992).

Order Xenarthra

Family Dasypodidae

Holmesina septentrionalis (Leidy)

Northern Pampathere

Material — Right astragulus, UGAMNH2012; right calcaneus,

UGAMNH2159; right metacarpus II, UGAMNH1981; two phalanges,

UGAMNH1982-1983; numerous dermal plates, UGAMNH1954-1980, 1984-

2029, 2160, 2166.

Remarks — The species is known from numerous sites throughout

the South and Southeast. Its range is somewhat similar to that of

its modern relative, Dasypus novemcinctus, and Holmsina had a similar

preference for open woodlands (Kurten and Anderson 1980). Like its

modern counterpart, Holmsina probably fed on insects and various

invertebrates. Kurten and Anderson (1980) suggest this diet might

have restricted them to relatively warm climates where food was

available year round. Specimen UHAMNH2159, a right calcaneus, has

rodent and carnivore gnaw marks that occurred prior to fossilization.

Its stratigraphic range is early Irvingtonian to Wisconsinan (Kurten

and Anderson 1980).

20

Timothy S. Young and Joshua Laerm

Table 1. List of vertebrate species recovered from the St. Marks River.

The figure f indicates extinct forms.

Class Mammalia

Order Didelphimorphia

Family Didelphidae

Didelphis virginiana

Order Xenarthra

Family Dasypodidae

\Holmesina septentrionalis

Family Megalonychidae

^Megalonyx jeffersonii

Family Mylodontidae

^Glossotherium harlani

Order Primates

Family Hominidae

Homo sapiens

Order Lagomorpha

Family Leporidae

Sylvilagus sp.

Order Rodentia

Family Castoridae

Castor canadensis

Family Geomyidae

Geomys pinetis

Family Muridae

Microtus sp.

Microtus pinetorum

Neo fiber alleni

Ondatra zibethicus

Synaptomys australis

Order Carnivora

Family Mustelidae

M us tela sp.

Lutra canadensis

Mephitis mephitis

Family Canidae

Canis sp.

"\Canis dirus

Urocyon cinereoargenteus

Family Procyonidae

Procyon lotor

Family Ursidae

Ursus cf. U. americanus

Family Felidae

Felis sp.

^Smilodon sp.

Order Proboscidea

Family Mammutidae

■\Mammuthus jeffersonii

Family Elephantidae

\Mammut americanum

Order Perissodactyla

Family Equidae

Equus sp.

Family Tapiridae

~\Tapirus sp.

Order Artiodactyla

Family Tayassuidae

^Platygonus compressus

Family Suidae

Sus scrofa

Family Camelidae

"\Hemiauchenia macrocephala

^Palaeolama mirifica

Family Cervidae

Odocoileus virginianus

Family Bovidae

Bison sp.

Bison bison

Bos taurus

Class Aves

Order Podicipediformes

Family Podicipedidae

Podiceps auritus

Podilymbus podiceps

Order Pelecaniformes

Family Phalacrocoracidae

Phalacrocorax auritus

Order Ciconiformes

Family Ardeidae

Ardea herodias

Butorides virescens

Egretta caerulea

Family Threskiornithidae

Eudociums albus

Order Anseriformes

Family Anatidae

Aix sponsa

Anas acuta

St. Marks River Fauna

21

Table 1. Continued.

Anas americana

Anas discors

Anas platyrhynchos

Anas sp.

Ay thy a collar is

Ay thy a sp.

Branta canadensis

Bucephala albeola

Lophodytes cucullatus

Mergus merganser

Order Falconiformes

Family Accipitridae

Buteo jamaicensis

Pandion haliaetus

Order Galliformes

Family Phasianidae

Meleagris gallopavo

Order Gruiiformes

Family Rallidae

Fulica americana

Gall inula chloropus

Family Aramidae

Aramus guarauna

Order Strigiformes

Family Strigidae

Strix varia

Class Reptilia

Order Testudines

Family Chelydridae

Chelydra serpentina

Family Kinosternidae

Gen. et spec, indet.

Family Emydidae

Pseudemys concinna

Pseudemys floridanus

Pseudemys nelsoni

Trachemys scripta

Terrapene Carolina

Terrapene Carolina putnami

Family Testudinidae

Geochelone incisa

Geochelone sp.

Gopherus polyphemus

Family Trionychidae

Trionyx sp.

Order Squamata

Family Colubridae

Elaphe obsoleta

Gen et spec, indet.

Order Crocodilia

Family Alligatoridae

Alligator mississippiensis

Class Amphibia

Order Caudata

Family Sirenidae

Siren sp.

Order Anura

Gen. et spec, indet.

Class Osteichthyii

Order Lepisosteiformes

Family Lepisosteidae

Lepisosteus sp.

Order Amiiformes

Family Amiidae

Amia calva

Order Siluriformes

Family Ictaluridae

Py Iodic tis cf. P. olivaris

Family Ariidae

Ariopsis felis

Order Salmoniformes

Esocidae

Esox sp.

Order Perciformes

Family Percichthyidae

Morone saxatilis

Family Sparidae

Archosargus probatocephalus

Family Sciaenidae

Sciaenops ocellatus

Family Mugilidae

Mugil sp.

22 Timothy S. Young and Joshua Laerm

Family Megalonychidae

Megalonyx jeffersonii (Desmarest)

Jefferson's Ground Sloth

Material — A single phalanx, UGAMNH2135, and tooth,

UGAMNH2136.

Remarks — Jefferson's ground sloth occurred in woodlands where

it apparently fed on nuts, berries, leaves, and twigs (Stock 1925). It

is known from a number of sites in the Southeast including Florida

(Webb 1974a), Georgia (Ray 1967), South Carolina (Hay 1923, Roth

and Laerm 1980), and Tennessee (Guilday et al. 1969). It could have

tolerated a seasonally cool climate as evidenced by its

Pleistocene occurrence in what is now Canada and Alaska (McNab

1985). It is reported from Irvingtonian to Rancholabrean sites with a

terminal date of 13,890 years B.P., although Kurten and Anderson

(1980) suggest it may have survived even longer in Florida.

Family Mylodontidae

Glossotherium harlani (Owen)

Harlan's Ground Sloth

Material— Two teeth, UGAMNH2137-2138.

Remarks — This was an open plains and grassland species (Stock

1925). It is reported from Irvingtonian to Rancholabrean sites with a

terminal date of 13,890 years B.P., although Kurten and Anderson

(1980) suggest it may have survived even longer in Florida.

Order Primates

Family Hominidae

Homo sapiens Linnaeus

Human

Material — Cranial fragment, UF21280.

Remarks — This single specimen was recovered by Gillette (19766).

Unfortunately, the cranial fragment was not available for examination.

We are, therefore, unable to comment on the degree of mineralization.

No other human remains were recovered in our efforts. The presence

of considerable amounts of Native American cultural material (pottery

shards) as well as 18-20th century European-American artifacts

indicates the St. Marks River was a site of human occupation before

and after European contact.

St. Marks River Fauna 23

Order Lagomorpha

Family Leporidae

Sylvilagus sp. indet.

Material — Tooth fragment, UF21301.

Remarks — Rabbits are a common component of most Pleistocene

sites in Florida. It is surprising no more than a single tooth fragment

was encountered in the St. Marks River material. Two species of

rabbit occur in the St. Marks region today, the eastern cottontail,

Sylvilangus floridanus, and the more common swamp rabbit, S.

aquaticus. The former prefers heavy brushy, forested areas with open

areas nearby and edges of swamps. The latter is most common in

marshes, swamps, and bottomlands (Golley 1962).

Order Rodentia

Family Castoridae

Castor canadensis Kuhl

Beaver

Material — Left ulna, UGAMNH2126; right upper molar,

UGAMNH2125; right M3, UGAMNH2124; four molars, UF21294.

Remarks — Two beaver species occurred in Florida in the late

Pleistocene, Castoroides ohioensis and Castor canadensis. Both have

even been found in the same deposits (Webb 1974a); however, only

the latter is represented in the St. Marks River fauna. The beaver is

found in any suitable water habitat including rivers, streams,

lakes, and marshes (Lowery 1974). Its relative rarity in the St. Marks

may be related to the presence of Alligator. The stratigraphic range

is late Blancan to Recent (Kurten and Anderson 1980).

Family Geomyidae

Geomys pinetis Rafinesque

Southeastern Pocket Gopher

Material — A single lower fourth premolar, UF21291.

Remarks — Geomys pinetis is the only species of pocket gopher

in the Southeast. It is associated with the sandy soils of the Coastal

Plain (Golley 1962) and is present today in the uplands adjacent the

St. Marks River. It is known from late Irvingtonian to Recent

(Kurten and Anderson 1980).

Family Muridae

Microtus sp. indet.

Material— Left M3, UGAMNH2127.

Remarks — This fragment, while certainly Microtus, could not be

24 Timothy S. Young and Joshua Laerm

referred to a species with confidence. We follow Jones et al. (1992)

in their use of the familial name Muridae.

Microtus pinetorum (LeConte)

Pine Vole

Material— Right M2, UGAMNH2128.

Remarks — This molar compares well to modern Microtus

pinetorum. Regionally, the pine vole can be found in a wide range

of habitats from hardwood and pine forests to overgrown fields

(Golley 1962). The stratigraphic range is Sangamonian to Recent (Kurten

and Anderson 1980).

Neofiber alleni True

Round-tailed Muskrat

Material— right M2, UGAMNH2121; right M3, UGAMNH2123;

right M3, UGAMNH2122; maxilla, UF21293.

Remarks — Neofiber alleni is a semi-aquatic mammal that prefers

permanent bodies of water with emergent aquatic vegetation (Frazier

1977). Although it has a restricted range today, essentially extreme

northern Florida and south Georgia, during the Pleistocene it ranged

as far west as Kansas (Hibbard 1943). It is reported from late Irvingtonian

to Recent (Kurten and Anderson 1980). The stratigraphic range is

Illinoian to Recent (Kurten and Anderson 1980).

Ondatra zibethicus (Linnaeus)

Muskrat

Material— Right dentary with M1 AND M2, UGAMNH2120;

dentary, UF21292.

Remarks — The muskrat, like the round-tailed muskrat, is a semi-

aquatic mammal that prefers permanent bodies of water (Nelson and

Semken 1970). There is not overlap in the range of the two species

today. However, Martin and Webb (1974) indicate they were sympatric

in at least two late Pleistocene Florida faunas, Devils Den and

Ichetucknee River. The occurrence of the two species in the St. Marks

River fauna is not overly suggestive that they were sympatric here in

the past because of the apparently heterochronous deposition at St.

Marks. Furthermore, although the muskrat does not presently occur in

the St. Marks River or Apalachicola River drainages, it is known

from the extreme western panhandle and the Upper Coastal Plain of

Georgia, a distance of 120 km.

St. Marks River Fauna 25

Synaptomys australis Simpson

Florida Bog Lemming

Material— Left mandible with Mb UF21295.

Remarks — The specimen referred to in Gillette's (1976b) review

of the St. Marks River is the only record of this species at the site.

In Florida it is known primarily from Sangamonian and

Wisconsinan assemblages, although elsewhere it is known from the

Illinoian through the Wisconsinan (Kurten and Anderson 1980). Its

presence at Devils Den suggests it might have persisted until about

8,000 years B.P. (Martin and Webb 1974), although this radiocarbon

date is considered suspect. The Florida bog lemming is similar

morphologically to S. cooperi, the northern bog lemming, but differs

considerably in size; it is about 35% larger than S. cooperi. Kurten

and Anderson (1980) suggest it might represent a clinal variate of S.

cooperi. It was an inhabitant of moist bogs and damp meadows (Burt

1928).

Order Carnivora

cf. Order Carnivora, gen. et sp. indet.

Material— A left coronoid, UGAMNH1881.

Remarks — This specimen, though carnivore-like, could not be identified

to the familial level.

Family Mustelidae

Mustela sp. indet.

cf. Weasel

Material— A single left humerus, UGAMNH1738 and right P3,

UGAMNH1736.

Remarks — Two species of weasel, Mustela frenata and M. vison,

are common to the region today. Both are known from the

Irvingtonian to Recent and are represented in regional fossil sites

(Webb 1974a). However, fossil weasels have been reported from very

few sites in Florida (Martin 1974, Webb 1974a).

Lutra canadensis (Shreber)

River Otter

Material— Left humerus, UGAMNH1741.

Remarks — This material compares well to modern Lutra

canadensis. The stratigraphic range includes early Irvingtonian to Recent,

and the species is represented in numerous regional sites (Kurten

and Anderson 1980). The species occurs in woodlands near rivers

and streams but is also known from tidal creeks and marshlands

(Lowery 1974).

26 Timothy S. Young and Joshua Laerm

Mephitis mephitis (Schreber)

Striped Skunk

Material — Right mandible, UGAMNH1746; left humerus,

UGAMNH1737.

Remarks — This material compares well to modern Mephitis

mephitis, which can be found in mixed woodlands, brushlands, or

prairies but generally in reasonable proximity to water (Lowery 1974).

The stratigraphic range is mid Blancan to Recent (Kurten and

Anderson 1980).

Family Canidae

Canis sp. indet.

Material— Left ilium, UGAMNH1739; right dentary,

UGAMNH1878, 1880; right scapula, UGAMNH1879.

Remarks — None of these elements could be identified beyond

the generic level. They are well mineralized, suggesting they are not

modern C. familiaris contaminants. Several species of Canis are known

from late Pleistocene sites in Florida. These include C. lupus, the

gray wolf; C. rufus, the red wolf; C. latrans, the coyote; and C.

dims, the dire wolf. Martin (1974) has concluded that only two species,

C. rufus and C. dirus, are common to middle and late Pleistocene

deposits of Florida. Canis lupis is typical of Irvingtonian deposits,

whereas C. dirus is representative of the Rancholabrean.

Canis dirus Leidy

Dire Wolf

Material— left radius, UGAMNH1877.

Remarks — Canis dirus is known from a number of late Pleistocene

sites in Florida (Webb 1914a) and is one of the more common species

of mammals at numerous Rancholabrean sites throughout North America.

It is thought to have inhabited a wide range of habitats because it

was a hunter and scavenger (Kurten and Anderson 1980, Lundelius et

al. 1983). The stratigraphic range is early Illinoian to Wisconsinan

(Kurten and Anderson 1980). The most recent terminal date for extinction

is given at about 8,000 years B.P. in Florida (Martin and Webb

1974), but somewhat earlier (approximately 9,000-10,000 year B.P.)

elsewhere (Kurten and Anderson 1980).

Urocyon cinereoargenteus (Shreber)

Gray Fox

Material— Right dentary, UGAMNH1743; left frontal,

UGAMNH1744.

St. Marks River Fauna 27

Remarks — This material is not well mineralized, which suggests

that it is a modern contaminant. However, Urocyon cinereoargenteus

would be expected in this fauna. It can be found in a wide range of

habitats today, but brushy and woody areas probably best describe

the preferred habitat in the South and Gulf Coast area (Lowery 1974).

The stratigraphic range in Florida is Middle Rancholabrean to Recent

(Martin and Webb 1974). Elsewhere it is known as early as the

Blancan (Kurten and Anderson 1980).

Family Procyonidae

Procyon lotor (Linnaeus)

Racoon

Material— Three left dentaries, UGAMNH1742, 1747, 1750. A

partial skeleton is represented by UF21296.

Remarks — The University of Georgia material is not well

mineralized, which suggests it could be a modern contaminant,

since Procyon lotor is part of the modern fauna. In the Florida panhandle

today, the racoon is an inhabitant of forested bottomland swamps. It

fossil record in Florida extends from the Late Irvingtonian to Recent

(Martin and Webb 1974).

Family Ursidae

Ursidae gen. et sp. indet.

Material— Three phalanges, UGAMNH1745, 1749, 1752.

Remarks — Generic identity of this material is uncertain. In addition

to the modern black bear, Ursus americanus Pallas, several extinct

species of bears are known from the Pleistocene of Florida. These

include the cave bear, Tremarctos floridanus (Gidley), and the lesser

short-faced bear, Arctodus pristinus Leidy, all of which persisted at

least until the late Wisconsin (Kurten and Anderson 1980).

Ursus cf. U. americanus Pallas

cf. Black Bear

Material — A single right dentary with M1? UGAMNH1751.

Remarks — This specimen is well mineralized, but it is too worn

for positive identification. The black bear can be found in forests

and bottomland swamps throughout much of the Southeast (Golley

1962, Lowery 1974). It is represented in numerous late Pleistocene

sites. The stratigraphic range is early Irvingtonian to Recent (Kurten

and Anderson 1980).

28 Timothy S. Young and Joshua Laerm

Family Felidae

Felis sp. indet.

Material— Left radius, UGAMNH1740.

Remarks — This specimen is a large Felis, but it is too worn for

positive identification. Webb (1974a) states that several species of

Felis are known from the Late Pleistocene of Florida, and include F.

atrox Leidy, F. concolor Linnaeus, F. onca (Linnaeus), F. pardalis

Linnaeus, F. rufus Schreber, and F. yagouaroundi Geoff roy. Another

possibility is Felis amnicola, a new, small cat described by Gillette

(1976a). The description is based on several specimens from various

localities in Florida and possibly Georgia.

Smilodon sp. indet.

Material— Left scapho-lunar, UGAMNH1748.

Remarks — The sabertooth cats are reported from a dozen or more

late Pleistocene sites in Florida (Webb 1974a, Kurten and

Anderson 1980). Slaughter (1963) proposed a series of successional

changes in Smilodon species throughout the North American

Pleistocene. Webb (1974a) concurs that records of Smilodon in Florida

support such a successional outline: Smilodon gracilis is a late Blancan

and early Irvingtonian; S. fatalis is representative of late Irvingtonian

and early Rancholabrean sites; and that S. floridanus is typical of the

late Rancholabrean. The temporal span reflected by other faunal elements

from the St. Marks would be more suggestive of the latter species;

however, given the similarity of these species, more precise identification

is impossible from the limited available material. Smilodon could probably

have been found in habitats ranging from grassland to woodland (Merriam

and Stock 1932, Lundelius et al. 1983).

Order Proboscidea

Proboscidea gen. et sp. indet.

Material— Sesamoid, UGAMNH1098; tusk fragment, UF21255; skull

fragment, UF21256; leg fragment, UF21257; vertebral fragment UF21258.

Remarks — These specimens are very definitely proboscidean, but

assignment to species is impossible.

Family Mammutidae

Mammut americanum (Kerr)

American Mastodon

Material — Axis fragment, UGAMNH1614; tooth fragments,

UGAMNH1612, 1613, 1615, 1616, UF21267 and 21276; tusk fragments,

UF21277-21278; proximal humerus, UF21279; calcaneus, UF21290.

St. Marks River Fauna 29

Remarks — The morphology of the elements is consistent with its

identification as Mammut americanum. Dreimanis (1968) suggested that

M. americanum inhabited coniferous forests. The stratigraphic range is

early Blancan to Wisconsinan (Kurten and Anderson 1980).

Family Elephantidae

Mammuthus jeffersonii (Osborn)

Jefferson's Mammoth

Material — Tooth fragments, UGAMNH1607-1611; tooth fragments,

UF21259-21262, 21264-21266; mandibular symphysis, UF21263.

Remarks — The morphology of the tooth fragments and

mandibular symphysis is consistent with its identification as

Mammuthus jeffersonii. Jefferson's mammoth probably inhabited open

grasslands (Stock 1963, Harrington et al. 1974). The stratigraphic range

is Illinoian to Wisconsinan (Kurten and Anderson 1980).

Order Perissodactyla

Family Equidae

Equus sp. indet.

Horse

Material — left astragalus, UGAMNH1035; cervical vertebra,

UGAMNH1170; left upper cheek tooth, UGAMNH1045; right upper

cheek tooth, UGAMNH1031; right lower cheek tooth, UGAMNH1048;

cheek tooth, UGAMNH1046, 1047, 1062; right deciduous P2,

UGAMNH1042; left deciduous P2, UGAMNH1061; right cuneiform,

UGAMNH1049; left femoral head, UGAMNH1034; right distal

humeral epiphysis, UGAMNH1054; left I3, UGAMNH1036; right

I2, UGAMNH1041; lower incisor, UGAMNH1056; left I3, UGAMNH1059;

left I1, UGAMNH1060; incisive fragment, UGAMNH1032; left upper

molar, UGAMNH1038; right M2,UGAMNH1039; right M3,

UGAMNH1044; left M3, UGAMNH1057; upper molar fragment

UGAMNH1029; left navicular, UGAMNH1063; medial phalanges

UGAMNH1030, 1050, 1053; distal phalanx, UGAMNH1051; proximal

phalanx, UGAMNH1055; left P2, UGAMNH1037; right P2,

UGAMNH1040; left upper premolar, UGAMNH1043; right lower

premolar, UGAMNH1058; left scapula, UGAMNH1052; sesamoid,

UGAMNH1033; medial phalanx, UF21228; teeth, UF21229-21238; teeth

UF21240-UF21254; cheek tooth, axis, and pelvis, UF21297.

Remarks — Equus is well represented in St. Marks River. A portion

of the material is poorly mineralized and probably represent

contaminants of the modern E. caballus. However, the majority of

elements are well fossilized, and it is likely that most of the material

30 Timothy S. Young and Joshua Laerm

is of late Pleistocene origin. Given the uncertain relationships of late

Pleistocene horses in general and the likelihood of heterochronous

deposition, we did not assign the material to a particular species.

Pleistocene Equus was generally a grassland species (Kurten and Anderson

1980).

Family Tapiridae

Tapiridae gen. et spec, indet.

Material— Left dentary, UGAMNH2068.

Remarks — This edentulous specimen could not be assigned to

Tapirus with confidence, although the morphology is similar.

Tapir us sp. indet.

Tapir

Material — Right upper deciduous premolar, UGAMNH2070, left

upper deciduous premolar, UGAMNH2071; right fibula, UGAMNH2069.

Remarks — The available material, while certainly Tapirus, could

not be referred to a species with confidence. Tapirs occur in wet

woodlands (Simpson 1945, Gray and Crammer 1961).

Order Artiodactyla

Family Tayassuidae

Platygonus compressus LeConte

Flat-headed Peccary

Material— Axis, UGAMNH2072.

Remarks — The material has the diagnostic characters of Platygonus

compressus which is thought to have wide environmental tolerances,

but was probably most associated with open woodlands (Martin and

Guilday 1967, Ray et al. 1970). The stratigraphic range is

Sangamonian to Wisconsinan (Kurten and Anderson 1980).

Family Suidae

Sus scrofa Linnaeus

Pig

Material— Left maxilla with P3 and P4, UGAMNH1159; right

humeral fragment, UGAMNH1160; right femoral diaphysis,

UGAMNH1162; right radial fragment, UGAMNH1161; left humeral

fragments, UGAMNH1163, 1158; right ilial fragment, UGAMNH1157;

distal humeral fragment, UGAMNH1178; left femur, UGAMNH1156.

Remarks — None of the pig material showed evidence of significant

mineralization. The pig was introduced during historic times and

represents a domesticate. Specimen UGAMNH1178 shows marks of a

saw.

St. Marks River Fauna 31

Family Camelidae

Camelidae gen. et sp. indet.

Material— Proximal phalanges, UGAMNH1647, 1648; right

scaphoid, UGAMNH1649; right astragalus, UGAMNH1650; right

scapula, UGAMNH1651; left proximal femoral fragment, UGAMNH1652.

Remarks — These specimens have distinctive camelid familial

characters, but cannot be assigned to a particular species.

Hemiauchenia macrocephala (Cope)

Large-headed Llama

Material— Proximal phalanges, UGAMNH2151, 2152.

Remarks — The identification of these elements to Hemiauchenia

macrocephala is based on the size of the phalanges. According to

Webb (1974/?), H. macrocephala was a plains and grasslands inhabitant.

The stratigraphic range is Wisconsinan to Recent (Kurten and Anderson

1980). Because this species has such a limited stratigraphic range, at

least a portion of the deposit can be correlated to the Wisconsinan.

Paleolama mirifica (Simpson)

Stout-legged Llama

Material — Left proximal metacarpal fragment, UGAMNH2146; right

humerus, UGAMNH2145; left metatarsus, UGAMNH2158; left M3, UGAMNH2147;

metapodial, UGAMNH2148; right distal humerus, UGAMNH2149; left

pisiform, UGAMNH2150.

Remarks — The stratigraphic range is late Irvingtonian to

Wisconsinan (Kurten and Anderson 1980). Webb (1974b) reports them

to be an inhabitant of grasslands and savannahs. Specimen

UGAMNH2158, a left metatarsus, has longitudinal cracks indicative

of weathering prior to fossilization.

Family Cervidae

Odocoileus virginianus (Zimmerman)

White-tailed Deer

Material— Antler fragments, UGAMNH1103, 1106, 1134, 1148;

left astragalus, UGAMNH1101, 1124, 1210, 1757; right astragalus,

UGAMNH1071, 1113, 1125, 1677, 1758, 2162; right calcaneus,

UGAMNH1150, 1181, 1204, 1756; left calcaneus, UGAMNH1069, 1073,

1077, 1079, 1205, 1653, 2153, 2154, 1667, 1678; right

cubonavicular, UGAMNH1074, 1111; right dentary with P1? P2, P3,

M2, M3, UGAMNH1081; left dentary with Mlt UGAMNH1131; right

dentary with Mh M2, M3, UGAMNH1108, 1126; right dentary with

P2, P3, M1? M2, M3, UGAMNH1191; right proximal femoral fragment,

UGAMNH1184; femoral diaphysis, UGAMNH1064; left femoral head,

32 Timothy S. Young and Joshua Laerm

UGAMNH1129; left femoral diaphysis, UGAMNH2163; right distal

femur, UGAMNH11139; left distal femoral fragment, UGAMNH1085,

1088; left femoral lesser trochanter, UGAMNH1130; right femoral

diaphysis, UGAMNH1139, 2164; right frontal with antler,

UGAMNH1068, 1099; left frontal with antler, UGAMNH1012, 1082,

1118, 1121, 1666; frontal with antler pedicle, UGAMNH1196, 1203;

right humeral fragments, UGAMNH1075, 1100, 1122, 1133, 1142, 1661,

1668; left humeral fragments, UGAMNH1070, 1093, 1136, 1137, 1143,

1185, 1186, 1235, 1679; right ilial fragments, UGAMNH1076, 1090,

1681; left ilial fragments, UGAMNH1079, 1086, 1147; left ischial

fragments, UGAMNH1105, 1119, 1682; right lunate, UGAMNH1112;

right maxilla with P4, M1, UGAMNH1206; left metacarpal fragments,

UGAMNH1087, 1102, 1114, 1115, 1146, 1180, 1198, 1663; right metacarpal

fragments, UGAMNH1072, 1084, 1092, 1097, 1116, 1665; metacarpal

diaphysial fragments, UGAMNH1104, 1193-1195, 1670; right metatarsal

fragments, UGAMNH1091, 1183, 1201, 1659, 1669, 1675, 1759; metatarsal

diaphysial fragments, UGAMNH1117, 1199, 1200, 1208, 1212, 1656,

1672, 1673, 1676;

left metatarsal fragments, UGAMNH1120, 1192, 1654, 1655, 1662,

1664, 1671; right M1, UGAMNH1109, UGAMNH1151; left M3,

UGAMNH1289; right M3, UGAMNH1190; left petrous, UGAMNH1110,

1128, 1202, 1214; medial phalanx, UGAMNH1080, 1141, 1209, 1213,

1753, 1754; proximal phalanx, UGAMNH1078, 1109, 1127, 1182,

UGAMNH1211, 1755; left radial fragment, UGAMNH1065; right radial

diaphysial fragment, UGAMNH1067; left radial fragments, UGAMNH1207,

2165; right radial fragments, UGAMNH1144; sacrum, UGAMNH1089;

left scapular fragments, UGAMNH1140, 1188; right scapular fragments,

UGAMNH1094, 1145; right distal tibial fragments, UGAMNH1095,

1123, 1658, 1680; left distal tibial fragments, UGAMNH1096, 1657;

left proximal tibial fragment, UGAMNH1674; right proximal ulnar fragment,

UGAMNH1132; left proximal ulnar fragment, UGAMNH1197; thoracic

vertebral fragment, UGAMNH1760; lumbar vertebral fragment,

UGAMNH1187; cervical vertebral fragment, UGAMNH1066; atlar

fragments, UGAMNH1083, 1149; axial fragment, UGAMNH1660; antler,

UF21289; five mandibles, UF21298.

Remarks — The deer material shows a considerable range of

mineralization. A significant portion is poorly mineralized and probably

represents modern contaminants. The remaining material, however, is

well mineralized, but mineralization alone is a poor indicator of possible

Pleistocene age. The stratigraphic range of species is middle Blancan

to Recent (Kurten and Anderson 1980). Odocoileus is a woodland

and forest edge species (Golley 1962, Lowery 1974, Lundelius et al.

1983).

St. Marks River Fauna 33

Family Bovidae

Bovidae gen. et sp. indet.

Material— Proximal phalanges, UGAMNH1621, 1627, 1631,

1632; left lunate, UGAMNH1633, 1634; left lunar, UGAMNH1625;

left scapual spine, UGAMNH1635; rib head, UGAMNH1636; right

scapula, UGAMNH1622; right P4, UGAMNH1623; left distal humerus,

UGAMNH1624; left proximal femur, UGAMNH1626; right distal humeral

epiphysis, UGAMNH1628; metatarsal diaphysial fragment,

UGAMNH1629; left proximal tibial fragment, UGAMNH1630; tooth

fragments, UF21239, 21281, 21282, 21285, 21288; distal humerus, UF21283;

horn core tip, UF21284.

Remarks — These elements are definitely bovid but the available

material does not permit specific distinction.

Bison bison (Linnaeus)

Bison

Material— -Right M2, UGAMNH1620; left P3, UGAMNH1619; left

P2, UGAMNH1618; right M2, UGAMNH1617; molar UF2299.

Remarks — While Jones et al. (1992) have employed Bos bison

for the American bison, we continue the traditional use of Bison

bison. Two species of bison are known from Florida. The giant

bison, B. latifrons, is known from Illinoian and Sangamonian and

survived up until the late Wisconsinan. The American buffalo or

bison, B. bison, was widespread throughout the Wisconsinan through

the Recent (Kurten and Anderson 1980). Bison is typically associated

with grasslands, though in the Southeast may well have ranged into

woodlands (Golley 1962, Stock 1963). It became extinct in the

southeastern United States early in the 19th Century.

Bos taurus Linnaeus

Cow

Material — Right proximal humeral diaphysis, UGAMNH 10863; left

scapula, UGAMNH1155; left ilium, UGAMNH1152, 1154; right metatarsal

diaphysis, UGAMNH117; right proximal humerus,

UGAMNH1177; right distal humerus, UGAMNH1176; orbital portion

of right maxilla, UGAMNH1 153; right proximal tibia,

UGAMNH1171; right distal femoral epiphysis, UGAMNH1172; right

astragalus, UGAMNH1173; distal phalanx, UGAMNH1174; metapodial,

UF21300.

Remarks — Bos taurus was introduced into North America sometime

after 1492. All elements were poorly mineralized. The presence of

cow indicates the site has modern contaminants.

34 Timothy S. Young and Joshua Laerm

CLASS AVES

Order Podicipediformes

Family Podicipedidae

Podiceps auritus (Linnaeus)

Horned Grebe

Material— Distal portion of left ulna, USNM209968.

Remarks — Today the species winters in coastal areas and

infrequently occurs in freshwater (Sprunt 1954).

Podilymbus podiceps (Linnaeus)

Pied-billed Grebe

Material— Right humerus, USNM210293, 210294, 210301, 210302;

humerus, USNM210311; right proximal humerus, USNM210304; left

humerus, USNM210307; left tibial fragments, USNM210292, 210308,

210309, 210315, 210322, 210325, 210327; right tibia, USNM210297,

210300, 210312, 210316, 210317; left ulna, USNM210296, 210303,

210305, 210306, 210310, 210328, 210329; right ulna USNM210313,

210321; radius, USNM210298; right coracoid, USNM210319; left

coracoid, USNM210320; left femur, USNM210298; right

carpometacarpus, USNM210321; left carpometacarpus, USNM210323;

scapula, USNM210324; pedal phalanx, USNM210326; tarsometatarsus,

USNM210314, 210318; left tibia, USNM210306.

Remarks — The species inhabits freshwater marshes and ponds,

but also is associated with saltwater in winter (Sprunt 1954).

Order Pelecaniformes

Family Phalacrocoracidae

Phalacrocorax auritus (Lesson)

Double-crested Cormorant

Material— Left radius, USNM209845; left ulna, USNM209844;

scapula, USNM209859; anterior sternum, USNM209843; left coracoid,

USNM209858; phalanx 1 of digit II, USNM209861, 209852; phalanx

2 of digit II, USNM209856; left humerus, USNM209846; right

humerus, USNM209851; proximal radius, USNM209860; right ulna,

USNM209852; distal tibia, USNM209848; right femur, USNM209854;

right mandible, USNM209855; left mandible, USNM209850; sternal

fragment, USNM209857; left coracoid, USNM209849; right coracoid,

USNM209847.

Remarks — This species is distributed in large rivers and lakes as

well as brackish and saltwater systems (Sprunt 1954).

St. Marks River Fauna 35

Order Ciconiformes

Family Ardeidae

Ardea herodias Linnaeus

Great Blue Heron

Material— Cervical vertebrae, USNM210282, 210283, 210285,

210287; right mandible, USNM210281; mandible fragments,

USNM210280, 210286, 210288; maxilla fragment, USNM210279; right

coracoid, USNM210291; right proximal humerus, USNM210289; distal

tarsometatarsus, USNM210290; right carpometacarpus, USNM210279.

Remarks — The great blue heron has wide ecological tolerances,

occurring in freshwater swamps and riparian habitats as well as

saltwater marshes (Sprunt 1954).

Butorides striatus (Linnaeus)

Green-backed Heron

Material— Right humerus, USNM209966.

Remarks — Butorides striatus and B. virescens, sometimes regarded

as separate species, are recognized as geographic races of B. striatus

by the American Ornithologists Union (1983). It occurs along lake

margins, streams, ponds, and freshwater and saltwater marshes (Sprunt

1954).

Egretta caerulea (Linnaeus)

Little Blue Heron

Material — Mandibular tip with right ramus, USNM209862.

Remarks — Freshwater swamps and saltwater marshes are the

preferred habitats (Sprunt 1954).

Family Threskiornithidae

Eudocimus albus (Linnaeus)

White Ibis

Material — Right humerus, USNM209971; left proximal coracoid,

USNM209972.

Remarks — Eudocibus albus is associated with swampy forests,

marshy sloughs, and saltwater marshes (Sprunt 1954).

Order Anseriformes

Family Anatidae

Aix sponsa (Linnaeus)

Wood Duck

Material— Right carpometacarpus, USNM209931, 209938, 209939,

209944; left carpometacarpus, USNM209934; right ulna,

36 Timothy S. Young and Joshua Laerm

USNM209927, 209946; left ulna, USNM209926, 209940, 209945; left

humerus, USNM209928, 209930, 209932, 209933; right humerus,

USNM209941; radius, USNM209929, 209936; scapula, USNM209942,

209943; proximal tibia, USNM209935; right coracoid, USNM20992;

right femur, USNM209925; right tarsometatarsus, USNM209937.

Remarks — The species is common today in freshwater woodland

rivers, ponds, and marshes (Sprunt 1954).

Anas sp. indet.

Material— Right ulna, UGAMNH2078.

Anas acuta Linnaeus

North Pintail

Material— Left coracoid, USNM209965.

Remarks — The pintail is associated with freshwater marshes,

ponds, and lakes (Sprunt 1954).

Anas americana Gmelin

American Wigeon

Material— Left humerus, USNM210270; left ulna, USNM210267,

210273; right scapula, USNM210278; scapula USNM210272, 210274;

right coracoid, USNM210268, 210275, 210276; right ulna, USNM210277;

phalanx 1 of digit II, USNM210269; radius, USNM210271.

Remarks — This species is an inhabitant of freshwater marshes,

ponds, and shallow lakes (Sprunt 1954).

Anas discors Linnaeus

Blue-winged Teal

Material— Left carpometacarpus, USNM209865, 209866, 209872-

209874; right carpometacarpus, USNM209870, 209871; right humerus,

USNM209869; right coracoid, USNM209868; left ulna, USNM209864.

Remarks — Sprunt (1954) reports the species from freshwater ponds

and lakes.

Anas platyrhynchos Linnaeus

Mallard

Material— Left humerus, USNM209910, 209911, 209914; right

humerus, USNM209913, 209918; right scapula, USNM209916; left scapula,

USNM209917; left coracoid, USNM209919; right coracoid, USNM209912;

right carpometacarpus, USNM209920; furcula,

USNM209915.

St. Marks River Fauna 37

Remarks — The mallard prefers freshwater lakes and marshes

(Sprunt 1954).

Aythya sp. indet.

Material — Right carpometacarpus, UGAMNH2073; left distal

tibiotarsus, UGAMNH2077.

Aythya collaris (Donovan)

Ring-necked Duck

Material— Humerus shaft, USNM209899; left humerus,

USNM209884, 209878, 209886, 209890, 209894, USNM209910; right

humerus, USNM209877, 209893, 209903; left ulna, 209885, 209888,

209897; right ulna, USNM209878, 209892, 209905-209907; left tibia,

USNM209880, 209908; right tibia, USNM209909; left

carpometacarpus, 209898, 209900; right tarsometatarsus,

USNM209881, 209889; left tarsometatarsus, USNM209887;

tarsometatarsus, USNM209902; right coracoid, 209895, 209896; proximal

radius, USNM209882; distal radius, USNM209883; radius, USNM209904;

right scapula, USNM209891; cervical vertebra, USNM209901.

Remarks — This species is associated most commonly with wooded

lakes, ponds, and rivers, but also is reported from saltwater systems

(Sprunt 1954).

Branta canadensis (Linnaeus)

Canada Goose

Material — Right coracoid, UGAMNH2074; right tarsometatarsus,

USNM209875; right distal carpometacarpus, USNM209876.

Remarks — Both USNM specimens from the 1970s are noted by

Storrs Olson (personal communication) as small and possibly represent

either a small subspecies or juveniles. The UGAMNH specimen from

1987 is large. Sprunt (1954:53) states the center of abundance in

Florida for modern Branta canadensis is the St. Marks Refuge. This

coracoid could possibly be assigned to Branta cf. B. dickeyi on the

basis of size. Steven Emslie (Point Reyes Bird Observatory, personal

communication) examined the St. Marks River specimen and thought

it could be assigned to B. dickeyi. Measurements of the coracoid are

larger than modern B. canadensis, but there is some overlap. Emslie

(personal communication) reported a large B. dickeyi from the early

Pleistocene of Florida. We refer the coracoid conservatively to B.

canadensis. The species prefers freshwater lakes, rivers, and marshes

(Sprunt 1954).

38 Timothy S. Young and Joshua Laerm

Bucephala albeola (Linnaeus)

Bufflehead

Material — Right carpometacarpus, USNM209969.

Remarks — The bufflehead is most common in saltwater bays and

estuaries, and rarely in freshwater lakes and ponds (Sprunt 1954).

Lophodytes cucullatus (Linnaeus)

Hooded Merganser

Material — Right proximal humerus USNM209975; right distal

humerus USNM209976, 209980; right humerus, USNM209977; left humerus,

USNM209978; left ulna, USNM209979.

Remarks — The species occurs in freshwater wooded ponds,

rivers, and lakes (Sprunt 1954).

Mergus merganser Linnaeus

Common Merganser

Material— Left distal tarsometatarsus, USNM209863.

Remarks — The common merganser inhabits wooded freshwater rivers

and ponds but winters in saltwater bays (Sprunt 1954).

Order Falconiformes

Family Accipitridae

Pandion haliaetus (Linnaeus)

Osprey

Material — Right distal tarsometatarsus, USNM209967.

Remarks — The species prefers fresh and saltwater marshes, lakes,

and bays (Sprunt 1954).

Buteo jamaicensis (Gmelin)

Red-tailed Hawk

Material— Left distal humerus, USNM209970.

Remarks — The red-tailed hawk is most common in deciduous

forests adjacent to open grasslands (Sprunt 1954).

Order Galliformes

Family Phasianidae

Meleagris gallopavo Linnaeus

Wild Turkey

Material — Left tarsometatarsus, UGAMNH2075; right proximal tibiotarsus,

USNM209921; right proximal femur, USNM209922;

tarsometatarsus shaft, USNM209923.

Remarks — The species is known from drier swamps, open pine,

and hardwoods as well as prairies (Sprunt 1954).

St. Marks River Fauna 39

Order Gruiformes

Family Rallidae

Fulica americana (Gmelin)

American Coot

Material— Left distal tarsometatarsus, UGAMNH2076; left tibia,

USNM209947, 209951; left distal tibiotarsus, USNM209956, 209958;

right distal tibiotarsus, USNM20949, 209952, 209962; right tibiotarsus,

USNM209961; tibiotarsus shaft, USNM209955; left ulna, USNM209954,

209966; right carpometacarpus, USNM209950, 209963; right distal femur,

USNM209948; distal humerus, USNM209953; right scapula, USNM209959;

left coracoid, USNM209960.

Remarks — The American coot is primarily associated with open

freshwater ponds and marshes (Sprunt 1954).

Gallinula chloropus (Linnaeus)

Common Moorhen

Material— Right tarsometatarsus, USNM210259, 210260, 210255;

right tibiotarsus, USNM210257; radius, USNM210256; left phalanx 1

of digit II, USNM210258.

Remarks — This species prefers freshwater marshes and ponds with

heavy aquatic vegetation (Sprunt 1954).

Family Aramidae

Aramas guarauna (Linnaeus)

Limpkin

Material— Left tarsometatarsus, USNM210262, 210266; right

tarsometatarsus, USNM210261; right distal tarsometatarsus,

USNM210265; left distal tibiotarsus, USNM210263; right distal

tibiotarsus, USNM210264.

Remarks — The limpkin is associated with open, freshwater

swamps and marshes (Sprunt 1954).

Order Strigiformes

Family Strigidae

Strix varia Barton

Barred Owl

Material — Right proximal femur, USNM209973; right tibiotarsus

shaft, USNM209974.

Remarks — The barred owl occurs in low, wet woodlands and

swampy forests (Sprunt 1954).

40 Timothy S. Young and Joshua Laerm

CLASS REPTILIA

Order Testudines

Family Kinosternidae

Kinosternidae gen. et sp. indet.

Material— Nuchal, UGAMNH2038, 2047; right peripheral 1,

UGAMNH2041; left peripheral 2, UGAMNH2042; left peripheral 4,

UGAMNH2052; right peripheral 4, UGAMNH2044; left peripheral 9,

UGAMNH2053; right peripheral 10, UGAMNH2048; plastron fragment,

UGAMNH2050, 2051; right pleural 1, UGAMNH2039; left pleural 1,

UGAMNH2045; right pleural 2, UGAMNH2043; right pleural 6,

UGAMNH2040; pleural fragments, UGAMNH2046, 2049.

Remarks — None of the kinosternid material could be referred to

genus or species.

Family Chelydridae

Chelydra serpentina (Linnaeus)

Snapping Turtle

Material— Right peripheral, UGAMNH2034, 2037; left peripheral

4, UGAMNH2035; peripheral UGAMNH2036.

Remarks — This material compares well with modern Chelydra

serpentina. The species prefers permanent freshwater systems (Conant

1975).

Family Emydidae

Emydidae gen. et spec, indet.

Material— Right epiplastron, UGAMNH1350, 1351, 1355, 1356,

1366, 1368, 1370, 1402, 1403, 1405, 1444, 1462, 1482, 1510, 1530,

1535, 1538, 1546, 1548-1550, 1554, 1872; left epiplastron,

UGAMNH1235, 1268, 1282, 1285, 1307, 1343, 1345, 1346, 1349,

1359, 1362, 1375, 1380, 1390, 1394, 1445, 1511, 1551; left humerus,

UGAMNH1498; right hypoplastron at inguinal notch,

UGAMNH1217, 1237, 1241, 1242, 1281, 1308, 1316, 1322, 1324,

1325, 1357, 1379, 1382, 1418, 1419, 1467, 1469, 1501, 1601, 1871;

right hypoplastron at axillary notch, UGAMNH1238, 1240, 1301,

1352, 1358, 1404, 1452, 1474, 1503, 1547, 1553, 1555, 1565; right

hypoplastron, UGAMNH1164, 1167, 1215, 1216, 1219, 1221, 1236,

1283, 1333, 1344, 1354, 1376, 1388, 1389, 1429, 1433, 1470, 1473,

1480, 1495, 1521, 1533, 1572, 1575, 1584, 1600, 1867, 1870; left

hypoplastron at axial notch, UGAMNH1290, 1361, 1369, 1566,

1602; left hypoplastron at inguinal notch, UGAMNH1168, 1220,

1269, 1280, 1284, 1300, 1302, 1413, 1447, 1449, 1516, 1522, 1527,

1559, 1567, 1582; left hypoplastron, UGAMNH1218, 1222-1225, 1239,

1278, 1279, 1293, 1320, 1342, 1377, 1396, 1423, 1456, 1464, 1471,

St. Marks River Fauna 41

1475, 1509, 1513, 1519, 1526, 1532, 1560, 1564, 1569, 1571, 1574,

1578, 1583; neural 1, UGAMNH1328, 1439, 1545, 1558; neural 2,

UGAMNH1568, UGAMNH1588; neural 3, UGAMNH1573, 1581; neural

6, UGAMNH1461, 1577; neural 7, UGAMNH1260, 1341, 1579, 1874;

neural 8, UGAMNH1271; neural 9, UGAMNH1410; neural, UGAMNH1233,

1258, 1259, 1277, 1291, 1309, 1310, 1312, 1363, 1364, 1372, 1384,

1392, 1393, 1406, 1409, 1414, 1440, 1441, 1494, 1524, 1563, 1570,

1576, 1580, 1589, 2139, 2141; nuchal, UGAMNH 1261- 1264, 1313,

1411, 1417, 1427, 1457, 1486, 1504, 1508, 1518, 1595; right periphal

1, UGAMNH1165, 1321, 1399, 1398, 1451, 1489, 1525, 1528, 1592,

1866; left peripheral 1, UGAMNH1231, 1245, 1303, 1319, 1454, 1505,

1562; right peripheral 2, UGAMNH1381, 1397, 1407; left peripheral

2, UGAMNH1275, 1298, 1400; right peripheral 3, UGAMNH1169,

1294, 1442, 1531; left peripheral 3, UGAMNH1416, 1421, 1472; right

peripheral 4, UGAMNH1540; left peripheral 4, UGAMNH2167; right

peripheral 5, UGAMNH1395; left peripheral 5, UGAMNH1425, 1591;

right peripheral 6, UGAMNH1244, 1517; left peripheral 6, UGAMNH1274,

1296, 1446;

right peripheral 7, UGAMNH1246, 1428, 1594; left peripheral

7, UGAMNH1329, 1432, 1453, 1455; right peripheral 8, UGAMNH1232,

1552, 2143; left peripheral 8, 1424, 1542; right peripheral 9, UGAMNH1166,

1373, 1484, 1490; left peripheral 9, UGAMNH1249, 1299, 1492;

right peripheral 10, H1273, 1276; left peripheral 10, UGAMNH1248;

right peripheral 11, UGAMNH1326, 1332, 1429, 1587, 1597; left

peripheral 11, UGAMNH1297, 1304, 1408, 1435; peripheral

UGAMNH1000, 1243, 1247, 1311, 1420, 1426, 1434, 1442, 1449,

1450, 1536, 1593, 1604; right pleural 1, UGAMNH1234, 1334, 1336,

1374, 1385, 1437, 1485, 1493, 1502, 1554, 1875; left pleural 1,

UGAMNH1365, 1371, 1378, 1391, 1438, 1468, 1487, 1507, 2142;

left pleural 2, UGAMNH1292; right pleural 2, UGAMNH1465,

UGAMNH1491; left pleural 3, UGAMNH1431; right pleural 3,

UGAMNH1436, UGAMNH1430; left pleural 4, UGAMNH1460; right

pleural 5, UGAMNH1596; left pleural 5, UGAMNH1492; right

pleural 6, UGAMNH1488; right pleural 7, UGAMNH1340; Pleural,

UGAMNH1265-1267, 1270, 1286-1289, 1295, 1305, 1306, 1314, 1318,

1330, 1339, 1348, 1352, 1367, 1383, 1387, 1401, 1483, 1537, 1543,

1585, 1586, 1590, 1598, 1599, 1603, 1868, 1873; pygal, UGAMNH1317,

1323, 1422, 1458, 1476, 1556, 1561; left scapula, UGAMNH1496,

1497; right scapula, UGAMNH1665; suprapygal, UGAMNH1499;

right xiphiplastron, UGAMNH 1226- 1230, 1250, 1251, 1253, 1255,

1257, 1331, 1347, 1360, 1415, 1463, 1472, 1479, 1481, 1869, 1876;

left xiphiplastron, UGAMNH1252, 1254, 1256, 1315, 1335, 1337, 1338,

1386, 1512, 1514, 1515, 1523, 1539, 1544, 1459, 1478.

42 Timothy S. Young and Joshua Laerm

Remarks — Most of the emydid material could only be identified

to the familial level. Species level identification is difficult and requires

nearly complete elements. Almost all the material was well mineralized.

We are confident that the majority represents Pleistocene deposition

as opposed to Recent.

Pseudemys concinna (LeConte)

River Cooter

Material— Left peripheral 3, UGAMNH1882; left peripheral 4,

UGAMNH1885; right peripheral 7, UGAMNH1884; right peripheral

11, UGAMNH1883.

Remarks — Pseudemys concinna is distinguished by its distinctive

carapace. It is most common in slow streams and rivers (Conant

1975).

Pseudemys floridana (LeConte)

Cooter

Material— Left peripheral 3, UGAMNH2030; left pleural 3,

UGAMNH2031; left pleural 4, UGAMNH2033; nuchal, UGAMNH2032.

Remarks — The species is most commonly associated with

permanent bodies of freshwater including swamps and rivers (Conant

1975).

Pseudemys nelsoni Carr

Florida Redbelly Turtle

Material — Entoplastron, UGAMNH1904; right epiplastron,

UGAMNH1920; right hypoplastron axial notch, UGAMNH1889, 1913,

1938; left hypoplastron axial notch UGAMNH1928, 1940; right

hypoplastron inguinal notch, UGAMNH1908, 1943; left hypoplastron

inguinal notch, UGAMNH1897, 1905; neural 7, UGAMNH1901;

neural, UGAMNH1887; nuchal, UGAMNH1914, 1953; right peripheral

1, UGAMNH1899, 1906; right peripheral 2, UGAMNH1929; right

peripheral 3, UGAMNH1937; left peripheral 3, UGAMNH1917; right

peripheral 4, UGAMNH1930; left peripheral 5, UGAMNH1900; left

peripheral 7, UGAMNH1890, 1898; right peripheral 8, UGAMNH1942,

1950; left peripheral 8, UGAMNH1945, 1948; right peripheral 9,

UGAMNH1915; left peripheral 9, UGAMNH1886; left peripheral 10,

UGAMNH1946, 1947; right peripheral 11, UGAMNH1506, 1918, 1919,

1941; left peripheral 11, UGAMNH1944; peripheral, UGAMNH1907,

1909, 1949; right pleural 1, UGAMNH1534, 1892, 1910; left pleural

1, UGAMNH1922, 1951, 2140; left pleural 2, UGAMNH1917; left

pleural 3, UGAMNH1912; left pleural 4, UGAMNH1933; right pleural

5, UGAMNH1934; pleural, UGAMNH1891, 1893-1895, 1903, 1916,

St. Marks River Fauna 43

1921, 1923-1927, 1931, UGAMNH1932, 1935, 1936, 1939; suprapygal,

UGAMNH1888; right xiphiplastron, UGAMNH1896; left xiphiplastron

UGAMNH1902, 1952.

Remarks — This is a species associated with freshwater sloughs,

marshes, streams, and ponds (Conant 1975).

Trachemys scripta (Schoepff)

Slider

Material— Entoplastron, UGAMNH1763, 1792, 1801, 1831; right

hypoplastron axial notch, UGAMNH1780; left hypoplastron axial

notch, UGAMNH1828, 1833; right hypoplastron inguinal notch,

UGAMNH1774; left hypoplastron inguinal notch, UGAMNH1819;

neural 1, UGAMNH1790, 1834; neural 3, UGAMNH1789; neural 8,

UGAMNH1846; neural, UGAMNH1766, 1769, 1819, 1837, 1842; nuchal,

UGAMNH1520, 1764, 1773, 1776, 1778, 1782, 1784, 1787, 1791,

1823, 1841, 1844; right peripheral 1, UGAMNH1768, 1770, 1826,

1827; left peripheral 1, UGAMNH1783, 1840; right peripheral 2,

UGAMNH1765, 1798, 1802; left peripheral 2, UGAMNH1776, 1806;

right peripheral 3, UGAMNH1794; left peripheral 3, UGAMNH1796;

left peripheral 5, UGAMNH1775; left peripheral 8, UGAMNH1835,

1836, right peripheral 9, UGAMNH1779, 1793; left peripheral 10,

UGAMNH1767, 1820, 1839; right peripheral 11, UGAMNH1781, 1843;

left peripheral 11, UGAMNH1762, 1803-1805, 1816, 1825, 1832, 1845;

peripheral, UGAMNH1785, 1824, 1829; left pleural 1, UGAMNH1807;

right pleural 2, UGAMNH1799; left pleural 2, UGAMNH1800; left

pleural 4, UGAMNH1788; pleural, UGAMNH1771, 1772, 1786, 1808-

1815, 1821, 1822, 1838; pygal, UGAMNH1795, 1797, 1818, 1830.

Remarks — This material has the distinctive sculpted appearance

of Pleistocene Trachemys scripta. All the material is well mineralized.

It occurs in freshwater ponds, streams, and rivers (Conant 1975).

Terrapene Carolina (Linnaeus)

Eastern Box Turtle

Material— Right and left epiplastron, UGAMNH1703; left

hypoplastron, UGAMNH2144; left and right hypoplastron and

xiphiplastron, UGAMNH1697, 1698; right hypoplastron at hinge,

UGAMNH1686, 1727; left hypoplastron at hinge, UGAMN1685, 1687,

1690, 1705, 1731; right hypoplastron at inguinal notch, UGAMNH1713;

right hypoplastron, UGAMNH1714; left and right hypoplastron,

UGAMNH1696, 1715; hypoplastron, UGAMNH1716; neural 1, pleural

and peripheral 1 and 2, UGAMNH1699; neural 1 and left and right

peripheral 1, UGAMNH1732; neural 5 and 6, UGAMNH1730; neural,

UGAMNH1707; nuchal, UGAMNH1704, 1725; right peripheral 1 and

44 Timothy S. Young and Joshua Laerm

2, UGAMNH1728; left peripheral 1 and 2, UGAMNH1726; right peripheral

1, 2 and 3, UGAMNH1692; left peripheral 1, 2, 3 and pleural 1,

UGAMNH1720; left peripheral 3, UGAMNH1722; left

peripheral 3 and 4, UGAMNHA1721; right peripheral 3 and 4,

UGAMNH1708; right peripheral 5, UGAMNH1688, 1706; left

peripheral 5, UGAMNH1691; right peripheral 6 and 7 and pleural 4

and 5, UGAMNH1710; right peripheral 6, 7, and 8, UGAMNH1712;

left peripheral 7, UGAMNH1684; left peripheral 8, UGAMNH1702;

left peripheral 8, 9, and 10, UGAMNH1694; right peripheral 9, 10,

and 11, UGAMNH1733; left peripheral 10, UGAMNH1689; right

peripheral 10 and 11, UGAMNH1695, UGAMNH1718; left peripheral

10 and 11, UGAMNH1719, 1734; right peripheral 10 and 11 and

pygal, UGAMNH1711; left peripheral 11, UGAMNH1701; left

peripheral 11 and pygal, UGAMNH1709; left and right peripheral 11

and pygal, UGAMNH1717; right peripheral 11, UGAMNH1724; left

pleural 2 and peripheral 4 and 5, UGAMNH1700; pygal,

UGAMNH1683, UGAMNH1723; left xiphiplastron, UGAMNH1327,

1729; left and right xiphiplastron, UGAMNH1693.

Remarks — Terrapene Carolina can be distinguished from its extinct

relative T. Carolina putnami based on smaller size. It is a terrestrial

woodland species (Conant 1975).

Terrapene Carolina putnami Hay

Giant Box Turtle

Material — Right epiplastron, UGAMNH1860; left hypoplastron at

inguinal notch, UGAMNH1855; left hypoplastron and epiplastron and

entoplastron, UGAMNH1863; right hypoplastron and xiphiplastron,

UGAMNH1864; neural 1 and pleural and peripheral 1, UGAMNH1856;

nuchal, UGAMNH1865; left peripheral 3 and 4, UGAMNH1858; right

peripheral 6 and 7, UGAMNH1859; right peripheral 10 and 11 and

pygal, UGAMNH1862; right peripheral 1, UGAMNH1849; left

peripheral 2, 3, and 4, UGAMNH1848; left peripheral 4, 5, and 6,

UGAMNH1861; right peripheral 6, UGAMNH185; left peripheral 7

and 8, UGAMNH1852; left peripheral 8 and 9, UGAMNH1853; right

peripheral 9, UGAMNH1847; left peripheral 10 and 11,

UGAMNH1854; left pleural 2 and 3 and peripheral 4 and 5,

UGAMNH1857; right pleural 2 and 3, UGAMNH1850.

Remarks — This extinct giant subspecies is common in late

Pleistocene deposits of Florida where it occurred in coastal marshes

and lowland savannahs. (Auffenberg 1958, Kurten and Anderson 1980).

It is readily distinguishable on the basis of its large size.

St. Marks River Fauna 45

Family Testudinidae

Testudinidae gen. et sp. indet.

Material— Osteoderms, UGAMNH1645, UGAMNH1646.

Remarks — These specimens represent a large tortoise, but the

osteoderms are not diagnostic.

Geochelone sp. indet.

Material— Pleural, UGAMNH1638; left hypoplastron,

UGAMNH1639; right pleural 2, UGAMNH1640; left pleural 4,

UGAMNH1641.

Remarks — The available material, while certainly Geochelone,

could not be referred to a species with confidence.

Geochelone incisa (Hay)

Material— Right peripheral 7, UGAMNH1642; nuchal

UGAMNH1643; right peripheral 5, UGAMNH1644.

Remarks — This material compares well with the series of G.

incisa in the collections of the Florida Museum of Natural History

and corresponds to Auffenberg's (1963) description. The was apparently

an open grassland inhabitant thought to require a frost free winter

(Kurten and Anderson 1980); however, Martin and Guilday (1967)

disagree.

Gopherus polyphemus (Daudin)

Gopher Tortoise

Material— Nuchal, UGAMNH1637.

Remarks — This material compares well with modern Gopherus

polyphemus which ranges in dry sandy soils (Conant 1975).

Family Trionychidae

Trionyx sp. indet.

Material — Carapacial fragment, UGAMNH1761.

Remarks — The available material, while certainly Trionyx because

of the distinctive pattern on the bone, could not be referred to a

species with confidence.

Order Squamata

Family Colubridae

Colubridae gen. et spec, indet.

Material— Vertebrae, UGAMNH2054-2061.

46 Timothy S. Young and Joshua Laerm

Elaphe obsoleta (Say)

Rat Snake

Material— Vertebra, UGAMNH2055.

Remarks — This material compares well with modern Elaphe

obsoleta which may be found in woodlands and grasslands (Conant

1975).

Order Crocodilia

Family Alligatoridae

Alligator mississippiensis (Daudin)

American Alligator

Material — Left angular, UGAMNH1015; distal phalanx,

UGAMNH1001, right dentary (without teeth), UGAMNH1012; dermal

scutes, UGAMNH1003-1011 (1010 and 1011 exhibit crossmends),

UGAMNH1023; right femur, UGAMNH1020, 1022; left humerus,

UGAMNH1019; fused parietals, UGAMNH1025; left scapula,

UGAMNH1014; right scapula, UGAMNH1016, 1018; teeth,

UGAMNH1002, 1024, 1026, 1028, 1029; vertebra, UGAMNH1017;

frontal, UGAMNH1013; left jugal, UGAMNH1021.

Remarks — This material has the distinctive Alligator mississippiensis

morphology and it compares well with modern examples. Alligators

occur in both fresh and brackish waters (Conant 1975).

CLASS AMPHIBIA

Order Caudata

Family Sirenidae

Siren sp. indet.

Material— Vertebrae, UGAMNH2129-2131, 2161.

Remarks — The available material compares well with modern

Siren.

Order Anura

Anura gen. et sp. indet.

Material— Vertebrae, UGAMNH2132-2134-right humerii.

Remarks — The available material, while certainly frog, could not

be referred to a genus or species with confidence.

CLASS OSTEICHTHYES

Order Lepisosteiformes

Family Lepisosteidae

Lepisosteus sp. indet.

Material— Scales, UGAMNH2 109-21 11.

St. Marks River Fauna 47

Remarks — The scales, while certainly Lepisosteus, could not be

referred to a species with confidence. Lepisosteus occurs in freshwater

and estuarine habitats (Hoese and Moore 1977, Lee et al. 1980).

Order Amiiformes

Family Amiidae

Amia calva Linnaeus

Bowfin

Material— Left dentary, UGAMNH2088; left frontal,

UGAMNH2089; cervical vertebra, UGAMNH2090.

Remarks — This material compares well with modern specimens

of Amia calva. The bowfin is a freshwater and estuarine species

(Hoese and Moore 1977, Lee et al. 1980).

Order Siluriformes

Family Ictaluridae

Ictaluridae gen. et sp. indet.

Material— Spine, UGAMNH2112; vertebra, UGAMNH2113.

Remarks — The available material, while certainly catfish, could

not be referred to a genus or species with confidence.

Pylodictis cf. P. olivaris (Rafinesque)

Flathead Catfish

Material — Left proximal coracoid, UGAMNH2119.

Remarks — The morphology of the single element is very similar

to modern specimens of P. olivaris and distinct from the other known

regional ictalurids available for comparison. The specimen at hand

shows some evidence of mineralization, but mineralization is not

extensive. The species occurrence in the St. Marks River is outside

its reported range which extends from northeastern Mexico east

throughout Gulf of Mexico drainages to Mobile Bay (Lee et al.

1980 et seq.). However, in recent times the species has undergone

introductions and populations are now known from at least the

Appalachicola-Chatahoochee System (M. and B. J. Freeman,

University of Georgia, personal communication). Uyeno and Miller

(1962) reported some specimens of P. olivaris from the Trinity River

Terrace, Texas. The deposit was dated to the Sangamon (late

Pleistocene); however, that site is within the present range of the

species. It is a freshwater species (Hoese and Moore 1977, Lee et al.

1980).

48 Timothy S. Young and Joshua Laerm

Family Ariidae

Ariidae gen. et sp. indet.

Material — Spine, UGAMNH2114-2116; cervical vertebrae,

UGAMNH2117, UGAMNH2118.

Remarks — These specimens show the characters of the marine

catfishes, although species identification is not possible.

Ariusfelis (Linnaeus)

Hardhead Catfish

Material— Spine, UGAMNH2091.

Remarks — This spine compares well with the distinctive Arius

felis morphology. This species is restricted to saltwater and estuaries

(Hoese and Moore 1977, Lee et al. 1980).

Order Salmoniformes

Family Esocidae

Esox sp. indet.

Material— Right dentary, UGAMNH2101, left dentary

UGAMNH2095, 2096, 2098-2100, 2102-2105; dentary,

UGAMNH2097; parasphenoid, UGAMNH2106; pharyngeal grinding

plates, UGAMNH2107, 2108.

Remarks — These specimens closely resemble both E. americanus

Gmelin and E. niger Lesueur. Both are considered freshwater species

(Lee et al. 1980 et seq.) and occur in regional waters today.

Order Perciformes

Family Percichthyidae

Morone saxatilis (Walbaum)

Striped Bass

Material— Right maxilla, UGAMNH2082; right premaxilla,

UGAMNH2083; right quadrate, UGAMNH2084, 2085; left quadrate,

UGAMNH2086; atlas, UGAMNH2087.

Remarks — This material compares well with modern examples

of Morone saxatilis which occurs in both coastal saltwater and estuaries

(Hoese and Moore 1977, Lee et al. 1980).

Family Sparidae

Archosargus probatocephalus (Walbaum)

Sheepshead

Material — Right dentary, UGAMNH2079; left preoperculum,

UGAMNH2080; tooth, UGAMNH2081.

Remarks — This material compares well with modern examples of

St. Marks River Fauna 49

Archosargus probatocephalus. The sheepshead is a coastal salt-

water and estuary species (Hoese and Moore 1977, Lee et al. 1980).

Family Sciaenidae

Sciaenops ocellatus (Linnaeus)

Red Drum

Material— Quadrate, UGAMNH2092.

Remarks — This material compares well with modern examples of

Sciaenops ocellatus. It is a coastal saltwater species, but is also

associated with estuaries (Hoese and Moore 1977, Lee et al. 1980).

Family Mugilidae

Mugil sp. indet.

Material— Vertebrae, UGAMNH2093, UGAMNH2094.

Remarks — The available material, while certainly Mugil, could

not be referred to a species with confidence. Mugil is a coastal saltwater

species (Hoese and Moore 1997, Lee et al. 1980).

RESULTS AND DISCUSSION

Chronology and Environment of Deposition

Of several thousand separate skeletal elements recovered from

the St. Marks River, 1,162 were referable to specific taxa. Included

are 37 species of mammals, 3 birds, 13 reptiles, 2 amphibians, and

9 fish. An additional 23 species of birds were identified from the

1972 collection made by Storrs Olson. Of all species we reported,

14 mammals and 2 reptiles are restricted to the Pleistocene. The

remaining are representative of the modern extant regional fauna.

With the exception of modern contaminants, the latter are acceptable

Pleistocene species; however, they more probably represent a mixture

of Holocene and Pleistocene material. This is reflected in the range

of mineralization observed in many species. In all cases those species

known only from the Pleistocene are well mineralized. However,

several species with both a Pleistocene and Recent occurrence such

as horse and deer exhibit both well mineralized and, what appears

to be, very recent unmineralized condition. Modern contaminants such

as cow and pig are unmineralized. In general, mineralization is no

criterion of Pleistocene deposition. The problem of apparent

heterochronous deposition and separation of Pleistocene and Holocene

materials is exacerbated by the apparent rapid mineralization that

can occur in reducing environments. Neill (1957) noted that rapid

mineralization of organic remains in Florida creates the illusion that

Recent material is of older age. Nonetheless, the St. Marks River

50 Timothy S. Young and Joshua Laerm

fauna is clearly mixed and reflects heterochronous deposition over

time beginning no later than the late Pleistocene (Wisconsinan) and

extending through the Recent.

We compared the St. Marks River faunal list and a modern

regional faunal list of the Apalachicola River system (Means 1976).

Of the 344 species listed by Means, 29% of the mammals, 10% of

the birds, 19% of the reptiles, 5% of the amphibians, and 2% of

the fish are represented in the St. Marks River fauna. This bias

toward mammals probably reflects taphonomic factors associated with

the larger size of mammalian elements in a fluvial environment. Small,

more fragile vertebrates (birds, reptiles, amphibians, and fish) are clearly

under-represented in the St. Marks River fauna. This bias is reflected

also in the mammalian fauna where chiropteran, insectivoran, and small

rodent remains are conspicuously absent.

While many of the species recovered from the St. Marks River

are eurytopic and provide only limited information regarding the

environment of deposition, a number are stenotopic and are considered

good environmental indicators.

Mammals — The mammalian fauna, in particular, is very useful

in assessing the chronology and paleoenvironment of the St. Marks

River. The reason for this is two-fold. First, mammals are the most

numerous and have the largest component of extinct forms. Second,

Florida has an extremely rich and well-documented late Pleistocene

as well as modern mammalian fauna upon which comparisons to the

St. Marks River fauna can be made.

Thirteen (35%) of the mammalian fauna of the St. Marks River

is represented by extinct forms. These include Holmsina septentrionalis,

Megalonyx jeffersonii, Glossotherium harlani, Canis dims, Smilodon

sp., Synaptomys australis, Tapirus, sp., Equus sp., Platygonus

compressus, Hemiauchenia macrocephala, Paleolama mirifica,

Mammut americanum, and Mammuthus jeffersonii. This closely

approximates the relative percentage of extinct mammals from a

number of Rancholabrean faunas from elsewhere in Florida (Martin

and Webb 1974). The temporal span of the extinct forms ranges

from Blancan through Recent. However, they all share a late

Wisconsinan chronology. Those species representing extant forms,

although individually some exhibit a longer stratigraphic history, also

share a late Wisconsinan chronology. With few exceptions, all the

extant species are represented in the local fauna today.

Comparison of the known and inferred habitat preferences or

requirements of the extant and extinct mammalian species suggests

the depositional environment was heterogeneous. On one hand there

are a number of essentially woodland species: Didelphis, Holmsina,

St. Marks River Fauna 51

Megalonyx, Lutra, Mephitis, Urocyon, Ursus, Tapirus, Platygonus,

Odocoileus, and Mammut. However, grassland species are well represented

also: Glossotherium, Mephitis, Geomys, Equus, Hemiauchenia,

Paleolama, Bison, and Mammuthus. From a simple listing it might

appear that grassland species are about as common as woodland

species. However, when compared by the number of identified

specimens per taxon, woodland species are more prevalent. Despite

criticism, this method is reliable for a comparison of relative abun-

dances of species (Grayson 1984). In addition, a number of species

indicate proximity of water: Didelphis, Lutra, Procyon, Ursus, Castor,

Neofiber, Ondatra, Synaptomys, and Tapirus are all typically

associated with moist, riparian, or standing water habitats.

Birds — Storrs Olson's collection from the St. Marks River have

never been published. He was kind enough to provide a list of the

birds identified and has permitted us to include it in the present

discussion. Olson (personal communication) felt that "there was very

little of interest among the birds" mainly because the list of avian

species recovered from the St. Marks River is essentially similar to

the modern fauna (Means 1976). As a whole, birds are uninstructive

concerning the dating of the St. Marks River fauna. They do, however,

provide considerable information relating to the environment of

deposition.

The St. Marks River avian fauna is clearly biased toward large

species with predominantly salt and freshwater marshland habitat

preferences: Podiceps, Podilymbus, Phalacrocorax, Ardea, Butorides,

Egretta, Eudocimbus, Aix, Anas, Aythya, Branta, Bucephala,

Lophodytes, Padion, Fulica, Gallinula, and Aramus. In addition, a

number of the species are typically associated with woodlands or

woodland riparian habitats: Aix, Mergus, Buteo, and Strix.

Conspicuously absent are the passeriforms. This probably represents

the taphonomic bias referred to above. While a significant number

of the birds are often present in saltwater marsh habitats, there are

no shorebird (charadriform) species present.

Reptiles and Amphibians — Many turtles, but few other reptiles,

are reported from the St. Marks River. Emydid turtles, in particular,

are well represented and make up approximately 90% of the recovered

reptilian material. In fact, in numbers alone they make up well over

one third the individual elements in the fauna. The emydid turtle

species identified from the 1987 collection were Pseudemys concinna,

P. floridanus, P. nelsoni, Trachemys scripta, and Terrapene Carolina,

all of which are found in the area today. Pseudemys and Trachemys

are indicative of a freshwater environment, while Terrapene is

terrestrial. An extinct, large, late Pleistocene subspecies of Terrapene

52 Timothy S. Young and Joshua Laerm

Carolina, T. c. putnami, is represented in the St. Marks River fauna

by a number of elements. It was probably limited to the Coastal

Plain and Savannah habitats (Auffenberg 1958) and is represented in

many late Pleistocene sites in Florida. Other aquatic turtles recovered

include one chelydrid, C. serpentina, and a number of unreferrable

kinosternid fragments. Terrestrial testudinoid turtles present at the site

are Geochelone incisa, Geochelone sp., and Gopherus polyphemus.

Geochelone incisa represents a definite late Pleistocene species, as

does Terrapene Carolina putnami. Gopherus polyphemus occurs in the

area today.

Only two snakes, Nerodia sp. and Elaphe obsoleta, were

identified from the 1987 collection. Both snakes occur in the area

today. No lizards were identified from any of the fossil collections.

Two amphibians were recovered, one caudate and one anuran, neither

of which could be identified to species.

With the exception of the two late Pleistocene components, the

herpetofauna is representative of the modern regional fauna and

includes both lower Coastal Plain riverine and marshland species,

as well as terrestrial forms.

Fishes — The fish fauna described includes both freshwater and

marine forms. Ariopsis felis, Morone saxatilis, Archosargus

probatocephalus, Sciaenops ocellata, and Mugil sp. although

typically marine are also estuarine tolerant. The freshwater fishes

include Pylodictis cf. P. olivaris, Lepisosteus sp., Esox sp., and Amia

calva. Of these, Lepisosteus sp., Esox sp., and A. calva tolerate

estuarine, but not marine, conditions (Hoese and Moore 1977).

In conclusion, the aquatic community suggests a mixed freshwater

and marine, or more likely an estuarine environment, similar to the

lower half of the St. Marks River drainage today. The terrestrial

fauna indicates a wooded riparian environment also similar to that

found in the St. Marks River drainage today. However, the presence

of Hemiauchenia, Bison, Equus sp., and Mammumthus coupled with

Geomys, Geochelone, and Gopherus suggests that more open, semi-

forested savannah habitats were also represented. This is consistent

with other late Pleistocene (Rancholabrean) faunas from the

panhandle of Florida, some of which are considered below.

Faunal Comparison

The Chipola River sites (IA and HA) — This is a river deposit

similar to the St. Marks River and contains similar species including

Didelphis virginiana, Holmesina septentrionalis, Castor canadensis,

Procyon lotor, Bison sp., Equus sp., Mammut americanum,

Odocoileus virginianus, and Hemiauchenia macrocephala (Webb

St. Marks River Fauna 53

1914a). Although no formal paleontological description of the site

exists, the species present in that assemblage indicate a mixed

woodland/grassland environment (Webb 1974a).

The Aucilla River IA site — The site is also similar to the St.

Marks River in depositional and temporal characters. No published

paleontological description exists for this site either, but from the

fauna a habitat of woodland and marsh can be assumed. It includes

Didelphis virginiana, Holmesina septentrionalis, Glossotherium cf. G.

harlani, Ondatra zibethicus, Castor canadensis, Neochoerus

pinckneyi, Sylvilagus floridanus, Canis dims, and Tremarctos

floridanus (Webb 1974a).

Wakulla Springs — This, too, is similar to the St. Marks River

in depositional and temporal characters. Included are Mammuthus sp.,

Mammut americanum, and Bison bison antiquus (Webb 1974a). No

formal paleontological description of the site exists.

Generally there are only slight differences between the St.

Marks River and other Florida panhandle, riverine deposits. These

differences can probably be attributed to a number of causes including

collection by amateurs, undersampling, taphonomic events, or other

collecting biases.

Compared to the other Rancholabrean faunas from peninsular

Florida (Martin and Webb 1974, Webb 1974a, Webb and Wilkins

1984), the St. Marks River assemblage probably is not representative

of the full late Pleistocene fauna that existed in the area. For example,

more than 50 species of mammals are known to have been present

in Florida during the time of accumulation of the Ichetucknee River

fauna, Columbia County, Florida (Martin and Webb 1974). As shown

by Martin and Webb (1974) mammalian faunal diversity was

considerably elevated in peninsular Florida during Rancholabrean time,

and it is highly likely that is was the case along the rich fluvio-

estuarine environment of the panhandle during the same period.

ACKNOWLEDGMENTS— Thanks are due Chris McKensie, Locke

Rogers, Brad Newsom, Tim Gaudin, and Luis Insignares for their

efforts below and above the surface of the St. Marks River. Chris

McKensie deserves added thanks for the many long hours he spent

screening and sorting bones. The United States Department of

Agriculture provided an advance copy of their publication on the

soils of Wakulla County. Gary Morgan and Russ McArty at the

Florida Museum of Natural History were more than just helpful.

The gracious offer of Storrs Olson to make available his unpublished

records of the avian material he collected and analyzed is greatly

appreciated. Robert Martin, Robert Frey, and Elizabeth Reitz provided

54 Timothy S. Young and Joshua Laerm

many helpful critical comments on earlier drafts. Funds for this study

were provided through Department of Zoology and the Museum of

Natural History, the University of Georgia.

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Accepted 29 July 1992

58

AUTUMN LAND-BIRD MIGRATION

ON THE BARRIER ISLANDS OF NORTHEASTERN

NORTH CAROLINA

by

Paul W. Sykes, Jr.

For three consecutive years Sykes investigated the autumn migration

of land birds in the Bodie Island and Pea Island area of coastal North

Carolina. During a 102-day period in 1965, he recorded 110,482 individual

birds of 148 species. He was able to correlate major influxes of migratory

species with specific weather patterns. His data show seasonal peaks of

southward movement for the land-bird species that pass along the North

Carolina coast in large numbers. In addition, Sykes recorded five species

native to the western United States. Three of these vagrants provided the

first reports of Swainson's Hawk, Sage Thrasher, and Western Meadowlark

for North Carolina.

1986 49 pages Softbound ISBN 0-917134-12-5

Price: $5 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: LAND-BIRD MIGRATION, N.C. State Museum of Natural

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

No Decline in Salamander (Amphibia: Caudata) Populations:

A Twenty-Year Study in the Southern Appalachians

Nelson G. Hairston, Sr., and R. Haven Wiley

Department of Biology, University of North Carolina

Chapel Hill, North Carolina 27599-3280

ABSTRACT — Identical observations, conducted 1-4 times per year

for 15-20 years at two locations in the southern Appalachians,

have yielded quantitative data on populations of six species of

salamanders. Although the numbers have fluctuated for various

reasons, there has been no trend in the numbers of any of the

species. The "world-wide decline of amphibian populations" has not

occurred in the two localities studied.

Recently, much attention has been given to a decline in many

populations of amphibians (Barringer 1990, Blaustein and Wake 1990,

Phillips 1990). There is a suggestion by some authors that there is a

general cause for a supposed "world-wide" decline. We do not deny

that many amphibian species have decreased in abundance. Among

the causes that have been suggested are acid precipitation (Harte and

Hoffman 1989, Beebee et al. 1990) and ultraviolet increase due to

ozone depletion (Barringer 1990, Blaustein and Wake 1990, Phillips

1990). The same authors have considered overcollecting and rejected

it as a general cause. Habitat destruction is also widely mentioned.

The last cause is common to all species, except for some pioneering

ones, and would not apply only to amphibians. The situation is regard-

ed by many herpetologists as very serious, so much so that the World

Conservation Union (IUCN), Species Survival Commission, has activated

a Declining Amphibian Populations Task Force. This group has estab-

lished local subgroups throughout the United States and elsewhere in

the Americas to promote research on the problem.

If there has been a general cause for the decline in amphibian

populations, all amphibian populations should be involved; if they are

not, the original claim of a "world-wide decline" must be modified,

either by eliminating some taxonomic groups, some ecologically distinc-

tive species (e.g., those lacking aquatic stages), or some geographic

regions. Study of apparently exceptional cases might give clues to the

causes of declines that have been observed.

METHODS

In September 1971 and 1972 N.G.H. and classes from the

University of Michigan studied the distribution of the colors of

Brimleyana 18:59-64, June 1993 59

60 Nelson G. Hairston, St., and R. Haven Wiley

Plethodon hybrids along altitudinal transects at the Coweeta Hydrologic

Laboratory, near Franklin, North Carolina, and in 1972 they also

recorded the ecological distribution of four species of Desmognathus

(Hairston 1973). In 1973 and 1974, N.G.H. studied a zone of intergrada-

tion between two forms of Plethodon jordani and the altitudinal

replacement of that species and P. glutinosus at Heintooga Overlook

in the Great Smoky Mountains National Park near the junction of the

Balsam Mountains and the Great Smoky Mountains in Haywood and

Swain counties, North Carolina. The name glutinosus is controversial

for this form; Highton (1983) proposed the name teyahalee, which

we believe to be misapplied (Hairston 1992).

N.G.H. also continued the observation of Plethodon hybrids at

Coweeta in 1974. Beginning in September 1976 and for each year

thereafter, we have led one to four (usually two) undergraduate

classes of 15 students each to both localities and made carefully

repeated observations of the same kind.

At Heintooga, the students were instructed to capture 10

Plethodon, return with them to the vehicles, and examine the animals'

cheeks for the amount of red color; specimens were then returned to

the forest. The exercise was repeated at 3.2, 6.4, 9.7, 11.3, and 13.7

km along the National Park Service road to Round Bottom Camp

Ground, with species identifications made at the last two stops. The

elevations ranged from 1,600 m at the start to 1,350 m at the last

stop. At Coweeta, the same exercise was carried out at five elevations,

starting at 686 m and continuing up at 91.5-m intervals. We and the

students evaluated the amount of red on legs and the amount of

white on sides and back. Both of these exercises were performed at

night, beginning at dark. The Desmognathus exercise involved the

students collecting specimens and noting identification of each and

the distance from nearest surface water to where they were found.

The exercise requires a period of 2-2.5 hours in the afternoon.

Each class exercise on Plethodon began at dark (2000 hours)

and ended at approximately the same time each night (2330 hours at

Heintooga and 2230 hours at Coweeta). The Desmognathus exercise

began at 1230 hours and ended at approximately 1430-1500 hours.

Thus, there has been no tendency to expend extra effort to observe

the same number of salamanders.

RESULTS

There has been no consistent trend in the number of individuals

of any of the seven populations over the 15-20 years of the study

(Figs. 1-3). All seven series show fluctuations greater than more exact

studies showed over shorter periods near the sites reported here

Salamander Populations

61

(Hairston 1987). There are several known causes for the fluctuations,

which occurred over much shorter intervals than the mean generation

times (5-10 years) for the different species (Hairston 1987). Some

were due to cold weather, when Plethodon tend to remain underground

(first class, September 1981); others were due to exceptionally

enthusiastic classes (September 1977). None of the fluctuations in

numbers observed can be attributed to a real change in the number

of salamanders actually present. The mean generation time for P.

jordani is 9.8 years and for glutinosus it is at least a year longer

(Hairston 1987). Thus, fluctuations in numbers seen at shorter intervals

do not represent real changes.

iocv

P jordoni

Fig. 1. Population history of Plethodon jordani and P. glutinosus in the

Heintooga locality, as shown in numbers observed by successive classes from

1976 to 1990. Broken lines represent preliminary observations not exactly

equivalent to class date. Arrows show means of all class data; standard

error not given because the counts might not be independent because of the

longevity of individual salamanders and the likelihood that at least some of

the same ones were observed on successive class exercises.

62

Nelson G. Hairston, St., and R. Haven Wiley

8a

60

'40

Plethodon Hybrids

Fig. 2. History of the Plethodon population at Coweeta. Symbols are the

same as in Figure 1.

D. quadramoculatus

<*40

A ochrophaeus

Fig. 3. Population histories of four species of Desmognathus at 686-m

elevation, Coweeta Hydrological Laboratory. Symbols are the same as in

Figure 1.

Salamander Populations 63

DISCUSSION

As far as we know, this is the longest series of continuous

quantitative observations on any amphibian populations. Other

multiple-year studies include 13 years for Savannah River Ecology

Laboratory studies (Pechmann et al. 1991) and the same duration for

Taricha rivularis in California (Twitty 1966). In the former, changes

could be explained by drought, and the latter was completed long

before the supposed general decline of amphibian populations.

Our observations bear on some of the suggested causes for long-

term declines in amphibian populations. There is considerable evidence

that the observed dieback and decline of spruce-fir forests in the

southern Appalachians is due to atmospheric pollution (Bruck 1988,

Dall et al. 1988, Zedaker et al. 1988). As the salamander populations

have remained essentially in steady states, acid rain and ozone depletion

cannot be universal causes of all declines in amphibian populations.

As the great majority of records of population declines are based

on anecdotal evidence, we remain skeptical of the generality of these

declines until similar long-term records are produced. We are also

convinced that over-collecting by biological supply companies and by

some herpetologists has been underrated as a possible cause of observed

declines.

ACKNOWLEDGMENTS— Vie thank successive officials of the

Great Smoky Mountains National Park for permission to carry out

the study at Heintooga. We also thank Wayne Swank, Director, Coweeta

Hydrologic Laboratory for permission to carry out the studies there.

Without the dedicated efforts of the hundreds of students in our classes,

as well as those of the teaching assistants, the data could not have

been collected.

LITERATURE CITED

Barringer, M. 1990. Where have all the froggies gone? Science 247:1033-

1034.

Blaustein, A. R., and D. B. Wake. 1990. Declining amphibian populations:

a global phenomenon? Trends in Ecology and Evolution 5:203-204.

Beebee, T. J. C, R. J. Flower, A. C. Stevenson, S. T. Patrick, P. G.

Appleby, C. Fletcher, C. Marsh, J. Natkanski, B. Rippey, and R. W.

Batterby. 1990. Decline of the natterjack toad Bufo calamita in

Britain: paleoecological, documentary and experimental evidence for

breeding site acidification. Biological Conservation 53:1-20.

Bruck, R. I. 1988. Research site: Mount Mitchell (southern Appalachians).

Decline of red spruce and Fraser fir. United States Department of

Agriculture Forest Service, General Technical Report 120: 133-143.

64 Nelson G. Hairston, Sr., and R. Haven Wiley

Dall, C. W., J. D. Ward, H. D. Brown, and G. W. Ryan. 1988. Proceedings

of the United States/Federal Republic of Germany research symposium:

effects of atmospheric pollutants on the spruce-fir forest of the eastern

United States and the Federal Republic of Germany. United States

Department of Agriculture Forest Service, General Technical Report

NE 120:107-110.

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

21.

Hairston, N. G. 1987. Community ecology and salamander guilds. Cambridge

University Press, New York, New York.

Hairston, N. G. 1992. On the validity of the name teyahalee as applied to

a member of the Plethodon glutinosus complex (Caudata: Plethodontidae):

a new name. Brimleyana 18:59-64.

Harte, J., and E. Hoffman. 1989. Possible effects of acidic deposition on

a Rocky Mountain population of the tiger salamander, Amby stoma tigrinum.

Conservation Biology. 3:149-158.

Highton, R. 1983. A new species of woodland salamander of the Plethodon

glutinosus group from the southern Appalachian mountains. Brimleyana

9:1-20.

Pechmann, J. H. K., D. E. Scott, J. W. Gibbons, R. D. Semlitsch, L. J.

Vitt, and J. P. Caldwell. 1991. Declining amphibian populations: the

problem of separating human impacts from natural fluctuations. Science

253:892-895.

Phillips, K. 1990. Frogs in trouble. International Wildlife 20:4-11.

Twitty, V. 1966. Of scientists and salamanders. W. H. Freeman, San

Francisco, California.

Zedaker, S. M., N. S. Nicholas, C. Eagar, P. S. White, and T. E. Burk.

1988. United States Department of Agriculture Forest Service, General

Technical Report NE 120:123-131.

Accepted 7 August 1992

On the Validity of the Name teyahalee as Applied to a Member

of the Plethodon glutinosus Complex

(Caudata: Plethodontidae): A New Name

Nelson G. Hairston, Sr.

Department of Biology, University of North Carolina,

Chapel Hill, North Carolina 27599-3280

ABSTRACT — The name Plethodon teyahalee (Hairston) cannot be

applied to the member of the P. glutinosus complex as designated

by Highton (1983). Biochemical data show that the population from

which the type of teyahelee was taken consists of hybrids between

local populations representing the P. jordani and P. glutinosus com-

plexes, and thus cannot be applied to a member of either of those

two species under Article 23(h) of the International Code of Zoo-

logical Nomenclature (1985). A new name, Plethodon oconaluftee,

is proposed, and a new type is designated.

Plethodon glutinosus, a salamander distributed widely over the

eastern United States, has recently been divided into 16 species on

the basis of allozyme frequencies (Highton 1983, 1989). Most of

these forms occupy non-overlapping distributions, and it is not known

at present whether they are allopatric or parapatric. The form that is

distributed west of the French Broad River throughout southwestern

North Carolina and immediately adjacent parts of Tennessee, Georgia,

and South Carolina is one of the few that overlaps any adjoining

species of the complex without hybridization. In extreme southeastern

Tennessee and extreme southwestern North Carolina, it overlaps P.

aureolus and P. glutinosus (sensu stricto). Highton (loc. cit.) has

appropriated the name teyahalee for this representative of the glutinosus

complex.

In 1950 I described a form from Teyahalee Bald in the Snowbird

Mountains of southwestern North Carolina as P. jordani teyahalee,

believing it to be closely related to other subspecies of P. jordani

(Hairston 1950). The presence of red spots on the legs of some

individuals indicated the population's relationship to P. j. shermani

of the Nantahala Mountains, and the greenish-yellow spots on the

sides appeared to make it unique. Subsequent collectors have failed

to find any specimens with the greenish-yellow spots, and Highton

(1962), in a review of the genus, argued that they could be explained

as follows: "Sometimes the lateral pigment of large specimens (of

glutinosus) is more yellowish than in small ones, but structurally the

pigment appears the same." He did not comment on the detailed

differences between the white spots of P. glutinosus and those of

Brimleyana 18:65-69, June 1993 65

66 Nelson G. Hairston, Sr.

some populations of P. jordani figured by Hairston and Pope (1948).

His conclusion was that only a representative of glutinosus is present

on Teyahalee Bald and that it has genetically swamped a pre-existing

form of jordani (Highton and Henry 1970); Highton 1972, 1989),

using that as his justification for appropriating the name teyahalee.

We have known for more than 50 years that the high-altitude

red-legged form of P. jordani and the low-altitude white-spotted form

then known as glutinosus are hybridizing at intermediate elevations

throughout the Nantahala Mountains, a short distance from Teyahalee

Bald (Bishop 1941, Highton and Henry 1970). As the hybrid zone in

the Nantahala Mountains is spreading toward higher elevations

(Hairston et al. 1992), Highton's interpretation appears reasonable. More

recently, some hybridization has been found at other localities, but

not in the area between the Tuckaseegee and French Broad rivers,

nor in the western two-thirds of the Great Smoky Mountains, nor in

the Cheoah, Max Patch, or Sandy Mush mountains, nor in the southern

95% of the Balsam Mountains, i.e., not in more than half of the

distribution of this representative of the glutinosus complex.

The important question is the status of the population of Plethodon

on Teyahalee Bald. Allozyme data presented by Peabody (1978) show

that these animals are intermediate between neighboring populations

of jordani and the low-altitude representative of the glutinosus complex.

In fact, the calculated values of Nei's Genetic Identity are more

similar to the nearest populations of jordani than they are to the

nearest populations of the glutinosus complex (Table 1). The genetic

swamping is thus so incomplete that the entire population on Teyahalee

Bald must be regarded as hybrids, and judging from the history in

the adjacent Nantahala Mountains have been hybrids since at least

1938 (Bishop 1941) and probably earlier (Hairston et al. 1992).

Table 1. Genetic identities (Nei's I [Nei 1972]) among the Teyahalee Bald

population, the nearest populations of the Plethodon glutinosus complex, and the

nearest populations of the P. jordani complex. Note that both jordani and

glutinosus are represented at Cheoah and Unicoi West. Data from Peabody (1978).

Validity of the Name teyahalee 67

The situation on Cheoah requires comment. No hybridization occurs

there, and the samples of the two species are therefore distinct. That

representative of the P. jordani complex is more distantly related to

the other four populations than they are to each other. The average

genetic identity between it and them is 0.857 (range = 0.813-0.895);

the average identity among the other four populations is 0.932 (range

= 0.900-0.967). The population on Teyahalee Bald is closely related

to those four representatives of P. jordani, but not to the Cheoah

representative.

It appears, therefore, that what I described as Plethodon jordani

teyahalee was a hybrid, and under Article 23(h) of the International

Code of Zoological Nomenclature the name teyahalee cannot be used

for that part of the glutinosus complex to which it was applied by

Highton (1983, 1989), because that is one of the parent species. To

avoid future confusion I have collected a new type for this form

from an area where hybridization with P. jordani is unknown, and I

propose the name Plethodon oconaluftee.

The following synonomic list is taken from Highton (1989):

Plethodon glutinosus (Green): Brimley (1912) (part), Highton (1970)

(part) [actually Highton and Henry (1970)]. Plethodon jordani

teyahalee Hairston (1950:269). Plethodon jordani Blatchley: Highton

(1962). Plethodon (glutinosus) glutinosus (Green): Bishop (1941)

(part). Plethodon teyahalee Hairston: Highton (1984) [actually

Highton 1983].

Holotype— GSMNP 33339, an adult female collected 16 May

1991, by N. G. Hairston, Sr., Pisgah National Forest, beside Forest

Service Road 140 near the North Fork of the French Broad River at

an elevation of 930 m on the south-facing slope of the Balsam

Mountains, Transylvania County, North Carolina. Snout to posterior

angle of vent, 75 mm; numerous very small white spots on back and

top of tail, a few on top of head; numerous irregularly shaped

white spots on sides and cheeks; underside dark throughout, including

throat and chin, which have a number of irregular white spots.

Paratype — GSMNP 33340, an immature female (about 3 years

old) collected in same place as the type on 17 May 1991 by M. P.

Hairston. Snout to posterior angle of vent, 39 mm; dorsum, sides,

head, and cheeks as for type; belly dark, throat and chin paler than

in type, with many melanin-free spots, but with white pigment only

in a few lateral ones. Both types have been deposited in the collections

of the Great Smoky Mountains National Park.

The following diagnosis and distribution are quoted from Highton

(1989), which I use because we discuss the same taxonomic entity:

"Diagnosis: A large, light-chinned species with very small white dorsal

68 Nelson G. Hairston, Sr.

spots, reduced lateral spotting, and often with small red spots on the

legs. The unique combination of genetic alleles that distinguishes P.

teyahalee from other species of the P. glutinosus group is Pgi allele

c and Trf allele a are characteristic of P. teyahalee populations but

are usually rare or absent in the other species." (Highton 1989:54)

("teyahalee" used because of the direct quotation).

"Distribution: West of the French Broad River in the Blue

Ridge physiographic province of southwestern North Carolina

and in immediately adjacent Tennessee. It also occurs in

northern Rabun County, Georgia, and in Oconee, Pickens,

Anderson, and Abbeville counties, South Carolina." (Highton

1989:54).

ACKNOWLEDGMENTS— I thank Richard Highton for a friendly

discussion of the issues involved and for suggesting the locality

from which the types of P. oconaluftee were collected. Three anony-

mous reviewers made constructive suggestions.

LITERATURE CITED

Bishop, S. C. 1941. Notes on salamanders with descriptions of several

new forms. Occasional Papers of the Museum of Zoology. University

of Michigan. 451:1-21.

Hairston, N. G. 1950. Intergradation in Appalachian salamanders of the

genus Plethodon. Copeia 1950:262-273.

Hairston, N. G., and C. H. Pope. 1948. Geographic variation and speciation

in Appalachian salamanders (Plethodon jordani Group). Evolution

2:266-278.

Hairston, N. G, R. H. Wiley, C. K. Smith, and K. A. Kneidel. 1992.

The dynamics of two hybrid zones in appalachian salamanders of the

genus Plethodon. Evolution 46:930-938.

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

Plethodon. Bulletin of the Florida State Museum 6:235-367.

Highton, R. 1972. Distributional interactions among North American

salamanders of the genus Plethodon. Pages 139-188 in The

distributional history of the biota of the southern Appalachians

(P. C. Holt, editor). Research Division Monograph 4. Virginia

Polytechnic Institute and State University, Blacksburg.

Highton, R. 1983. A new species of woodland salamander of the

Plethodon glutinosus group from the southern Appalachian Mountains.

Brimleyana 9:1-20.

Highton, R. 1989. Biochemical evolution in the slimy salamanders of the

Plethodon glutinosus complex in the eastern United States. Illinois

Biological Monograph 57:1-153.

Validity of the Name teyahalee 69

Highton, R., and S. A. Henry. 1970. Evolutionary interations between

species of North American salamanders of the genus Plethodon.

Evolutionary Biology 4:211-256.

International Code of Zoological Nomenclature, Third edition. 1985.

International trust for zoological nomenclature, University of

California Press, Berkeley, California.

Nei, M. 1972. Genetic distance between populations. American

Naturalist 106:283-292.

Peabody, R. B. 1978. Electrophoretic analysis of geographic variation and

hybridization of two Appalachian salamanders, Plethodon jordani and

Plethodon glutinosus. Ph.D. Thesis. University of Maryland, College

Park.

Accepted 26 March 1992

70

ATLAS OF NORTH AMERICAN FRESHWATER FISHES

by

D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins,

D. E. McAllister, J. R. Stauffer, Jr., and many collaborators

This very useful book provides accounts for all 777 species of fish

known to occur in fresh waters in the United States and Canada. Each

account gives a distribution map and illustration of the species, along with

information on systematics distribution, habitat, abundance, size, and general

biology.

"[It] represents the most important contribution to freshwater fishes

of this continent since Jordan and Evermann's 'Fishes of North and Middle

America' over 80 years ago." — Southeastern Fishes Council Proceedings.

1980 825 pages Index Softbound ISBN 0-917134-03-6

Price: $25 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: FISH ATLAS, N.C. State Museum of Natural Sciences,

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

ATLAS OF NORTH AMERICAN FRESHWATER FISHES

1983 SUPPLEMENT

by

D. S. Lee, S. P. Platania, and G. H. Burgess

The 1983 supplement to the 1980 Atlas of North American Freshwater

Fishes treats the freshwater ichthyofauna of the Greater Antilles. In addition

to this bound supplement, there are 19 accounts, mostly species not described

in 1980, in looseleaf form to the added to the 1980 volume. Illustrated by

Renaldo Kuhler.

1983 67 pages Index Softbound ISBN 0-917134-06-0

Price: $5 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: FISH ATLAS, N.C. State Museum of Natural Sciences,

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

Differences in Variation in Egg Size for Several Species of

Salamanders (Amphibia: Caudata) That Use

Different Larval Environments

Christopher King Beachy1

Department of Biology, Western Carolina University,

Cullowhee, North Carolina 28723

ABSTRACT — Comparative descriptive data are provided on

variation of egg size in five species of salamanders. The species

differ in their use of larval habitats. Ambystoma maculatum uses

temporary, rain-filled pools in the southern Appalachian

Mountains. Desmognathus aeneus is a direct developer and is not

constrained by risk of larval desiccation. The remaining three

species, Eurycea wilderae, D. ochrophaeus, and D. santeetlah, have

permanent streams as their larval environment. Using the

coefficient of variation (CV), I document both variation within

individual clutches and variation at the interclutch level. The degree

of variation differs among individual clutches and among

species. Variation at the intraclutch level does not agree with

that predicted. However, variation at the interclutch level

conforms to the prediction that A. maculatum (which utilizes

ephemeral larval environments) exhibits the highest degree of

variation in egg size.

In many populations of biphasic amphibians, the key factor

underlying the timing of metamorphosis and larval survivorship is

the time for which the larval environment remains hospitable. In a

permanent larval habitat, where mortality from desiccation is unlikely,

the larval period of an amphibian can be long. For example,

paedomorphic species of salamanders inhabit permanent bodies of

water. However, many species of amphibians inhabit larval

environments that are temporary and unpredictable, and desiccation

is a threat to species that use those bodies of water. Because

environmental pressure to escape the larval environment can vary

from year to year, species that breed in temporary pools may exhibit

different reproductive strategies than species that use permanent

bodies of water.

Parental investment, one facet of the study of reproductive

strategies, has been the subject of theoretical and/or empirical studies

in amphibians (e.g., Wilbur 1977; Kaplan 1980, 1985; Crump 1981,

1984; Kaplan and Cooper 1984). These studies have documented the

'Present address: Department of Biology, The University of Southwestern Louisiana,

P.O. Box 42451, Lafayette, LA 70504-2451.

Brimleyana 18:71-82, June 1993 71

72 Christopher King Beachy

extensive variation in propagule size in amphibians. Such variation

has been interpreted as an "evolved tactic" (Capinera 1979) that

ensures that viable offspring are produced in variable habitats. In

many species of amphibians with complex life cycles, vitellogenesis

occurs in the terrestrial habitat. The female then may not be able

to receive environmental cues that indicate the size of eggs she

should produce to ensure survival of offspring in the aquatic

habitat. Because offspring can be exposed to a habitat that is variable,

there should be an optimal range in offspring size within an

individual female's clutch. As habitat variability decreases, the range

should decrease because of consistent selection for an optimal

phenotype.

Egg size also varies among clutches. A single female might

produce clutches with very different mean egg sizes (Kaplan 1987).

Kaplan and Cooper (1984) showed that in species that cannot "predict"

the stability of the environment in which their larvae will grow and

develop, the most efficient strategy will be to randomly produce a

few large eggs or many small eggs. Interclutch variation in egg size

in a female's lifetime (or within a population at one time) can

outweigh the intraclutch variation of a female's single clutch.

Egg size has been shown to influence characters that relate to

larval survival in salamanders (Kaplan 1980, 1985; Petranka 1984).

Kaplan (1985) showed that in the newt Taricha torosa (Rathke, 1883)

egg size can have profound effects on hatching size and growth

rate. Thus, egg size might affect timing of metamorphosis. If so,

then an optimal egg size can be selected for given a stable (or

permanent) larval environment. In an unreliable environment, an

optimal range of egg sizes might be the best strategy to maximize

parental fitness. In T. torosa, large eggs produce large hatchlings

that begin feeding sooner than smaller larvae. When fed ad libitum,

larger larvae will metamorphose at an earlier time and at a larger

size than conspecifics hatched from small eggs. In food-limited

situations, larvae from larger eggs still metamorphose at a larger

size but at a later time than larvae from small eggs. This interaction

among egg size, food availability, and habitat reliability suggests

that the optimal egg size can vary from season to season.

I present data on variation in egg size in five species of

salamanders. Three questions are explored: (1) Does a species that

uses temporary larval environments exhibit greater intraclutch

variation in egg size than species that use more permanent larval

habitats, such as mountain streams? (2) Is the total population

variation in egg size greater in a species that uses temporary larval

environments than in species that use mountain streams? (3) Is the

Salamander Egg Sizes 73

bulk of variation in egg size introduced at the intraclutch or

interclutch level, and is this related to habitat variability?

MATERIALS AND METHODS

Study Species

Ambystoma maculatum (Shaw, 1802) in the southern Appalachian

Mountains usually breeds in mid-winter in temporary, seasonal pools

that result primarily from heavy rains. Larvae emerge from eggs in

early spring, and metamorphosis occurs 60-120 days later (Bishop

1941, Shoop 1974). The collection sites I used in this study dry

completely 1-6 months after the rains, and on occasion they dry too

early for any larvae to transform (R. C. Bruce, personal

communication). Populations of A. maculatum in eastern North

America are known to lay one mass of eggs per clutch (Wilbur

1977, Pfingsten and Downs 1989) or two or more masses per clutch

(Bishop 1941, Pfingsten and Downs 1989). "Masses" will be referred

to as "clusters" in this article. It is unknown if the clutch of a

female A. maculatum at this locality consists of one or multiple

clusters. Eggs of A. maculatum were collected in March 1988 from

four temporary pools located in Blue Valley on the escarpment of

the Blue Ridge Mountains, Macon County, North Carolina.

Eurycea wilder ae Dunn, 1920, Desmognathus santeelah Tilley,

1981, and D. ochrophaeus Cope, 1859 lay eggs in and along

headwater streams. These sites represent permanent sources of water,

even during seasons of drought (W. Swank, Coweeta Hydrologic

Laboratory, personal communication). Fishes are uncommon in these

headwater streams. The permanence of these sources of water is evident

when one considers that several species of plethodontids have larval

periods in excess of 3 years, e.g., D. quadramaculatus (Holbrook,

1840) and Gyrinophilus porphyriticus (Green, 1827) (Bruce 1980,

1988a). Eurycea wilderae has a larval period of 1 or 2 years (Bruce

1988b). Females attach their eggs to the undersides of large rocks

where the clutch is exposed to running water. Eurycea wilderae

clutches were collected during February and March 1988 at Wolf

Creek on Cullowhee Mountain, in the Cowee Mountains, Jackson

County, North Carolina. Clutches were located by raking through

cobble of headwater seepages.

Desmognathus ochrophaeus clutches were collected from various

headwater streams in the Balsam Mountains in Haywood and Jackson

counties, North Carolina. The Balsams are a southern extension of

the Great Smoky Mountains, Swain County, North Carolina where

the D. santeetlah clutches were collected. Desmognathus santeetlah

and D. ochrophaeus females brood the eggs under moss on logs and

74 Christopher King Beachy

rocks in and along the edges of headwater streams and seepages.

These species of Desmognathus have larval periods less than one

year (Bruce 1989). Clutches of the D. ochrophaeus and D. santeetlah

were collected during July and August 1987.

Desmognathus aeneus Brown and Bishop, 1947 females oviposit

under moist logs and moss. This species is direct-developing (Wake

1966), and desiccation risk should represent less constraint to it.

Clutches of D. aeneus were collected in the vicinity of Standing

Indian Campground in the Nantahala Mountains, Macon County,

North Carolina, in May 1988.

Collection of Material

As soon as I collected them, I placed egg clutches in individual

plastic containers. If I found a brooding female with the clutch, I

collected her and placed her with it. The plastic containers were

placed in a cooler and returned to the laboratory where the egg

clutches were assigned Harrison developmental stages (Duellman and

Trueb 1986). I used a dissecting microscope equipped with an ocular

micrometer to measure egg diameters to the nearest 0.1 mm. Late

developmental stages were assigned based primarily on gill ontogeny.

Embryos of clutches in which the embryos were in later stages of

development were adjusted for developmental increases in size with

the transformation formulas of Kaplan (1979). Clutches of A.

maculatum and D. aeneus were all collected very early in

development. The other three species were collected at various

developmental stages, some of them at late stages. Embryo

diameters for the plethodontid species in late development were

recorded as the length of the longest axis of the embryo (Fig. 1).

Using this measurement protocol, I observed that plethodontid

embryos do not begin to increase in size until after Harrison stage

30 (Beachy 1988).

Analyses

Intraclutch variation was quantified by calculating the

coefficient of variation (CV) for each clutch. The CV is a statistic

that expresses the standard deviation as a percentage of the mean so

that groups having very different means can be compared. These

intraclutch CVs were subjected to a one-way ANOVA by species.

Although a single female of A. maculatum may oviposit several

clusters of eggs, I assumed that differences among these clusters do

not contribute significantly to the variance in CVs, and all A. maculatum

clusters were treated as though they were from different females

(however, see Results).

Salamander Egg Sizes

75

Fig. 1. Desmognathus santeetlah embryo at four different Harrison developmental

stages: (A) stage 25, (B) stage 35, (C) stage 40, (D) stage 45. Note spherical

orientation of embryo, even at late developmental stages. Bars = 2.0 mm.

Egg laying is not synchronous in plethodontid salamanders,

and this precludes collection of a large number of plethodontid

clutches at early stages of development. Therefore, I collected

clutches at various stages of development. To ensure that intraclutch

variation did not vary with development, a Wilcoxon's matched-pairs

signed-ranks test was conducted on 12 D. santeetlah clutches to test

if stage of development significantly influenced CV. Coefficients of

variation were calculated for two developmental stages (stage 45 and

an earlier stage, ranging from 9 to 41 for the clutches in question)

for each clutch, and no significant difference was found in CV due

76 Christopher King Beachy

to development ( x early stage = 5.43, x stage 45 = 5.96, n = 12,

Ts = 21, ns). This result was assumed to hold true for D. ochrophaeus

and E. wilderae. This assumption was not required for D. aeneus

and A. maculatum because these eggs were collected at early

developmental stages.

Interclutch variation was quantified by taking a mean egg

diameter for all clutches at, or prior to, Harrison stage 30 (see

above discussion on changes in size). These means were pooled

according to species, and a CV was calculated for each species.

These species CVs were squared and subjected to pairwise

F-tests (Lewontin 1966). Kaplan (1987) showed that Bombina

orientalis (Boulenger, 1890) females can produce clutches of

different mean egg sizes. By assuming that this is the case for the

species used in my study (i.e., the variation represented in a sample

of the population might mirror the variation introduced by a single

female in her lifetime), one can set predictions that are similar to

those for intraclutch variation.

To determine the relative contributions of intraclutch and

interclutch variation to the overall variance, egg size data for each

species were subjected to a one-way ANOVA, with individual

clutches as the factor. Relative contributions of intraclutch and

interclutch variation to overall variance for each species were

calculated using the factor and error sum-of-squares of the

ANOVA table (Sokal and Rohlf 1981).

I analyzed data with StatView512+TM following the

methods of Sokal and Rohlf (1981). In all analyses a = 0.05.

RESULTS

My hypothesis was that the variation in egg size would be

greatest in A. maculatum, the temporary pool breeder, and the lowest

in D. aeneus, which is not constrained by habitat variability and

thus should exhibit the greatest degree of canalization. Variation in

egg size should be intermediate in the other three species. Descriptive

statistics of egg size for the five species under study are presented

in Table 1.

Intraclutch Variation

Coefficients of variation of all clutches were analyzed with a

model I one-way ANOVA (Sokal and Rohlf 1981), with species as

treatment. Significant differences were found in intraclutch CV among

species. A Fisher's PLSD a posteriori test was employed to

determine the nature of the differences. The prediction that species

using ephemeral larval habitats will display larger variation in egg

size was not supported. Of all species, D. aeneus exhibited the

Salamander Egg Sizes

77

Table 1. Descriptive statistics of egg size for five species of

salamanders.3

aData are calculated using mean egg size per clutch. Only data for those

clutches at, or earlier than, Harrison developmental stage 30 are presented

(see text for explanation).

b Refers to clusters for A. maculatum.

E. wilderae

D. ochrophaeus

D. santeetlah

A. maculatum

D. aeneus

....

14

20

63

n = 37

n = 10

2 3 4 5 6 7

Intraclutch CV

8 9

10

Fig. 2. Coefficients of variation (CV) of intraclutch variation in egg size

for five species of salamanders. Number of clutches examined for each

species is indicated. Vertical lines indicate means not significantly

different using a model I ANOVA with species as treatment. (<x = 0.05).

78

Christopher King Beachy

lowest degree of variability, as expected. But A. maculatum did not

exhibit the highest degree of intraclutch variation as was expected

(Fig. 2).

Interclutch Variation

Because embryo diameter begins to increase in late developmental

stages, only those clutches of D. santeetlah, D. ochrophaeus, and E.

wilderae collected earlier than Harrison developmental stage 30 were

used in this analysis. All egg size data for A. maculatum and D.

aeneus were analyzed. Mean egg size was determined for each clutch,

and those data were used to calculate a CV of egg size for each

species (Table 1). In this analysis, A. maculatum did show the greatest

variation in egg size (Fig. 3). Except for E. wilderae, all species

CVs were significantly lower than that for A. maculatum. D. aeneus

was predicted to exhibit the lowest degree of variation in egg size;

only D. ochrophaeus has a lower CV, although this difference was

not significant.

A. maculatum

E. wilderae

D. santeetlah

D. aeneus

D. ochrophaeus

6 8

Pooled CV

10

12

14

Fig. 3. Coefficients of variation (CV) of pooled (interclutch) variation in

egg size for five species of salamanders. The species CVs are tested with

an F-test of the squared CVs (Lewontin 1966). Vertical lines indicate means

not distinguishable by pairwise F-tests (P > 0.05).

Salamander Egg Sizes

79

A. maculatum

E. wilderae

D. aeneus

D. ochrophaeus

D. santeetlah

40 60

Percent of variation

100

Fig. 4. Relative contributions of intraclutch and interclutch variation in

egg size to overall variance. For each species, a separate model II one-

way ANOVA of egg size was conducted with clutches as the factor, thus

a total of five ANOVAs. Contributions are calculated using the factor and

error sum-of-squares of the ANOVA table. Black bars represent intraclutch

variation, and hatched bars represent interclutch variation.

Interclutch and Intraclutch Contribution to Variation

Partitioning of the variance components demonstrated the

respective contributions of intraclutch and interclutch variation to the

sample variance in egg size (Fig. 4). For E. wilderae and A. maculatum,

interclutch contributions to the variance outweighed intraclutch

contributions. In all three desmognathines, interclutch and

intraclutch contributions were approximately equal.

As pointed out earlier, A. maculatum females may oviposit a

clutch that consists of several clusters. The mean cluster size reported

in Table 1 is lower than earlier reports of clutch size in A. maculatum

(Bishop 1941, Wilbur 1977, Pfingsten and Downs 1989). This suggests

that females of A. maculatum in the southern Appalachians may

oviposit clutches that consist of several clusters. Because interclutch

variation outweighs intraclutch variation in this species, the possibility

remains that several clusters might contribute to a single A. maculatum

clutch. This may confound the analysis of intraclutch variation, i.e.,

the intraclutch CVs for A. maculatum might be underestimates. However,

the pooled variation remains as an indicator of potential adaptive

variation in egg size at the interclutch level.

80 Christopher King Beachy

DISCUSSION

The potential role of variation in offspring size has long been

a topic of debate. Kaplan and Cooper (1984) suggested that egg size

variation in amphibians enables a female to produce viable offspring

in unpredictable environments. In addition, Kaplan and Cooper (1984)

submit that there are two levels at which variation in offspring size

may be introduced: within a single clutch and among successive clutches

of an individual female (i.e., at the intraclutch and interclutch levels).

I tested these hypotheses by comparing the amount of variation in

egg size observed in five species of salamanders that use larval

environments ranging (in terms of safety from desiccation) from

permanent to ephemeral.

These data add to the existing evidence showing that variation

in egg size in amphibians is extensive. In addition, it appears that

variation in offspring size differs among individual clutches and that

the degree of variation differs among species. The question to be

asked is whether the degree of variation is related to different life

history strategies employed by species using different environments.

Kaplan and Cooper (1984) proposed that variation in amphibian

egg size might reflect different reproductive strategies. In their attempt

to model parental investment, they included a consideration of the

extensive variation in propagule size seen in amphibians, insects,

and plants. Earlier models of parental investment lacked this aspect

(e.g., Smith and Fretwell 1974). A study by Crump (1981) emphasized

the potential role of variation in amphibian egg sizes. Crump found

no significant differences in egg size variation among species of

treefrogs that use habitats of differing variability. However, among

species that use temporary ponds, individual females produced clutches

that tended toward a platykurtic distribution of egg size (bet

hedging). Those species that breed in permanent ponds tended

toward a leptokurtic distribution (canalization).

However, Crump's evidence has been criticized as unconvincing

(McGinley et al. 1987). In a mathematical consideration of Smith and

Fretwell's and Kaplan and Cooper's models, McGinley et al. (1987)

suggested that variable environments do not necessarily select for

variable parental investment in offspring. Parental fitness can be maxi-

mized, even in heterogeneous environments, by investing equally in

all offspring. Is variation in egg size adaptive? Or are there factors

that prevent a female from investing equally in all offspring?

If one were to consider only the intraclutch variation that I

present here, the supposition that variation in egg size is correlated

with desiccation risk appears to garner little support. The data on

interclutch variation, however, suggest that species that use ephemeral

Salamander Egg Sizes 81

larval habitats might be able to introduce variation in egg size by

ovipositing successive clutches with different mean egg sizes. A.

maculatum did exhibit the largest degree of interclutch variation,

and the other species followed in nearly the predicted order.

Smith-Gill (1983) suggested that much adaptive variation can be

introduced at the whole organism level through developmental mechan-

isms, the mechanism in this case being vitellogenesis. Those develop-

mental mechanisms that provide for variation should be subject to

natural selection; i.e., those mechanisms should provide the amount of

variation that maximizes an individual's fitness. The variation in egg

size in the species examined provides some support for the hypothesis

that this variation is correlated with habitat variability and is possibly

adaptive. It remains to be seen if the hypothesis of adaptive variation

in egg size is supported when more species of amphibians are

investigated.

ACKNOWLEDGMENTS — This research represents a portion of

work submitted in partial fulfillment of the requirements for the

M.S. degree at Western Carolina University, Cullowhee, North

Carolina. R. C. Bruce, M. L. Crump, R. G. Jaeger, R. H. Kaplan,

C. L. Ory, and S. C. Walls reviewed versions of the manuscript. S.

G. Tilley kindly identified some of the clutches of Desmognathus. I

thank J. B. Bernardo, R. C. Bruce, and S. R. Voss for their help

in collecting clutches of eggs and for helpful criticisms during the

completion of this project. This work was supported by The

Highlands Biological Station and Louisiana Board of Regents

Doctoral Fellowship LEQSF (1988-1994)-GF-15.

LITERATURE CITED

Beachy, C. K. 1988. The correlation between egg size variation and

habitat variability in several species of salamanders: a test of the

adaptive "coin-flipping" hypothesis. M.S. Thesis, Western Carolina

University, Cullowhee, North Carolina.

Bishop, S. C. 1941. The salamanders of New York. New York State

Museum Bulletin 324:1-365.

Bruce, R. C. 1980. A model of the larval period of the salamander

Gyrinophilus porphyriticus based on size frequency distributions.

Herpetologica 36:78-86.

Bruce, R. C. 1988a. Life history variation in the salamander

Desmognathus quadramaculatus. Herpetologica 44:218-227.

Bruce, R. C. 1988b. An ecological life table for the salamander

Eurycea wilderae. Copeia 1988:15-26.

Bruce, R. C. 1989. Life history of the salamander Desmognathus

monticola, with a comparison of the larval periods of D. monticola

and D. ochrophaeus. Herpetologica 45:144-155.

82 Christopher King Beachy

Capinera, J. L. 1979. Qualitative variation in plants and insects: effect of

propagule size on ecological plasticity. American Naturalist 114:350-

361.

Crump, M. L. 1981. Variation in propagule size as a function of envi-

ronmental uncertainty for tree frogs. American Naturalist 117:724-737.

Crump, M. L. 1984. Intraclutch egg size variability in Hyla crucifer

(Anura: Hylidae). Copeia 1984:302-308.

Duellman, W. E., and L. Trueb. 1986. Biology of amphibians. McGraw-

Hill, New York, New York.

Kaplan, R. H. 1979. Ontogenetic variation in "ovum" size in two species

of Ambystoma. Copeia 1979:348-350.

Kaplan, R. H. 1980. The implications of ovum size variability for

offspring fitness and clutch size within several populations of

salamanders {Ambystoma). Evolution 34: 51-64.

Kaplan, R. H. 1985. Maternal influences on offspring in the California

newt, Taricha torosa. Copeia 1985:1028-1035.

Kaplan, R. H. 1987. Developmental plasticity and maternal effects of

reproductive characteristics in the frog, Bombina orientalis.

Oecologia 71:273-279.

Kaplan, R. H., and W. S. Cooper. 1984. The evolution of developmental

plasticity in reproductive characteristics: an application of the

"adaptive coin-flipping" principle. American Naturalist 123:671-689.

Lewontin, R. C. 1966. On the measurement of relative variability.

Systematic Zoology 15:141-142.

McGinley, M. A., D. H. Temme, and M. A. Geber. 1987. Parental

investment in variable environments: theoretical and empirical

considerations. American Naturalist 130:370-398.

Petranka, J. W. 1984. Incubation, larval growth, and embryonic and larval

survivorship of smallmouth salamanders {Ambystoma texanum) in

streams. Copeia 1984:862-868.

Pfingsten, R. A., and F. L. Downs, eds. 1989. Salamanders of Ohio. Ohio

Biological Survey Bulletin, New Series 7(2):1-315.

Shoop, C. R. 1974. Yearly variation in larval survival of Ambystoma

maculatum. Ecology 55:440-444.

Smith, C. C, and S. D. Fretwell. 1974. The optimal balance between size

and number of offspring. American Naturalist 108:499-506.

Smith-Gill, S. J. 1983. Developmental plasticity: developmental conver-

sion versus phenotypic modulation. American Zoologist 23:47-55.

Sokal, R. R., and F. J. Rohlf. 1981. Biometry. W.H. Freeman and

Company, San Francisco, California.

Wake, D. B. 1966. Comparative osteology and evolution of the lungless

salamanders, family Plethodontidae. Memoirs of the Southern

California Academy of Science 4:1-111.

Wilbur, H. M. 1977. Propagule size, number, and dispersion pattern in

Ambystoma and Asclepias. American Naturalist 111:43-68.

Accepted 23 March 1992

Helminth Parasites of the Eastern Box Turtle,

Terrapene Carolina Carolina (L.)

(Testudines: Emydidae), in North Carolina

Michael D. Stuart1 and Grover C. Miller

Department of Zoology, North Carolina State University

Raleigh, North Carolina 27695-7617

ABSTRACT — We examined 117 eastern box turtles, Terrapene Caro-

lina Carolina, for helminth parasites. Nine species (two trematodes,

six nematodes, and one acanthocephalan) were recovered, and 39%

of the turtles were infected with three-five species of parasites.

Infection rates were as follows: Oswaldocruzia sp. (82.9%), Spironoura

affinis (76.1%), Telorchis robustus (29.9%), Cosmocercoides dukae

(20.5%), Aplectana sp. (6.0%), Brachycoelium salamandrae (2.6%),

Physaloptera sp. (2.6%), Serpinema (=Camallanus) microcephalus (0.9%),

and Macracanthorhynchus ingens (0.9%). Ulcerations of the stomach

mucosa harbored larval Spironoura affinis. The presence of Spironoura

affinis, Telorchis robustus, and Serpinema microcephalus suggests a

close phylogenetic relationship of Terrapene to other emydid turtles.

The other helminth species are normally found in amphibians and

might represent parasites acquired in the turtle's evolutionary transi-

tion from an aquatic to a terrestrial lifestyle.

The box turtle, Terrapene Carolina (L.), is found throughout the

eastern United States. This small, terrestrial turtle has been studied

more thoroughly than most reptile species, perhaps because of its

ubiquity and innocuousness. The wealth of our knowledge on diet,

habitat preference, and behavior makes the box turtle an excellent

model for investigating parasite-host interactions (Stuart and Miller

1987).

This study was initiated to determine the following: (1) helminth

intensity and prevalence in box turtles in North Carolina; (2) correlations,

if any, of host age and sex with helminth intensity and prevalence;

(3) similarity of helminth fauna in host specimens from North Carolina

and elsewhere in the United States; and (4) helminth infection patterns

in relation to box turtle behavior and dietary habits.

MATERIALS AND METHODS

Turtles were collected from 13 North Carolina counties between

June 1982 and August 1989. Collecting was done primarily on the

Present address: Department of Biology, University of North Carolina at Asheville,

Asheville, NC 28804.

Brimleyana 18:83-98, June 1993 83

84 Michael D. Stuart and Grover C. Miller

Piedmont Plateau (n = 97), with small comparative samples taken

from the Blue Ridge Mountains (n = 16) and the Coastal Plain (n =

4). Ninety-seven turtles were collected either as roadkills or while

they were crossing highways. Twenty specimens were collected in the

field, in part with the aid of a border terrier dog trained to locate

turtles by scent. All turtles were sexed by secondary sexual characteristics

or grouped as juveniles if the plastron length was <100 mm. The

turtles were weighed to the nearest 0.1 g, measured both along the

straight length of the plastron and around the curve of the carapace,

and examined for helminth parasites. The entire visceral mass was

removed. The body cavity and each organ was examined separately.

The gastrointestinal tract was separated into distinct sections (esophagus,

stomach, intestines, and colon), and individual sections were cut length-

wise, washed, and examined separately. After removal of parasites

from the lumen, each section was scraped with a sharp blade. The

gut sections and the contents were digested in a pepsin-HCl solution

agitated constantly for 1 hour at 36C. The solution was decanted,

and the residue was examined for helminths with the aid of a stereoscopic

microscope.

Stomach ulcerations were removed and fixed in 70% ethyl alcohol,

5% formalin, or gluteraldehyde before sectioning for histological analysis.

Blood smears were strained with hemal blood film stain and examined

for the presence of microfilariae.

Helminths were considered prominent if the prevalence was >15%

and peripheral if the prevalence was <15%. We used an ANOVA to

test for significant differences in number of species of parasites between

sexes. A Kruskal-Willis test was used to test for significant differences

in number of individual parasites between sexes and geographical

regions of the state.

RESULTS

The box turtles we examined consisted of 43 males, 48 females,

and 26 juveniles. Stuart and Miller (1987) previously reported on

mass, sex and age structure, seasonal distribution, reproduction, and

food habits of 104 individuals from this collection. Of the 117 turtles,

3 were not infected, 27 were infected with 1 species of helminth

parasite, 41 were infected with 2 species, and 46 were infected with

>3 parasite species. The modal number of helminths was 22 with a

range of 0-303, exclusive of larval Spironoura affinis in stomach

ulcers. The helminth species we found and their prevalence and intensity

are shown in Table 1. No significant differences were found in helminth

species prevalence or intensity between host males, females, and juveniles.

Four helminth species exhibited a prominent infection rate of >15%:

Eastern Box Turtle Parasites

Table 1. Helminths collected from 117 turtles from North Carolina.

85

Oswaldocruzia sp. (82.9%), Spironoura affinis Leidy, 1856 (76.1%),

Telorchis robustus Goldberger, 1991 (29.9%), and Cosmocercoides dukae

(Holl, 1928) (20.5%). An additional five species were considered peripheral

with a prevalence of <15%. No extraintestinal helminths or microfilariae

were found.

We found a morphologically distinct and unnamed species of

Oswaldocruzia in the stomach of box turtles. Spicular morphology

differs substantially from O. pipiens Walton, 1929 from amphibian

hosts. Both the spicules and the dorsal ray of males are substantially

larger than those of O. pipiens, although the species in the box turtles

is smaller in all other respects. The prevalence of infection was 82.9%

(97/117), making this the most common helminth parasite encountered.

The mean intensity of infection was 7.3 worms per infected turtle

with no significant difference between any age or sex classes (P =

0.30).

Spironoura affinis infected 76.1% (89/117) with a mean intensity

of 32 nematodes per turtle. The range in intensity was 1-151. The

difference in intensity of infection with S. affinis was nearly significant

between adults and juveniles (P = .056); means for males were 20 ±

4.0 (SE), females 17 ± 4.6, and juveniles 34.7 ± 8.4. Pairwise contrasts

86 Michael D. Stuart and Grover C. Miller

using a Mann-Whitney U-test between males, females, and juveniles

indicated that the number of S. affinis in juveniles differed significantly

from that in females (P = 0.02) but that there was no significant

difference between males and females (P = 0.26), or between juveniles

and males (P = 0.13). Fifty-two (44%) of the 117 turtles had active

ulcer-like lesions in the fundal region of the stomach. Macroscopically,

the lesions or ulcers showed a raised area 1-2 cm in diameter with

a central opening 4-5 mm wide that extended into the stomach wall.

Microscopically, the lesions showed a moderate to dense lymphiod

infiltrate into the granulatomatous lining of the ulcer. When pressure

was applied to the base of these ulcers, masses of larval nematodes

were expressed. Comparison with adult and immature nematodes already

collected from the colon, particularly in regard to the shape of the

esophageal bulb and the developing lip structures, showed these larvae

to be Spironoura affinis. Many of the turtles had healed from previous

ulcers, which suggests that the damage is tolerated.

In our survey, 29.9% of the turtles (35/117) were parasitized by

Telorchis robustus, including 13 males, 15 females, and 7 juveniles.

The number of worms per turtle was substantial with a mean (and

range) of 62.2 (1-295) for males, 22.5 (1-93) for females, and 27.5

(1-80) for juvenile turtles. Telorchis robustus caused the only serious

health problem seen in our study. One turtle with 223 worms had a

partially telescoped intestine, apparently caused by the large worm

mass.

Two of the male turtles hosted nine and seven Brachycoelium

salamandrae (Froelich, 1789), respectively. One female harbored nine

worms. None was found in juvenile turtles. The total prevalence was

2.6% (3/117) with a mean of 8.3 worms per turtle.

Twenty-four of the box turtles were infected with Cosmocercoides

dukae (range = 1-241, x = 22.3). The genus Aplectana is closely

related to Cosmocercoides and is usually distinguished from the latter

by the absence of plectanes in the male. Both are normally parasites

of amphibians. Five (4.3%) turtles were infected with 1-15 sexually

mature nematodes, lacking plectanes on the males. Because of the

absence of plectanes, these worms were tentatively identified as Aplectana

sp., but additional work on the morphology and life cycle is needed

before a firm identification of the species can be made.

We also found two genera of spirurid nematodes. Serpinema

(-Camallanus) microcephalus (Dujardin, 1845) was in the stomach of

one turtle. This host was partially buried in the mud in a pool in an

intermittent stream. The turtle had possibly swallowed infected copepods.

Three turtle stomachs contained Physaloptera sp. Only two males were

recovered. Based on the small sample size, we could not determine

Eastern Box Turtle Parasites 87

whether these specimens were the species described from Terrapene

ornata by Hill (1941) as P. terrapenis Hill, 1941.

One turtle contained one immature specimen of the acanthocephalan

Macracanthorhynchus ingens (Linstow, 1879). This worm was not attached

to the stomach wall and was possibly a spurious parasite contracted

by the turtle having recently eaten an infected beetle.

DISCUSSION

The life histories of most of the parasites collected in this study

are poorly documented, and much of the literature is either contradictory

or limited in scope. In addition, major disagreement exists concerning

the appropriate nomenclature for many species or species complexes.

To help clarify existing information and to place our results in perspective

for future studies, a summary of nomenclatural problems and life

history data follows.

Platyhelminthes: Digenea

Brachycoelium salamandrae (Froelich, 1789) — Both Harwood (1932)

and Byrd (1937) described a number of species of Brachycoelium

from reptiles and amphibians in the southeastern United States. Rankin

(1938) reviewed the genus and concluded that the characters used to

describe the various species were too variable to be of specific diagnostic

value. Rankin (1945) also stated that the relative size of individual

flukes was dependent on the number of flukes infecting a particular

host. The worms were quite small when large numbers were present

but were substantially larger when ^ 20 were present. Rankin advocated

that Brachycoelium daviesi Harwood, 1932; B. dorsale Byrd, 1937; B.

georgianium Byrd, 1937; B. hospitale Stafford, 1900; B. louisianae

Byrd, 1937; B. meridionalis Harwood, 1932; B. mesorchium Byrd,

1937; B. obesum Nicoll, 1914; B. ovale Byrd, 1937; B. storeriae

Harwood, 1932; and B. trituri Holl, 1928 be reduced to synonyms of

B. salamandrae. Cheng (1958) disagreed, recognized all of the above

listed species, and described a new species, Brachycoelium elongatum

Cheng, 1958. Since that time, two additional species have been described:

B. stablefordi Cheng and Chase, 1961 and B. ambystomae Couch,

1966. All specimens found in the box turtles in our study were

identified with Cheng's keys as Brachycoelium salamandrae. Here, we

follow Rankin in treating this species complex as a single, extremely

variable species whose morphological features are influenced by numbers

and hosts.

Brachycoelium salamandrae has been reported from a wide range

of reptile and amphibian species. Rumbold (1928) reported B. salamandrae

as the only trematode he found in seven box turtles from North

Carolina, with an infection rate of 28% and an average of 0.28 worms

88 Michael D. Stuart and Grover C. Miller

per turtle. Raush (1947) examined 19 box turtles in Ohio and found

one turtle to host 27 specimens of B. salamandrae. Rankin (1945)

listed the species distribution as worldwide, but Yamaguti (1971) listed

only Palearctic and Nearctic hosts. Rankin (1945) noted a correlation

between the terrestrial habits of certain amphibian hosts and a high

level of prevalence, suggesting that terrestrial invertebrates were probably

involved in transmission. Denton (1962) reported snails and slugs,

Praticollela berlandieriana (Moricand), Derocerceras reticulatwn (=Agriolimax

agrestis) (Mier), and Mesodon thyroideus (Say) as suitable experimental

first intermediate hosts. Both motile and encysted cercariae were shed

in secreted mucus. Uninfected P. berlandieriana, D. reticulatum, Triodopsis

texasiana (=Polygyra texasiana) (Moricand), Anguispira alternata (Say),

and Bulimulus alternatus (Say) became infected within 2-10 days after

being exposed to infected first intermediate hosts, thus serving as

second intermediate hosts. Jordan (1963) and Jordan and Byrd (1967)

added Triodopsis caroliensis (Lea) and Mesodon inflectus (Say) to

the list of first intermediate hosts and T. caroliensis, M. inflectus,

Zonitoides aboreus (Say), Gastrocopta contracta (Say), Stenotrema

barbigerum (Redfield), Philomycus carolianus (Bosc), and Deroceras

laeve (Miiller) as second intermediate hosts. Cheng (1958) reported

development of nonencysted metacercariae in Ventridens ligera

{-Zonitoides ligerus) (Say). The definitive host presumably becomes

infected by consuming snails with encysted metacercariae, because both

Klimstra and Newsome (1960) and Stuart and Miller (1987) found

that gastropods comprise a large percentage of box turtle diets. Given

the broad range of first and second intermediate hosts that B.

salamandrae is capable of infecting and the high frequency of these

taxa in box turtle diets, it is surprising that the prevalence of infection

is so low.

Telorchis robustus Goldberger, 1911 — Wharton (1940) redefined

the genus Telorchis and its species. We used his species key to

identify Telorchis robustus from the box turtles in our study. Goldberger

(1911) described T. robustus from a box turtle collected in Maryland,

and Krull (1936) stated the trematode was common in Maryland box

turtles. Bennett and Sharp (1938) found T robustus in 38% (13/34)

of Terrapene c. triunguis (Agassiz) examined in Louisiana. The number

of worms ranged from three to nine, with an average of five worms

per turtle. They also reported T. robustus from 12% (8/65) Sternotherus

odoratus (Latreille) with an average infected of 10 worms per animal

and a range of 1-28. Rausch (1947) reported Telorchis sp. from 1 of

19 box turtles in Ohio and T. robustus in four of eight Clemmys

guttata (Schneider). The latter averaged two worms per turtle with a

maximum of four. Thirteen of 35 turtles in our study had ^25 worms

Eastern Box Turtle Parasites 89

in the small intestines, which is substantially more than found in

previous studies. The number of worms per turtle was substantial

with a mean (and range) of 62.2 (1-295) for males, 22.5(1-93) for

females, and 27.5 (1-80) for juvenile turtles. We do not know why

the mean number of this trematode is so much higher in North Carolina

box turtles than that reported from other localities or other species of

turtles.

Krull (1935, 1936) reported that Pseudosuccinea columella (Say)

became infected after eating trematode eggs (experimental infection)

and began to shed xiphidiocercariae within 28-32 days. Cercariae success-

fully penetrated and encysted as metacercariae in three snail species:

P. columella, Helisoma trivolvis (Say), and Lymnaea traskii (Lea).

Krull postulated that turtles were infected during the spring and early

summer when they ate snails in semi-flooded flats. He also noted

that metacercariae were never abundant, although snails had been

repeatedly exposed to thousands of cercariae. However, he did report

that one snail would occasionally acquire a much heavier infection

than others in the same group perhaps because they began feeding

more quickly than others. In light of more recent studies on host

immunity, genetic susceptibility might be a more reasonable cause

than a behavioral trait. In either case, these "super-infected" snails

will influence the range of worms in infected definitive hosts.

Nematoda

Oswaldocruzia sp. — Seven species of Oswaldocruzia have been

described from North American amphibians and reptiles (Baker 1977):

O. subauricularis Travassos, 1917; O. leidyi Travassos, 1917; O. pipiens

Walton, 1929; O. collaris Walton, 1929; O. waltoni Ingles, 1936; O.

euryceae Reiber, Byrd, and Parker, 1940; and O. minuta Walton,

1941. Oswaldocruzia subauricularis is a neotropical species and has

only been reported once in the United States. Baker (1977) redescribed

O. pipiens and regarded O. collaris and O. eurycea as synonyms of

O. pipiens. He also treated O. waltoni and O. minuta as species

inquirendae and O. leidyi as a nomen nudum. Accounts of develop-

mental and transmission patterns in Oswaldocruzia vary widely. Baer

(1952) stated that O. fillicollis (Goeze) (presumably referring to

Oswaldocruzia filiformis [Goeze, 1782] from amphibians molted twice

within the egg and was thus infective when the egg was consumed.

Baer also noted that L3 larvae might sometimes hatch and remain

ensheathed in the preceding molt until consumed. Baker (1978a) reported

that eggs from frogs and toads were laid in the 16-cell stage, and L}

larvae hatched within 24 hours of passage in the host's feces. Laboratory

cultured specimens developed to the ensheathed, infective L3 stage

within 3-4 days at room temperature. Anuran infection occurs via

90 Michael D. Stuart and Grover C. Miller

skin penetration. Larvae attached initially to the stomach mucosa but

migrated posteriorly to the intestine as they matured. Baker felt that

late summer and early fall, when marsh size was reduced and frog

density was highest, was the most important period for parasite

transmission and that O. pipiens could overwinter in its host. Hendrikx

(1981) studied the seasonal fluctuation of O. filliformis in Bufo bufo

L. in the Netherlands, and he reported L4 larvae embedded in the

stomach mucosa of overwintering hosts.

Oswaldocruzia specimens were only rarely found outside the stomach

in the box turtles of our study. An infective mode involving skin

penetration, while feasible in an amphibian, is somewhat more difficult

to accept in the heavily armored box turtle. While softer areas of

skin are found around the base of the legs and throat, chance nematode

access and penetration could scarcely account for the very high levels

of infection. The habits and habitats of the extremely wide range of

amphibian and reptile hosts of Oswaldocruzia provide additional reasons

to suspect an alternate route of infection. Storeria dekayi (Holbrook),

S. occipitomaculata (Storer), Anolis carolinensis (Voigt), many ranid

and bufonid species, Typholtriton spelaeus Stejneger, and Terrapene

Carolina use a broad spectrum of habitats including semi-fossorial,

arboreal, aquatic, semi-aquatic, cave-dwelling, and terrestrial. These habitat

differences alone would severely hamper transmission of a parasite

dependent on skin penetration to infect. Circumstantial evidence based

on host diversity and host diet suggests that Oswaldocruzia might use

an alternate life cycle with gastropods as intermediate hosts. In support

of this contention, we found a small male Oswaldocruzia completely

embedded in a piece of snail tissue taken from the stomach of a

freshly killed box turtle. While as yet unproven, morphological and

biological differences in the parasites and the different hosts suggest

that the species of Oswaldocruzia in box turtles is distinct from O.

pipiens.

Spironoura affinis Leidy, 1856 — Leidy (1856) erected the genus

Spironoura and listed two species: S. gracile from the stomach of

the red-bellied turtle, Pseudemys rubriventris (Le Conte) (=Emys serata)

and S. affine, later modified to S. affinis by Yamaguti (1961), from

the cecum of the box turtle, Terrapene Carolina {-Cistudo Carolina).

Yorke and Maplestone (1926) designated S. gracile as the type species

apparently because it appeared first in Leidy's manuscript. This species

has not, however, been collected since its description. Freitas and

Lent (1942) felt that S. gracile should be considered a species inquirendum

since Leidy's description was brief and incomplete. They proposed

revalidation of the genus Falcaustra Lane, 1915 with all of the species

of Spironoura transferred to this genus. Some authorities (Yamaguti

Eastern Box Turtle Parasites 91

1961, Skrjabin et al. 1964) rejected this, but Chabaud (1978) considered

Falcaustra to be the valid genus for this group. We follow Chapin

(1924) in rejecting Falcaustra and continue to use Spironoura because

Leidy's original description, although brief by today's standards, is

sufficient to distinguish the genus, giving Spironoura priority.

About 50 species of Spironoura have been described from fishes,

reptiles, and amphibians worldwide (Skrjabin et al. 1964). Mackin

(1936) published a thorough study of the anatomy of the genus Spironoura

and a key to the species from the United States. Five species of

Spironoura have been reported from Terrapene Carolina in the United

States: Spironoura affinis Leidy, 1856; S. longispicula (Walton, 1927);

S. cryptobranchi Walton, 1930; S. chelydrae (Harwood, 1930); and S.

concinnae Mackin, 1936. Canavan (1929) described S. procera from

the same host, Pseudemys rubriventris (=Emys serata), and from the

same locality where Leidy worked, i.e., Philadelphia. Harwood (1930)

is the only other investigator to have reported the presence of Spironoura

procera, but he said that it was not sufficiently distinct from S.

affinis to merit specific status. Harwood's (1932) work, plus our own

study of S. affinis and examination of the specimen from the U.S.

National Museum Helminthological Collection, Beltsville, Maryland

marked "Spironoura procera? (No. 52145)," suggest that S. affinis, S.

procera, and S. gracile are all members of the same species. We

also borrowed specimens identified as S. concinnae, collected from

Terrapene Carolina in Mississippi, from the U.S. National Museum

Helminthological Collection (No. 66152). Comparison with specimens

of S. affinis collected from the same host in North Carolina convinced

us that Caballero (1939) was correct in considering S. concinnae as a

synonym of S. affinis. We have followed Yamaguti's (1961) designation

of Spironoura affinis to minimize confusion. However, this diverse

and complex genus needs revision.

Cosmocercoides dukae (Holl, 1928) — Cosmocercoides dukae is a

common parasite of amphibians. Holl (1928) originally described C.

dukae (=Cosmocerca dukae) from a newt collected in Durham, North

Carolina. Harwood (1932), apparently unaware of Holl's work, described

the same species as Oxysomatium variablis and listed 10 species of

amphibian and reptile hosts. Harwood's experimental attempts to

demonstrate host infection by skin penetration were not successful,

but he felt that a direct life cycle was probable. Ogren (1953, 1959)

demonstrated the nematode's ability to complete its life cycle in a

variety of gastropod species, including Ashmunella rhyssa (Dall), Triodopsis

(=Polygyra) fosteri (Baker), Retinella sp. Fischer, and Deroceras sp.

Rafinesque. Anderson (1960) described the life cycle of C. dukae in

Discus cronkhitei (Newcombe), Zonitoides aboreus (Say), and Deroceras

92

Michael D. Stuart and Grover C. Miller

Table 2. Helminths reported from Terrapene Carolina.

Eastern Box Turtle Parasites

93

Table 2. Continued.

94 Michael D. Stuart and Grover C. Miller

Table 2. Continued.

Number

Infected/Examined Locality Reference

Physalopteridae

Physaloptera terrapenis Hill,

1941 7/47 in Ok. Hill (1941)

T. ornata

Spiruridae

Spiroxys constricta

(Leidy, 1856) 7/16 La. Acholonu and

Amy (1970)

Penn.? Leidy (1856)

S. contorta (Rudolphi, 1819) 11/63 111. Martin (1973)

Gnathostomatidae

Gnathostoma procyonis

Chandler, 1942 3/4 La. Ash (1962)

laeve (Miiller) (=D. gracile Rafinesque) and suggested that amphibian infections

occurred from ingestion of infected molluscs. Baker (19786) reported that

C. dukae larvae burrow through the skin of toads and migrate through the

body. This versatility in ability to use such different definitive hosts suggested

to Baker that C. dukae represented an early stage of parasitic adaptation.

Parasites and Host Diet

Surveys from various areas within the range of Terrapene Carolina

indicate that a broad range of helminth parasites infect the box turtle

(Table 2). In North Carolina, parasite presence appears to be regulated

primarily by diet. Recognizable items found in the gastrointestinal

tract of 72 turtles, in order of frequency of occurrence, were snails

(Gastropoda) - 59%, insects (Insecta) - 43%, sowbugs (Isopoda) -

40%, plant material (primarily fungi) - 32%, slugs (Gastropoda) -

7%, rodents (Mammalia) - 5.5%, earthworms (Oligochaeta) - 3%, and

millipeds (Diplopoda) - 3%. Cosmocercoides dukae, Brachycoelium spp.,

and Telorchis robustus have all been shown to use molluscan intermedi-

ate hosts (Krull 1935, 1936; Ogren 1953; Jordan and Byrd 1967).

Our evidence strongly suggests that Spironora affinis and Oswaldocruzia

sp. also might be capable of using molluscs to reach the definitive

host. The records for Serpinema microcephalia and Physaloptera sp.

probably represent rare or accidental infections. The life cycle of most

physalopterans is unknown; those that are known use invertebrate

intermediate hosts. The specimen of Macroacanthorthynchus ingens

indicates recent ingestion of a scarabid beetle. Considering the relatively

high incidence of insects and sowbugs in the diet and the semi-

aquatic nature of the turtles, the absence of acanthocephalans is some-

Eastern Box Turtle Parasites 95

what surprising. The poisonous fungi in the diet could conceivably

act as a periodic vermifuge, but that has not been investigated.

Parasite life cycles are often complex and may involve a variety

of strategies to get the parasite into the definitive host, including one

or more intermediate hosts. Successful transmission requires a congruence

between parasite life cycle and host behavior or ecology. Aho (1990)

discussed the importance of reptile- and amphibian-parasite systems in

understanding the ecological and evolutionary relationships which determine

parasite species distribution. The parasite presence/absence data accumu-

lated in our study suggest that Spironoura affinis has a long history

or relationship with Terrapene Carolina and the aquatic emydid turtles

from which the box turtle evolved. Nursery ulcers and molluscan

intermediate hosts could represent the nematode's adaptation to the

host's move from an aquatic to a terretrial habitat. The presence/

absence data also suggest that Terrapene Carolina acquired a number

of amphibian parasites in the ecological shift from water to land.

ACKNOWLEDGMENTS— This study was supported in part by

Science and Engineering Development Award 89SE01 from the North

Carolina Board of Science and Technology. We are indebted to J.

Ralph Lichtenfels for the loan of specimens from the U.S. National

Museum Helminthological Collection. We further thank James W. Petranka

for assistance with statistical analyses, John G. Petranka for updating

the molluscan nomenclature, and to Joyce McHenry, Jeanne Simmons,

and Kathy Haley for assistance in collecting specimens.

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96 Michael D. Stuart and Grover C. Miller

Ash, L. R. 1962. Development of Gnathostoma procyonis Chandler, 1942

in the first and second intermediate hosts. Journal of Parasitology

48:298-305.

Baer, J. G. 1952. Ecology of animal parasites. University of Illinois Press,

Urbana.

Baker, M. R. 1977. Redescription of Oswaldocruzia pipiens Walton, 1929

(Nematoda: Trichostrongylidae) from amphibians of eastern North America.

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Baker, M. R. 1978a. Development and transmission of Oswaldocruzia pipiens

Walton, 1929 (Nematode: Trichostrongylidae) in amphibians. Canadian

Journal of Zoology 56:1026-1031.

Baker, M. R. 19786. Transmission of Cosmocercoides dukae (Nematoda:

Cosmocercoidea) to amphibians. Journal of Parasitology 64:765-766.

Bennett, H. J., and C. H. Sharp. 1938. Helminth parasites of Sternotherus

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of the Louisiana Academy of Sciences 4:241-242.

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Caballero, E. 1939. Nematodos de los reptiles de Mexico. V. Anales del

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Canavana, W. P. N. 1929. Nematode parasites of vertebrates in the Phila-

delphia Zoological Garden and vicinity. Parasitology 21:63-102.

Chabaud, A. G. 1978. Keys to genera of the superfamilies Cosmocercoidea,

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Farnham Royal, Bucks, England.

Chapin, E. A. 1924. Note on Spironoura affine Leidy, 1856. Journal of

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Accepted 4 May 1992

Notes on the Spiny Softshell, Apalone spinifera

(Testudines: Trionychidae), in Southeastern Virginia

Joseph C. Mitchell

Department of Biology University of Richmond

Richmond, Virginia 231 73

AND

Ronald Southwick1

Virginia Department of Game and Inland Fisheries

6530 Indian River Road

Virginia Beach, Virginia 23464

ABSTRACT— The Gulf Coast spiny softshell turtle (Apalone spinifera

aspera), collected in Lake Whitehurst, Norfolk, Virginia, represents

the first record of this subspecies for the state. The eastern spiny

softshell (Apalone spinifera spinifera) occurs naturally in the Ten-

nessee and Ohio drainages in southwestern Virginia. Based on avail-

able evidence, the Norfolk population and a population of the nomi-

nate subspecies in Isle of Wight County, Virginia, also reported

here, should be considered isolated, introduced, and established

populations.

Apalone spinfera is a wide-ranging species in North America

occurring from the Rio Grande River northeastward through the midwest

to the Great Lakes region and eastward through the Carolinas to the

Atlantic Ocean (Ernst and Barbour 1972, Conant and Collins 1991).

Seven subspecies are recognized (Ernst and Barbour 1989, Iverson

1992), of which one, Apalone spinifera spinifera, occurs naturally in

the Tennessee River drainage in southwestern Virginia (Tobey 1985).

Recent discoveries of populations of this species in southeastern Virginia

raise questions about the occurrence of a second subspecies in the

state and demonstrate that introduced populations can become esta-

blished in this area.

On 25 June 1991, a large female A. spinifera spinifera (370

mm carapace length [CL], 283 mm plastron length [PL], and 5.5 kg

body mass) was discovered adjacent to a commercial fish rearing pond,

4.8 km east of Windsor, Isle of Wight County, Virginia; she was

released. Subsequent observations (5 July and 24 October 1991) and

captures (9 November 1991) revealed at least four other adults (a

male 262 mm CL, 184 mm PL, 1,550 g, The Living Museum, Newport

News, Virginia) and one juvenile (111 mm CL, 85 mm PL, National

'Present address: Virginia Department of Game and Inland Fisheries, 2006 South

Main Street, Suite C, Blackburg, VA 24060.

Brimleyana 18:99-102, June 1993 99

100 Joseph C. Mitchell and Ronald Southwick

Museum of Natural History, USNM 314209) in several rearing ponds

on the same site. The hatchery owner could not verify time or source

(e.g., shipments of commercial fish) of the turtles. He noted that they

have been present for several years.

The discovery of another population in southeastern Virginia is

comparatively more perplexing. In October 1988, an unidentified soft-

shell turtle was observed in Lake Whitehurst, City of Norfolk, Virginia

(T. Pitchford, personal communication, Virginia Beach Marine Science

Museum). Softshells were confirmed in this lake on 2 August 1991

when an adult male A. spinifera aspera (196 mm CL, 134 mm PL,

574 g body mass, USNM 314210) was collected on hook and line. A

large female was taken by another fisherman between 13 August and

1 September 1991 but it escaped, and its identity cannot be confirmed.

Three juveniles found beneath a boat dock on Lake Whitehurst near

Shore Drive on 21 September 1991 were given to the junior author

by a local boy who said that the turtles were coming out of the

sand. The juveniles averaged 51.0 + 2.0 (SD) mm CL (range = 49-

53), 35 ± 1.0 mm PL (range = 34-36), and 13 g (n = 1) body

mass. A recent hatchling (38.9 mm CL, 26.6 mm PL, 6 g) was

caught in the same area on 12 October 1991. Softshells apparently do

not overwinter in the nest in this area, although they may in the

upper midwest (Gibbons and Nelson 1978).

The nearest population of A. spinifer aspera is in Harnett County,

North Carolina, 290 km southwest of Norfolk (A. S. Braswell, personal

communication, North Carolina Museum of Natural Sciences). Our Lake

Whitehurst records establish A. s. aspera for the first time in Virginia

and indicate a substantial distributional hiatus between south-central

North Carolina and southeastern Virginia.

Is the Norfolk population introduced or is it a heretofore undocu-

mented natural population? Intensive sampling with large nets during

1977-91 by the Virginia Department of Game and Inland Fisheries

revealed no softshells. However, most sampling occurred in early spring

(March to early April) before most freshwater turtles and presumably

softshells (see Robinson and Murphy 1978) become active in this area

(J. C. Mitchell, unpublished data). This particular sampling effort yielded

few turtles of any species (R. Southwick, personal observation). Con-

siderable effort has been expended in North Carolina and Virginia to

determine the distributions of reptilian species within these states (W.

M. Palmer and A. L. Braswell, personal communications; J. C. Mitchell

and C. A. Pague, unpublished data). In addition, the area of southeastern

Virginia and northeastern North Carolina has been a favorite collecting

area for decades (e.g., Nemuras 1964; W. M. Palmer and R. de Rageot,

personal communication). If present earlier, softshells should have been

Spiny Softshell 101

found. Thus, the likelihood of intentional release of pet trade softshells

into Lake Whitehurst cannot be discounted. A presumably introduced

specimen of A. s. spinifera (George Mason University Collection, GMU

1676) was found in Bull Run Creek, Fairfax County, Virginia in July

1982.

A population of Apalone spinifera became established after 1910

in the Maurice River system in southern New Jersey (Conant 1961)

and apparently continues to persist (Conant and Collins 1991). Populations

in the Colorado River and several aquatic systems in California are

also considered introduced (Linsdale and Gressitt 1937, Webb 1962,

Stebbins 1985). These introductions indicate that the spiny softshell

can survive in areas outside of its natural range and establish viable

populations. Thus, until additional populations of A. s. aspera are

discovered in southeastern Virginia and northeastern North Carolina,

the Lake Whitehurst population should be considered an introduced

population.

ACKNOWLEDGMENTS— We thank Joe St. Martin and Andrew

Castellano for bringing the Lake Whitehurst softshells to our attention,

Tom Pitchford for his initial observation of a softshell in Lake Whitehurst,

and Sam Perry, of Perry's Minnow Farm, for allowing us access to

his property to study turtles. Alvin Braswell of the North Carolina

State Museum of Natural Sciences kindly provided locations of softshell

populations in North Carolina. Carl H. Ernst allowed access to the

George Mason University vertebrate collection.

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Conant, R. 1961. The softshell turtle, Trionyx spinifer, introduced and

established in New Jersey. Copeia 1961:355-356.

Conant, R., and J. T. Collins. 1991. Reptiles and amphibians of eastern

and central North America. Third Edition, Houghton Mifflin Company,

Boston, Massachusetts.

Ernst, C. H., and R. W. Barbour. 1972. Turtles of the United States.

Universtiy of Kentucky Press, Lexington.

Ernst, C. H., and R. W. Barbour. 1989. Turtles of the world. Smithsonian

Institution Press, Washington, D.C.

Gibbons, J. W., and D. H. Nelson. 1978. The evolutionary significance of

delayed emergence from the nest by hatchling turtles. Evolution 32:297-

303.

Iverson, J. B. 1992. A revised checklist with distribution maps of the

turtles of the world. Privately Printed, Richmond, Indiana.

Linsdale, J. M., and J. L. Gressitt. 1937. Soft-shelled turtles in the Colorado

River basin. Copeia 1937:222-225.

102 Joseph C. Mitchell and Ronald Southwick

Nemuras, K. T. 1964. Collecting notes — southeastern Coastal Plain, Virginia.

Bulletin of the Philadelphia Herpetological Society 12:35-36.

Robinson, K. M., and G. G. Murphy. 1978. The reproductive cycle of the

eastern spiny softshell {Trionyx spiniferus spiniferus). Herpetologica 34:137-

140.

Stebbins, R. C. 1985. A field guide to western reptiles and amphibians.

Houghton Mifflin Company, Boston, Massachusetts.

Tobey, F. J. 1985. Virginia's amphibians and reptiles, a distributional

survey. Virginia Herpetological Society, Purcellville, Virginia.

Webb, R. G. 1962. North American soft-shelled turtles (family Trionychidae).

University of Kansas Publications, Museum of Natural History 13:429-

611.

Accepted 15 May 1992

Range Expansion of the Tree Swallow,

Tachycineta bicolor (Passeriformes: Hirundinidae),

in the Southeastern United States

David S. Lee

North Carolina State Museum of Natural Sciences

P.O. Box 27647

Raleigh, North Carolina 27611

ABSTRACT— Since the late 1800s and early part of this century

when the tree swallow, Tachycineta bicolor, was a peripheral and

sporadic breeding species in the southeastern United States, its range

has expanded considerably. The precise reasons for the range expan-

sion of this and other swallows in the Southeast is unclear. Land

clearing, impoundments and other land use patterns, the re-introduc-

tion of beavers {Castor canadensis), and the use of bluebird (Sialia

sialis) boxes by swallows as nest sites appear to have facilitated

the expansion. Several different corridors of dispersal are noted; North

Carolina represents the current frontier of range expansion in the

Southeast.

The 1957 edition of the American Ornithologists' Union Check-

list (AOU 1957) stated that the breeding range of the tree swallow

(now Tachycineta bicolor) extended south to northwestern Tennessee,

northern West Virginia, Virginia, central Maryland, and northeastern

Pennsylvania. At the turn of the century the species rarely nested as

far south as southwestern Kentucky (Fig. 1). The breeding range has

expanded considerably since the 1950s, but this change in distribution

is not well documented. By the early 1980s the southern limits of

distribution were defined as northeastern Louisiana, westcentral Missis-

sippi, Tennessee, and North Carolina, but the tree swallow was

"generally sporadic or irregular as a breeder east of the Rocky Mountain

states and south of the upper Mississippi and Ohio Valleys, or along

the Atlantic coast south of Massachusetts" (AOU 1983).

At least in parts of the Southeast, tree swallows are currently

undergoing a rapid range expansion and indications are that they will

become common and widespread throughout much of the area during

the coming decades. This pattern of range expansion has already been

exhibited by several other species of swallows in the Southeast. The

majority of the information presented here is from North Carolina,

which is the current frontier of range expansion in the Southeast.

Brimleyana 18:103-113, June 1993 103

104

David S. Lee

Fig. 1. Early isolated breeding records of tree swallows {Tachycineta bicolor)

in the southeastern United States. See text for documentation sources.

METHODS

Information presented here is compiled from numerous published

sources such as state bird journals and from unpublished records

obtained from individuals and agencies monitoring bird distribu-

tions in the southeastern United States.

RESULTS

West Virginia — Hall (1983) notes that most West Virginia summer

records are from the mountains where nests have been found near

beaver ponds and other flooded sites. Definite nest records were given

for Hampshire, Preston, Tucker, Randolph, and Pocahontas counties.

Hall (1983) also reported nesting records outside the mountains from

the McClintic Wildlife Station, Mason County (several occasions), at

Belle, Kanawha County (1961), and Randolph County (1975). Addi-

tionally, there are summer sightings for Jefferson, Morgan, Pendle-

ton, Greenbrier, Mercer, Wood, and Gilmer counties in West Virginia.

These records indicate a considerable expansion since the 1957 AOU

Checklist.

Maryland — Stewart and Robbins (1958) reported tree swallows

as breeding commonly in the tidewater region of the southern

Delmarva and as being uncommon or rare breeders in other tide-

Tree Swallow Range Expansion 105

water areas of Maryland's Coastal Plain. This species was first reported

as breeding in this area in 1929 and 1933 (unpublished egg catalogue

of E. J. Court; Court 1936), but based on Virginia records (see

below) it was probably present some time before Court's report. A

disjunct portion of the breeding range was also noted in the Allegheny

Mountains (Garrett County) with five specific nesting localities listed

(Stewart and Robbins 1958). Despite a moderate amount of fieldwork

in western Maryland early in this century, the first report of mountain

nesting was not until 1920 (Eifrig 1920) with subsequent records in

1936 (Brooks 1936). The earliest documented nesting in Maryland

was 1894 (W. H. Fisher; in Stewart and Robbins 1958). Although it

is not known when tree swallows first started nesting in Maryland, it

was likely just before the turn of the century. The 1894 record is

for Baltimore County and is probably from Dulaney Valley north of

Towson, where Fisher did most of his fieldwork (Lee 1988). Interesting-

ly, this area was not considered part of the tree swallow's range in

1958. In 1968 the distribution was essentially unchanged (Robbins

and Van Velzen 1968). Since then, however, the species has expanded

throughout the state. It is still most numerous in the mountains and

in tidewater areas. Tree swallows are now locally common nesters in

the Maryland Piedmont (Maryland Breeding Bird Atlas data). State-

wide they were found in 393 of the 1,256 Atlas blocks (31.2%), and

they were found in every county in the state (C. S. Robbins, personal

communication). Colonization of most areas in Maryland seems to be

dependent on the species ability to nest in bluebird boxes.

Kentucky — Despite the state's location in relation to the historic

range of tree swallows (southern Illinois 1889, Missouri 1894, and

Tennessee 1918), the species was essentially unknown as a breeding

bird in Kentucky even by the mid-1960s (Mengel 1965). The only

record for the state before the 1960s is from 1889 to 1925 when

Pindar noted that tree swallows were present in Fulton County in the

summer. This county is in the extreme western tip of the state along

the Mississippi River and just north of Reelfoot Lake, the site of the

first record for Tennessee (Pindar 1889, 1925). Mengel (1965) visited

this area between 1941 and 1951 and, although he noted favorable

habitat, he did not locate nesting birds. Breeding was again noted in

1905 in the Cumberland River Valley (Lyon County at Long Run

Park in Jefferson County) in 1975 (Monroe et al. 1988). The species

is now recorded as nesting in scattered localities throughout the state.

Its nesting is mostly along edges of lakes and rivers where there are

many dead trees, and recently the species has been found using nest

boxes (Monroe et al. 1988).

106 David S. Lee

Virginia — In the early part of this century, Bailey (1913) recorded

the tree swallow as nesting only on the lower Delmarva portion of

Virginia where it bred mostly on islands. Murray (1952) summarized

the historical nesting status in Virginia as follows: "There is one

nesting record for Princess Ann County, June 15, 1927, upper end of

Back Bay, Lewis; and one for Aylett, King William County, no date

(Auk 14, 408). Palmer found a nest with young on Smith Island,

June 10, 1897; and reports one with eggs, May 1894 (Auk 14, 408).

Brooks states that it is a 'fairly common summer resident at the

higher elevations at least in Highland County (Raven 6, 11-12, 2).' "

By the late 1970s Larner et al. (1979) noted that the tree swallow

was locally common in the Virginia portion of the Delmarva Peninsula

and rare elsewhere in the Coastal Plain. In the Piedmont it was a

rare and local summer resident, and the first breeding record "in

recent years" was recorded in Madison County in 1976. Larner et al.

(1979) reported records in the mountains from Augusta, Highland,

Russell, and Tazewell counties. In the last decade this species has

greatly expanded its range in Virginia with confirmed nesting records

in 34 counties (65+ sites) throughout the state and expected or

presumed nesting in at least 18 additional counties (Sue Ridd, per-

sonal communication 1988; Virginia Breeding Bird Atlas).

Tennessee — The first nest was discovered at Reelfoot Lake,

Tennessee, on 22 May 1918 (Ganier 1964) but from that time until

1968 no additional nests were reported; there was only one other

observation of these swallows during the breeding season. Olson (1968)

found an active nest at Norris Lake in Anderson County in 1968,

and the same year Gray (1968) found a nest at Monsanto Ponds in

Maury County. The early records were from the western part of the

state adjacent to the Mississippi River. Since 1968 nests have been

reported in Tennessee almost annually. Nicholson and Pitts (1982)

summarized the distribution of nesting in Tennessee, noting that in

recent years tree swallows have nested throughout the state.

North Carolina — In North Carolina tree swallows were first report-

ed nesting in 1979 in the extreme northwestern corner of the state

(LeGrand and Potter 1980). The nest was in an abandoned wood-

pecker cavity along the New River in Ashe County (elevation 9,100

m). The second record was nearly 192 km southeast of this site, and

2.4 km north of Asheville in Buncombe County near the French

Broad River (elevation 600 m) in 1981 (Duyck 1981). In the subse-

quent decade the breeding range has expanded considerably.

Breeding Bird Atlas volunteers in North Carolina found tree

swallows nesting in the mountains in the southwestern part of the

state in Transylvania County (four locations) and in Henderson County.

Tree Swallow Range Expansion 107

Nesting was recorded in 1988 and 1989 and will probably continue

in the future. In 1988 and 1989 a pair nested in a purple martin

(Progne subis) gourd in Cowee Valley, Macon County, and in 1990

a pair nested in a bluebird box near Piney Creek, Alleghany County

(Chat 55:64-65). The Piedmont birds were seen nesting in woodpecker

cavities in trees killed during flooding to create Jordan Lake in

Chatham County in 1988. A nesting pair was found in Vance County

in 1990. To date these are the only confirmed nesting birds in the

Piedmont. In the Coastal Plain, tree swallows were found nesting in

eight locations in Currituck County (1989-90), and were seen during

the nesting season in Pamlico County (1988) and along the lower

Cape Fear River in Brunswick County (1990). In the Coastal Plain

pairs of tree swallows used bluebird boxes and natural cavities in

trees killed by impoundment and where natural flooding occurred

along rivers and in salt marshes.

DISCUSSION

Although present day land-use patterns provide fields and other

open areas suitable for swallow foraging, this alone does not account

for the current, explosive range expansion because land clearing was

widespread in the Southeast by the early 1800s. Nest sites are critical,

and the same land-clearing patterns that provide open areas can also

eliminate snags and other potential cavity-nest sites. The elimination

of beavers from most of the Southeast at the turn of the century

eliminated the potential for natural snags in areas impounded by beavers.

Dead trees resulting from flooding in tidewater areas and adjacent

salt marshes seem to have provided a natural dispersal route in coastal

areas. Therefore it is interesting, and not easy to explain, that, despite

the availability of natural habitat, tree swallows have only recently

(since 1913 in Virginia; late 1980s in North Carolina) expanded in

, tidewater areas. It is to be expected that coastal tree swallows will

also follow river systems inland and make use of dead trees in im-

poundments and beaver ponds in the Coastal Plain.

Phenology — Tree swallows are relatively early to late spring and

early to late fall migrants, making it difficult to distinguish resident

breeding birds from migrants. In Currituck County, North Carolina, in

June 1990 I observed resident birds using cavities before, during, and

after a fairly extensive northern migration of large numbers of transients.

Nicholson and Pitts (1982) noted tree swallows at Reelfoot Lake,

Tennessee, by mid-March, and territorial birds by late April when

northbound migrants were still present. Southward migration in the

southeastern United States begins in early July (Nicholson and Pitts

108 David S. Lee

1982, personal observations). Major fall migration occurs in October,

and birds winter in much of the Southeast, particularly along coastal

areas.

Nesting dates in the Southeast range from 12 May to 3 July

with no indication that pairs at southern latitudes nest any earlier

than those to the north. The earliest recorded egg date, for example,

is from Maryland (see below). Birds reported in from North Carolina

(NCSM records) nested from early May through mid-June. The following

is a list of nesting dates for the Southeast: 4 May-21 June, nesting

activity (15-20 May eggs laid), North Carolina (Duyck 1981); 6 June,

adult feeds fledglings, Tennessee (Williams 1976); 9 June, adult feed-

ing young, Tennessee (Nicholson and Pitts 1982); 16 June, adult and

fledged young, Tennessee (Nicholson and Pitts 1982); 9 June, adult

at nest, North Carolina (Chat 44:9); 11 June, young birds being

fed, North Carolina (Chat 44:9); early July, feeding young, Tennessee

(Nicholson and Pitts 1982); early May to mid-July (egg dates 24

April-5 July, nestling 20 May-27 July), Maryland (based on 320 Mary-

land nest records, C. S. Robbins, personal communication).

Pattern of Range Expansion — Apparently the expansion of nesting

into the outer Coastal Plain of Maryland, Virginia, and North Carolina,

the southern Appalachians, Maryland, to North Carolina, and the Pied-

mont regions of these states occurred independently (Fig. 2). Sites

along major rivers and those adjacent to cleared agricultural areas

were the first to be colonized (i.e., 1889-1918 Mississippi River low-

lands). Colonization occurred rather rapidly in the mountains (1920-36

Maryland, by 1929 in Virginia, and northern West Virginia pre-1957),

and accelerated in the last decade (northern North Carolina in 1979 to

southwestern North Carolina by 1988). Many of these sites were along

rivers in the Mississippi basin.

In the Coastal Plain the species was nesting on the Delmarva

as early as 1897, but was not common or widespread even by the

1950s and did not become so until the 1970s. Breeding individ-

uals did not invade northeastern North Carolina until the late 1980s.

Their occurrence in the Piedmont seems sporadic. The earliest

Piedmont record from the Southeast and Atlantic states is 1894 (Mary-

land), but the birds did not become established. Piedmont nesting was

not documented until 1961 (West Virginia) and 1976 (Virginia) and

was not widespread until the 1980s (Maryland and Virginia). This

swallow is still uncommon the North Carolina Piedmont where it is

known from only two sites. West of the Appalachians, this species

had a similar history in Tennessee where the first nest was reported

in 1918 with no new records until the second half of the 20th Century.

Yet the species was nesting throughout Tennessee by 1982.

Tree Swallow Range Expansion

109

I960

1980 -;

Fig. 2. The expanding breeding distribution of the tree swallow

(Tachycineta bicolor) in the southeastern United States (1894-1991).

Possible Explanation for Range Expansion — Tree swallows might

now be imprinted on bluebird boxes as nesting sites, and this imprint-

ing possibly has allowed them to expand their range into areas where

the lack of natural cavities would otherwise inhibit nesting.

Plasticity in food habits may also be important in the range

expansion of this swallow. Lee and Franz (1972) reported on a pre-

migration staging flock of several thousand tree swallows feeding over

a cornfield on the eastern shore of Maryland. Examination of the

stomachs of a dozen birds revealed corn flea beetles {Chaetocnema

pulicaria). No other food items were found. Food items were counted

in one stomach, and 170 corn flea beetles were found. Although I

am not aware of other direct observations of tree swallows benefiting

from agriculture or agricultural pests, at the minimum the clearing of

land for agricultural purposes has created foraging areas not available

in the precolonial period.

Several nesting sites are associated with man-made lakes and

large impoundments constructed for waterfowl. Many hardwood trees

killed by floodwaters contain cavities made by woodpeckers that provide

multiple nest sites where the swallows form small colonies of a dozen

or more pairs. Although the beaver was reintroduced by various state

wildlife agencies and has made an explosive come-back, this generally

precedes the period of range expansion of tree swallows outlined

110 David S. Lee

here (see Bonwil and Owens 1939, Mansuetti 1950, Lee 1988, and

Lee et al. 1982).

Similar Range Expansion of Other Species of Swallows — The

relatively rapid range expansion parallels the changing distribution of

other swallows in the Southeast. The barn swallow's (Hirundo rustica)

breeding range was largely to the north of North Carolina before

1942 (Pearson et al. 1942), but it bred locally in the mountains in

the northwestern corner of the state and along the coast. The range

expansion in the state was undocumented. By 1975 the species was

colonizing the outer Coastal Plain in southeastern North Carolina, and

today it occurs statewide (personal observation). Although they nest

in barns and under docks, the main factor allowing dispersal seems

to be the replacement of wooden bridges with concrete ones through-

out the state (1960s-70s); the swallows use the concrete bridges for

nest substrate.

At the beginning of this century cliff swallows (Hirundo

pyrrhonota) were only transients in North Carolina. In 1967 they

were reported nesting in the state, and by 1983 they had expanded

their range in the Piedmont to Greensboro, Guilford County (Hen-

drickson 1984). They had previously been reported nesting at other

Piedmont sites, all around reservoirs (Lake Cammack, Hyco Reservoir,

McGehee's Mill, and Jordan Lake). They also nest at Falls Lake,

Wake County (NCSM records, 1989). Like tree swallows, this species

did not simply expand its range from north to south or from the

west to the east as one might expect. They colonized scattered sites

in the Piedmont and later colonized suitable adjacent sites. McConnell

(1981) first reported nesting in the Mountains of North Carolina and

noted a preference for reservoir dams as nesting sites. This range

expansion, which started in the mid-1960s, also includes the Piedmont

regions of Virginia, South Carolina, and Georgia, and in southcentral

Florida (summary in Grant and Quay 1977).

Platania and Clark (1981) discuss the 1960-80 range expansion

of northern rough-winged swallows (Stelgidopteryx ruficollis) in the

North Carolina Coastal Plain and mapped the known breeding distri-

bution of the species in the state. They also reported nesting season

records of bank swallows (Riparia riparid) from Roanoke Island, Dare

County, but a colony apparently never formed there. Bank swallows

have nested sporadically in North Carolina. Earlier records were avail-

able from 1926 to 1940 in Henderson County (Nicholson 1951, Pickens

1954). Snavely (1978) reported a colony in Wilkes County that was

present from 1977 to 1978. This was the first recent nesting record

for the state, but the colony has since died out. Subsequently, Lee

and Hendrickson (personal observations) found a nesting colony near

Tree Swallow Range Expansion 111

Linville in the Mountains of North Carolina in 1991. All North Carolina

nest sites are in artificial banks made by large earth-moving equipment.

Thus, it appears that current land use practices have provided

open foraging habitats, and various man-made structures and land

modifications have provided suitable artificial nest sites, thus allowing

various species of swallows to expand their breeding range in the

Southeast.

CONCLUSIONS

Like other species of swallows, the tree swallow has expanded

its range considerably since the late 1800s. Earliest records show the

species to be a peripheral breeding species in the southeastern United

States, and nesting records are sporadically distributed both geo-

graphically and temporally (Fig. 1). Range expansion appeared to be

gradual through the 1960s and then explosive from the 1970s to the

present as the birds populated the Piedmont of Maryland, Virginia,

and portions of North Carolina (Fig. 2). The species expansion in the

mountains, Mississippi basin, and Coastal Plain occurred at different

rates. Overall the species has spread south over 600 km in this century

with approximately 220 km (35%) of this expansion occurring in the

last decade. Although various factors such as land clearing were

obviously necessary for the range expansion to occur, the timing of

these changes in land use does not appear to correspond directly with

the expansion in range.

Although land clearing probably benefited all species of swallows

nesting in the Southeast, the change of distribution is also related to

availability of nesting sites. Thus, the northern rough-winged swallow,

the species with the least demanding requirements for nesting sites,

was the first to expand its range. Barn swallows followed, expanding

into the Piedmont and Coastal Plain as wooden bridges were replaced

with concrete ones. Cliff swallows have nested in North Carolina

only since the 1960s, and their current distribution is discontinuous

and dependent on the large reservoirs constructed in the latter part of

this century. Tree swallows expanded their range with the re-introduction

of beavers and after their adoption of nest boxes. The bank swallow,

which has the most restricted distribution, relies on exposed banks

that are sporadically distributed and often not continuously available

because of erosion and invading plant communities.

ACKNOWLEDGMENTS— \ thank various participants of the North

Carolina Breeding Bird Atlas program administered by the North Carolina

State Museum of Natural Sciences for participation in our field studies.

Herb Hendrickson, Julie Angerman-Stewart, Eloise Potter, Maurice

112 David S. Lee

Graves, John Gerwin, Mary Kay Clark, Phil Crutchfield, and Norma

and Bill Siebenhiller each reported nesting tree swallows. C. S. Robbins

and G. A. Hall reviewed the manuscript and maps and provided addi-

tional information.

LITERATURE CITED

American Ornithologists' Union. 1957. Checklist of North American Birds.

Fifth edition. Allen Press, Lawrence, Kansas.

American Ornithologists' Union. 1983. Checklist of North American Birds.

Sixth edition. Allen Press, Lawrence, Kansas.

Bailey, H. H. 1913. The birds of Virginia. J. P. Bell Company, Lynchburg,

Virginia.

Bonwill, A., and H. Owens. 1939. The return of a native. Bulletin of the

Natural History Society of Maryland 10:35-45.

Brooks, M. G. 1936. Notes on the land birds of Garrett County, Maryland.

Bulletin of the Natural History Society of Maryland 7:6-14.

Court, E. J. 1936. Four rare nesting records for Maryland. Auk 53:95-96.

Duyck, B. E. 1981. Range expansion of nesting Tree Swallows. Chat

45:98-100.

Eifrig, C. W. G. 1920. Additions to the "birds of Alleghany and Garrett

counties, Maryland." Auk 37:598-600.

Ganier, A. F. 1964. A Tennessee nesting of the tree swallow. The Migrant

35:51.

Grant, G, and T. L. Quay. 1977. Breeding biology of cliff swallows in

Virginia. Wilson Bulletin 89:286-290.

Gray, D. R., III. 1968. Tree swallow nesting in Maury County. The

Migrant 39:61.

Hall, G A. 1983. West Virginia birds. Carnegie Museum of Natural

History. Special Publication Number 7. Pittsburgh, Pennsylvania.

Hendrickson, H. T. 1984. Osprey and cliff swallows found breeding in

Guilford County, N.C. Chat 48:92-93.

Larner, Y. R. and Committee 1979. Virginia's birdlife: an annotated check-

list. Virginia Avifauna Number 2. Virginia Society of Ornithology, Inc.

Lee, D. S. 1988. Wm. H. Fisher's Mammals of Maryland: a previously

unknown early compilation of the state's fauna. Maryland Naturalist

32:9-37.

Lee, D. S., and R. L. Franz. 1972. A note on the feeding behavior of

the tree swallow. Maryland Birdlife 28:99-100.

Lee, D. S., J. B. Funderburg, and M. K. Clark. 1982. A distributional

survey of North Carolina mammals. Occasional Papers of the North

Carolina Biological Survey 1982-10. North Carolina State Museum,

Raleigh.

LeGrand, H. E., and E. F. Potter. 1980. Ashe County breeding bird

foray— 1979. Chat 44:5-13.

Tree Swallow Range Expansion 113

Mansuetti, R. 1950. Extinct and vanishing mammals of Maryland and

District of Columbia. Maryland Naturalist 23:71-75.

McConnell, J. 1981. Cliff swallows nesting on Fontana Dam, N.C. Chat

45:102.

Mengel, R. M. 1965. The birds of Kentucky. Ornithological Monographs

Number 3.

Monroe, B. L., A. L. Stamm, and B. L. Palmer-Ball, Jr. 1988. Annotated

checklist of the birds of Kentucky. Kentucky Ornithological Society.

Murray, J. J. 1952. A checklist of the birds of Virginia. Virginia Society

of Ornithology.

Nicholson, D. J. 1951. Summer birds of Lake Summit, Henderson County,

N.C. Chat 15:39-41.

Nicholson, C. P., and T. D. Pitts. 1982. Nesting of the tree swallow in

Tennessee. The Migrant 53:73-80.

Olson, F. B. 1968. Tree swallows nesting in east Tennessee. The Migrant

39:59-60.

Pearson, T. G., C. S. Brimley, and H. H. Brimley. 1942. Birds of North

Carolina. North Carolina Department of Agriculture, Raleigh.

Pickens, A. L. 1954. The bank swallow in the Carolinas. Chat 18:53-54.

Pindar, L. O. 1889. List of the birds of Fulton County, Kentucky. Auk

6:310-316.

Pindar, L. O. 1925. Birds of Fulton County Kentucky. Wilson Bull. 37:77-

88, 163-169.

Platania, S. P., and M. K. Clark. 1981. Rough-winged swallow nesting in

coastal North Carolina. Chat 45:100-102.

Robbins, C. S., and W. T. Van Velzen. 1968. Field list of the birds of

Maryland. Maryland Avifauna Number 2. Maryland Ornithological

Society.

Snavely, R. R. 1978. Bank swallows nesting in North Carolina. Chat

42:83-84.

Stewart, R. E., and C. S. Robbins. 1958. Birds of Maryland and the

District of Columbia. North American Fauna Number 62. United

States Fish and Wildlife Service, Washington, DC.

Williams, M. D. 1976. The season-central plateau and basin region.

Migrant 47:99-100.

Accepted 30 July 1992

114

POTENTIAL EFFECTS OF OIL SPILLS ON SEABIRDS

AND SELECTED OTHER OCEANIC VERTEBRATES

OFF THE NORTH CAROLINA COAST

by

David. S. Lee and Mary C. Socci

Based primarily on data gathered offshore during the past 15 years

by the staff of the North Carolina State Museum of Natural Sciences, this

book presents information regarding distribution and susceptibility to oil

pollution for 25 oceanic species: 14 birds, 6 mammals, and 5 turtles. An

overlay can be placed on range maps to demonstrate the proximity of

species occurrence to the oil lease sites off Cape Hatteras.

1989 64 pages Softbound ISBN 0-917134-18-4

Price: $8 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: OIL SPILL BOOK, N.C. State Museum of Natural Sciences,

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

An Observation of Fea's Petrel, Pterodroma feae

(Procellariiformes: Procellariidae), Off the

Southeastern United States, With Comments on the Taxonomy

and Conservation of Soft-plumaged and Related Petrels

in the Atlantic Ocean

J. Christopher Haney1

Biology Department, Woods Hole Oceanographic Institution,

Woods Hole, Massachusetts 02543,

Craig A. Faanes

U.S. Fish and Wildlife Service, 203 West Second Street,

Grand Island, Nebraska 68801,

AND

William R. P. Bourne

Zoology Department, Aberdeen University, Tilly drone Avenue,

Aberdeen AB9 2TN, Scotland

ABSTRACT — The soft-plumaged petrel and related species {Pterodroma

spp.) remain one of the most poorly known seabird taxa in the

Atlantic Ocean, and there is cause for serious concern over the

continued survival of two North Atlantic forms. Soft-plumaged pet-

rels were formerly considered to be a single, albeit morphologi-

cally variable, complex of one species. However, taxonomists now

generally consider the complex to contain at least three species

including the nominate. We report a marine occurrence of a North

Atlantic species, probably Fea's petrel P. feae, from the South At-

lantic Bight off the coast of the southeastern United States. We

describe morphological characteristics for separating the various forms

and consider the recent at-sea sightings in relation to dispersal fac-

tors such as seasonal wind regimes and coassociation with other

seabird species that regularly disperse to western sectors from east-

ern sectors in the North Atlantic Ocean.

Gadfly petrels in the genus Pterodroma are known to disperse

widely over the world's oceans, often at considerable distances from

their natal colonies (Bourne 1967). Soft-plumaged petrels, P. mollis

(Gould), are medium-sized gadfly petrels breeding in the Atlantic Ocean,

the southern Indian Ocean, and the South Pacific Ocean. Although

widely distributed, the number of island colonies is limited, and relative-

present address: Wildlife Technology Program, School of Forest Resources, Pennsylvania

State University, College Place, DuBois, Pennsylvania 15801-3199.

Brimleyana 18:115-123, June 1993 115

116 J. Christopher Haney et al.

ly few at-sea observations of either soft-plumaged or related northern

populations have been made away from their breeding areas (Collar

and Stuart 1985). We report such a sighting from the waters off the

southeastern United States and provide some field marks (see also

Enticott 1991) that may be useful for differentiating forms within a

difficult taxonomic complex formerly regarded as a single species (Bourne

1983a).

Sighting description — On 9 November 1984, two of us (J.C.H.

and C.A.F.) observed an unusual gadfly petrel with other pro-

cellariiforms during a census of seabirds at the edge of the continental

shelf off the Georgia coast. A mixed flock (30-40 individuals) of the

black-capped petrel, Pterodroma hasitata (Kuhl), Cory's shearwater,

Calonectris diomedea (Scopoli), greater shearwater, Puffinus gravis

(O'Reilly), and Audubon's shearwater, Puffinus Iherminieri Lesson, accom-

panied by pomarine jaegers, Stercorarius pomarinus (Temminck), and

herring gulls Larus argentatus Pontoppidan, was seen feeding over a

fish school near the western frontal boundary of the Gulf Stream at

31°39'N, 79°24'W. This was approximately 145 km due east of St.

Catherine's Island, Georgia (depth 250 m, surface temperature 25.5C).

We initially noticed a gadfly petrel without a white nape or rump in

the feeding flock at 1625 EST. During the next 15 minutes, we

watched from opposite ends of the stationary research vessel while

the bird flew and foraged with other seabirds. It was seen from as

close as 30 m through 9 x 35 and 10 x 40 binoculars.

J. C. Haney noted that, compared with the high bounding flight

of nearby black-capped petrels, this gadfly petrel had more rapid wing-

beats and flew closer to the ocean surface. It was not noticeably

different in size from the black-capped petrel, and like that species, it

alternated banking and gliding with first the dorsal and then the ventral

surface exposed to the observers. The bird's overall appearance was

dark gray above and white below. The gray tail was wedge-shaped

and slightly paler than the back, without light-colored upper tail coverts.

The crown appeared darker than the nape and hindneck, and the

forehead was white. No dark facial mask around the eye was observed.

There was conspicuous mottling or streaking along the bird's flanks.

This field mark was very obvious and set the bird apart from nearby

black-capped petrels which have clear white flanks. The bird did not

have a complete breast band, although it did have a short, ventrally-

projecting light gray bar on both sides of the neck in front of the

wings. C. A. Faanes noticed that the underwing coverts were noticeably

gray to the base of the primaries. The primary feathers appeared

white-based, reminiscent of a jaeger in flight, suggesting that the bird

may have been molting its wing coverts at the time. The upperwings

Fea's Petrel 117

were medium gray, darker on the wing coverts. The bill was dark

gray or black.

Taxonomy and identification — Various field guides on board indi-

cated that the bird must belong to one of the North Atlantic gadfly

petrels closely related to the soft-plumaged petrel P. mollis. The wholly

dark underwing eliminated all other Pterodroma that have been docu-

mented as occurring in the North Atlantic (i.e., Pterodroma hasitata,

P. cahow [Nichols and Mowbray], P. arminjoniana [Giglioli and

Salvadori]). The gray coloration, contrast between back and tail, dark

bill, and wholly white underparts ruled out similar petrels from other

oceans (i.e., Lugensa brevirostris Lesson, Pterodroma incerta Bonaparte,

Procellaria cinerea Gmelin). The soft-plumaged petrels, formerly treated

as four or more subspecies (e.g., Mathews 1934, Harrison 1983), have

recently been reclassified as three distinct species, Pterodroma madeira

Mathews, P. feae (Salvadori), and the nominate P. mollis (Bourne

1983a, Imber 1985, Zino and Zino 1986; see also Sibley and Ahlquist

1990, Warham 1990). All three forms may appear to have various

shades of dark gray-brown above, depending upon feather wear and

lighting conditions (Enticott 1991). The nominate soft-plumaged petrels

from the southern hemisphere are normally darkest, with variable gray

markings extending across the upper breast and shading into the white

face and chin above but contrasting sharply with the white belly

below (Fig. 1). Thus, these birds appear to have a dark head and

neck but pale face contrasting with white underparts. Some southern

soft-plumaged petrels have a prominent "W" mark on the wing, some

have noticeably paler rumps, and in some, the underwing shows variable

amounts of contrasting white coloration. The North Atlantic forms are

known as the Freira petrel (P. madeira) and Fea's petrel (P. feae:

see Bannerman and Bannerman 1966; known also as "gon-gon" petrel

on the Cape Verde Islands [Bourne 1983a, Collar and Stuart 1985]).

These two species are both normally paler and grayer above than the

nominate, with the entire body white below (Fig. 2, personal observation,

J. Enticott personal communication). The amount of black around the

eye in these forms ranges from conspicuous to essentially lacking. In

general, photographs we examined indicated that both the underwings

and upperwings are more uniformaly colored in the North Atlantic

species.

The Freira petrel and Fea's petrel are difficult to separate on

the basis of plumage alone (Bourne 1983a, Fisher 1989). Fea's petrel

is larger and is known to nest (in winter) only on the Cape Verde

Islands and on Bugio (in the Desertas group), although there are

recent sight and vocalization records from the Azores (Bibby and del

Nevo 1991) and from Great Salvage Island 300 km south of the

118 J. Christopher Haney et al.

k*

Fig. 1. Specimen of nominate Pterodroma mollis, showing underparts, collected

from Antipodes Island, 13 February 1969, by J. Warham (Department of

Zoology, University of Canterbury, Christchurch 1, New Zealand).

Madeiras (James and Robertson 1985). Measurements of wing and

body length in Fea's petrel overlap extensively with the black-capped

petrel (Cramp and Simmons 1977, Harrison 1983). When we (J.C.H.

and C.A.F.) compared specimens of P. feae and P. hasitata, we con-

cluded that size differences between these two species would be difficult

to detect under field conditions. The Freira petrel (a summer breeder)

is significantly smaller than either the Fea's petrel or the soft-plumaged

petrel (Cramp and Simmons 1977, Fisher 1989). Bourne (1983a) stated

the P. madeira has less mottling on the flanks than P. feae. However,

other accounts differ (see Cramp and Simmons 1977, Harrison 1983),

and such relative field marks may not be useful at sea in the absence

of direct comparison and experience.

Status and distribution — We believe that the bird we observed

must have been P. feae for the following reasons. First, the lack of

a complete breast band eliminated southern soft-plumaged petrels. None

of >290 specimens of the northern forms or numerous living examples

examined by P. and F. Zino (personal communication) have a marked

breast band, although a small minority of individuals with breast bands

have been reported at sea off Madeira (taxonomic identity unknown;

B. Zonfrillo personal communication). Generally, there is at least 1

cm of white between the two sides of the partial breast band in the

northern species (F. Zino personal communication). Second, the large

Fea's Petrel

119

Fig. 2. Photograph of Pterodroma madeira, showing distinctive dark underwing

and pure white underparts, Madeira, 16 June 1969, P. A. Zino (Quinta da

Vista Alegre, Rua do Dr. Pita 5, 9000 Funchal, Madeira).

120 J. Christopher Haney et al.

size of the bird we observed (near or identical to black-capped petrel)

indicates the Fea's petrel. Third, the population of the smaller Freira

petrel is apparently reduced to only a few dozen pairs (A. and F.

Zino personal communication), and it has not been recorded conclu-

sively away from the breeding sites. Therefore, it seems less likely to

be seen at sea. Finally, the Fea's petrel has been found wandering a

similar distance before, e.g. to Israel where it was collected on 8

February 1963 (Bourne 1983b), although the majority of pelagic records

for this form have been near the Canary Islands and off western

Africa (Bourne and Dixon 1973, 1975; Lee 1984).

Neither the soft-plumaged nor related petrels have yet been

accepted onto the official list of North American birds (cf. A.O.U.

1983, 1985). Lee's (1984) observation of a soft-plumaged petrel off

North Carolina on 3 June 1981 was the first report for this continent;

that individual had a complete breast band, a trait that points to a

southern origin. It is notable that Lee's observation occurred during

austral winter, a period when the southeast trade winds extend north

across the equator and are liable to drift southern-hemisphere seabirds

to the northwest Atlantic (comparable seasonal patterns for northward

dispersal by seabirds occur in the Indian Ocean, see Ash [1983]).

Similarly, the more northern Fea's petrel appeared off North America

to the west of its breeding sites at a time when the northeast tradewinds

still extend north. This pattern starts to deteriorate early in the northern

winter after which time the westerlies move south to replace the

northeast trades (at a time when the Fea's petrel was recorded as

vagrant to Israel). In addition to our and Lee's (1984) records of

soft-plumaged-type petrels off eastern North America, additional sight

reports, including photographs of the nominate soft-plummed petrel,

were made during May 1992, also off North Carolina (D. S. Lee

personal communication, Anon. 1992).

DISCUSSION

We are aware of the inherent uncertainty in sight records of

pelagic seabirds, especially Pterodroma, and we advocate photographic

documentation when possible. However, we disagree with recent sug-

gestions (Legrand 1985) that records of rare gadfly petrels off the

eastern United States be supported with voucher specimens. Most

countries, including the United States, have little jurisdiction over and

limited protection for rare pelagic birds beyond coastal waters. Standard

collection practices could further reduce the numbers of Atlantic petrels,

some of which are now composed of only a few dozen breeding

pairs. This risk seems unwarranted.

Fea's Petrel 121

The soft-plumaged petrel complex remains one of the least known

and seriously threatened seabird taxa in the Atlantic Ocean. There is

cause for serious concern over the continued survival of the North

Atlantic species. The total population of Freira petrels (P. madeira)

may consist of no more than 50 pairs nesting at the higher elevations

of Madeira, making it Europe's rarest bird (Buckle and Zino 1989).

During the late 1980s, several failed breeding attempts were recorded,

possibly because of interference by rats and/or predation by feral cats

(A. Zino personal communication). The population of Fea's petrels

(P. feae), which is unlikely to total more than several hundred pairs,

is subject to human predation arising from medicinal use of the bird's

body fat (Cramp and Simmons 1977, Collar and Stuart 1985). Breeding

in dispersed colonies in burrows, earth screes, and rocky outcrops,

this ground-nesting species is also vulnerable to predation by cats,

rats, and other feral mammals; deforestation, vegetation destruction,

and soil erosion by goats; and possible competition with rabbits for

nesting burrows (Collar and Stuart 1985:42).

As in several other eastern and southern Atlantic procellariiforms

(for examples, see Lee 1979, 1984; Haney and Wainright 1985), disper-

sal into the western Atlantic Ocean by petrels of the soft-plumaged

complex could prove to be a regular occurrence overlooked owing to

the species' rarity rather than casual vagrancy. Fea's petrels have

been observed accompanying groups of Cory's shearwaters in the eastern

Atlantic at other seasons (Lambert 1980). The mixed-species flock in

which we encountered the Fea's petrel was dominated by Cory's shear-

waters, a far more common species that occupies a sympatric breeding

range with Fea's petrel in the eastern North Atlantic. If such interspeci-

fic associations are typical, then the petrel we observed may have

followed, or have been locally attracted to, large flocks of trans-

Atlantic migrant shearwaters during its wanderings to the offshore

waters of the southeastern United States.

ACKNOWLEDGMENTS— We thank M. Lecroy of the American

Museum of Natural History for the loan of specimens of P. feae,

and E. McGhee of the University of Georgia Museum of Natural

History for curatorial assistance. J. Warham, P. Meeth, B. Zonfrillo,

S. M. Lister, and F. and P. A. Zino supplied additional information,

including reference photographs of soft-plumaged and related petrels

for comparison. We also thank D. H. White, F. Zino, and three

anonymous reviewers for valuable comments on earlier drafts. The

University of Georgia Marine Extension Service and crew of the R/V

"Bulldog" extended logistic support during our offshore surveys. Addi-

tional support was received from The Pew Charitable Trusts under

122 J. Christopher Haney et al.

the Marine Policy Center's program "Changing Global Processes and

Ocean Conservation" at Woods Hole Oceanographic Institution (WHOI).

This is WHOI Contribution No. 7763.

LITERATURE CITED

American Ornithologists' Union. 1983. Checklist of North American birds.

Sixth Edition. Allen Press, Lawrence, Kansas.

American Ornithologists' Union. 1985. Thirty-fifth supplement to the

American Ornithologists' Union checklist of North American birds.

Auk 102:680-686.

Anonymous. 1992. A soft-plumaged petrel in the U.S.A. Birding World 5.

Ash, J. S. 1983. Over fifty additions of birds to the Somalia list including

two hybrids, together with notes from Ethiopia and Kenya. Scopus

7:54-79.

Bannerman, D. A., and W. M. Bannerman. 1966. Birds of the Atlantic

islands. Volume 3, Oliver & Boyd, Edinburgh, United Kingdom.

Bibby, C. J., and A. J. del Nevo. 1991. A first record of Pterodroma

feae from the Azores. Bulletin of the British Ornithological Club

111:183-186.

Bourne, W. R. P. 1967. Long-distance vagrancy in the petrels. Ibis 109:141-

167.

Bourne, W. R. P. 1983a. The soft-plumaged petrel, the gon-gon and the

Freira, Pterodroma mollis, P. feae, and P. madeira. Bulletin of the

British Ornithological Club 103:52-58.

Bourne, W. R. P. 19836. A gon-gon Pterodroma (mollis) feae in Israel.

Bulletin of the British Ornithological Club 103:110.

Bourne, W. R. P., and T. J. Dixon. 1973. Observations of seabirds 1967-

1969. Sea Swallow 22:29-60.

Bourne, W. R. P., and T. J. Dixon. 1975. Observations of seabirds 1970-

1972. Sea Swallow 24:65-88.

Buckle, A., and F. Zino. 1989. Saving Europe's rarest bird. Roundel

67:112-116.

Collar, N. J., and S. N. Stuart. 1985. Gon-gon, Pterodroma feae (Salvadori

1900), Freira, Pterodroma madeira Mathews 1934. Pages 39-46, 52-58

in: Threatened birds of Africa and related islands. ICPB/IUCN Red

Data Book, Part 1, Cambridge, United Kingdom.

Cramp, S., and K. E. I. Simmons. 1977. The birds of the western Palearctic,

Volume 1. Oxford Universtiy Press, London, United Kingdom.

Enticott, J. W. 1991. Identification of soft-plumaged petrel. British Birds

84:245-264.

Fisher, D. 1989. Pterodroma petrels in Madeira. Birding World 2:283-287.

Haney, J. C, and S. C. Wainright. 1985. Bulwer's petrel from the South

Atlantic Bight. American Birds 39:868-870.

Harrison, P. 1983. Seabirds: an identification guide. Houghton-Mifflin Com-

pany, Boston, Massachusetts.

Fea's Petrel 123

Imber, M. J. 1985. Origins, phylogeny and taxonomy of the gadfly petrels

Pterodroma spp. Ibis 127:197-229.

James, P. C, and H. A. Robertson. 1985. Soft-plumaged petrels Pterodroma

mollis at Great Salvage Island. Bulletin of the British Ornithological

Club 105:25-26.

Lambert, K. 1980. Beitrage zur Vogelwelt der Kapverdischen Inseln. Beitr.

Vogelkd. 26:1-18.

Lee, D. S. 1979. Second record of the South Trinidad petrel {Pterodroma

arminjoniana) for North America. American Birds 33:138-139.

Lee, D. S. 1984. Petrels and storm-petrels in North Carolina offshore

waters: including species previously unrecorded for North America.

American Birds 38:151-163.

Legrand, H. 1985. South Atlantic coast region. American Birds 39:157.

Mathews, G. M. 1934. The soft-plumaged petrel Pterodroma mollis and

its subspecies. Bulletin of the British Ornithological Club 54:178-179.

Sibley, C. G., and J. E. Ahlquist. 1990. Phylogeny and classification of

birds: a study in molecular evolution. Yale University Press, New Haven,

Connecticut.

Warham, J. 1990. The petrels: their ecology and breeding systems. Academic

Press, London, United Kingdom. 440 pp.

Zino, P. A., and F. Zino. 1986. Contribution to the study of the petrels

of the genus Pterodroma in the archipelago of Madeira. Boletim do

Museu Municipal do Funchal 38:141-165.

Accepted 17 March 1992

124

ENDANGERED, THREATENED, AND

RARE FAUNA OF NORTH CAROLINA

PART I. A RE-EVALUATION OF THE MAMMALS

Edited by Mary Kay Clark

This book is a report prepared by a committee appointed in 1985 by

the North Carolina State Museum of Natural Sciences to re-evaluate the

list of mammals presented in Endangered and Threatened Plants and

Animals of North Carolina (John E. Cooper, Sarah S. Robinson, and John

B. Funderburg, editors. N.C. State Mus. Nat. Hist., Raleigh, 1977), which

is now out of print. Committee members were Mary Kay Clark, David A.

Adams, William F. Adams, Carl W. Betsill, John B. Funderburg, Roger A.

Powell, Wm. David Webster, and Peter D. Weigl. The report treats 21

species listed in the following status categories: Endangered (5), Threatened

(1), Vulnerable (6), and Undetermined (9). Most species accounts discuss

the animal's physical characteristics, range, habitat, life history and eco-

logy, special significance, and status (including the rationale for the evalua-

tion and recommendations for protection) and provide a range map and

an illustration of the animal's external characters. Ruth Brunstetter and

Renaldo Kuhler illustrated the book. An introductory section contributed

by Ms. Clark discusses the changes in status that occurred in the decade

between 1975 and 1985. It also mentions efforts to protect marine mam-

mals and includes a checklist of the cetaceans known from North Carolina.

1987 52 pages Softbound ISBN 0-917134-14-1

Price: $5 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: ETR MAMMALS, N.C. State Museum of Natural Sciences,

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

Cotton Mice, Peromyscus gossypinus LeConte

(Rodentia: Cricetidae), in the Great Dismal Swamp

and Surrounding Areas

James L. Boone1

Museum of Natural History, Institute of Ecology, and

Savannah River Ecology Laboratory

University of Georgia, Athens, Georgia 30602

AND

Joshua Laerm

Museum of Natural History, University of Georgia,

Athens Georgia 30602

ABSTRACT — Livetrapping of small mammals was conducted in the

Great Dismal Swamp and other areas of North Carolina in 1990.

Five Peromyscus gossypinus were caught in the Dismal Swamp pro-

per, and 42 were caught in the Chowan Swamp adjacent to the

Dismal Swamp. These are the first published records of P. gossypinus

taken in the Dismal Swamp region since the 1930s.

Rose et al. (1990) suggested that the cotton mouse, Peromyscus

gossypinus LeConte, could be extinct in the Great Dismal Swamp of

Virginia and North Carolina. With the exception of two specimens

collected in 1933 by Dice (1940), virtually none has been captured

there since the turn of the century despite the efforts of Handley

(1979) in the 1950s and Rose et al. (1990) in the 1980s. Our recent

collections and genetic analyses show P. gossypinus exists in the Dismal

Swamp, and that based on capture rate it is uncommon in the Swamp

proper, but is relatively abundant in areas adjacent to the southern

section of the Swamp.

Separating P. gossypinus from P. leucopus Rafinesque (white-

footed mouse) can be difficult both for live and museum specimens.

Dice (1940) states that in eastern Virginia size characteristics but not

color can be used to separate these species. Our studies (unpublished

data) show that several cranial and external characters from adult

specimens are required for consistent species identification with

discriminant analysis. However, a fixed allozyme difference at the

Glucose-6-Phosphate Isomerase locus (GPI or PGI, Enzyme Commis-

sion No. 5.3.1.9), and nearly fixed differences at the Albumin and

alpha-Glycerolphosphate dehydrogenase (a-GPD or GPD), Enzyme

Commission No. 1.1.1.8) loci separate these two species (Price and

Kennedy 1980; Robbins et al. 1985; Boone 1990; Boone unpublished

data).

'Direct correspondence to James Boone at the Museum of Natural History.

Brimleyana 18:125-129, June 1993 125

126 James L. Boone and Joshua Laerm

METHODS

We captured small mammals with Sherman livetraps in north-

eastern North Carolina for studies examining subspecific affinities,

population genetics, and Lyme disease (Magnarelli et al. 1992) in P.

gossypinus. On 26 and 27 April 1990, we trapped in the Dismal

Swamp along Highway 158 from 6.3 to 10.6 km east of Highway 32

(east of Sunbury, Gates County, North Carolina) for 587 trapnights.

On 13 June 1990, we placed 200 traps in the Chowan Swamp between

2.9 and 5.3 km south of Gatesville (Gates County, North Carolina).

On 28 and 29 April 1990, we trapped along the Cashie River in and

around Windsor (Bertie County, North Carolina) for 350 trapnights,

and we placed 150 traps in and around Richlands (Onslow County,

North Carolina) on 30 April 1990.

Locations of trap lines and specific traps were selected to maxi-

mize the capture of P. gossypinus based on our understanding of

its habitat preference and ecology learned from the capture of more

than 2,100 cotton mice from throughout its entire range. Although

these mice can be caught almost anywhere, they seem to exist in

highest densities in thick, undisturbed (anthropogenic or natural), sea-

sonally flooded, riparian woodlands near water. On coastal barrier

islands where these habitats do not exist, they seem to occur most

densely in undisturbed old-growth oak-palmetto (Quercus sp. and

Serenoa repens) forests. Traps were set on, in, and under logs, in

trees, under stumps, in the rotten bases of trees, on the edges of

ponds, on floating debris in flooded forests, as well as in old build-

ings and trash piles. More than one trap was set in particularly pro-

mising sites.

We used allozyme markers to identify the Peromyscus. Genetic

analysis was performed with standard horizontal starch gel electro-

phoretic and protein staining techniques on blood, liver, and muscle

for 42 enzyme and protein loci. Techniques were similar to those of

Selander et al. (1971) as described in Boone (1990).

Body mass was measured to the nearest 0.1 g. Age (juvenile,

subadult, or adult) was determined by pelage color, and reproduc-

tive status was determined by examination of external and internal

reproductive structures. Non-adult and pregnant females were deleted

from morpholo-gical comparisons.

RESULTS

Peromyscus gossypinus was captured in each of the four areas

examined, and P. leucopus was captured in all areas except Rich-

lands (Table 1). Additionally, one golden mouse (Ochrotomys nuttalli

Harlin) and one juvenile Virginia opossum (Didelphis virginiana

Great Dismal Swamp Cotton Mice

127

Table 1. Peromyscus gossypinus and P. leucopus captured in North Carolina,

1990.

Location

Species

P. gossypinus

P. leucopus

Captures/ Captures/

dumber « nnn Number 1 nnn

cauSht tracts cau8ht trap'nThts

Gates County Dismal Swamp

Gates County Chowan Swamp

Bertie County Windsor

Onslow County Richlands

Kerr) were captured in the Dismal Swamp, and one Blarina was

captured at Windsor.

We found that Dice's (1940) suggestion that these Peromyscus

species can be distinguished by size is not strictly true. Our comparison

of genetic markers and morphology indicates that although P. gossypinus

tends to be larger and heavier than P. leucopus, there is considerable

overlap. For the mice caught east of Sunbury, body mass of P. gossypinus

ranged from 20.9 to 35.5 g (x = 26.3 g, n = 5), whereas P. leucopus

ranged from 14.6 to 24.6 g (x = 19.1 g, n = 26). In the Chowan

Swamp, body mass of P. gossypinus ranged from 17.1 to 36.8 g,

( x = 25.9 g, n = 42); the one adult P. leucopus weighed 15.9 g. In

the Windsor area, P. gossypinus ranged from 19.2 to 37.9 g (x =

28.4 g, n = 22), and P. leucopus ranged from 17.1 to 24.1 g (x =

20.4 g, n = 4). The P. gossypinus from Richlands ranged from 21.2

to 39.4 g (x = 29.2 g, n = 33). Therefore, if Rose et al. (1990)

used size to identify Peromyscus, some of the specimens identified as

P. leucopus by might actually have been P. gossypinus.

DISCUSSION

Our results probably differ those of Rose et al. (1990) as a

result of different trapping location, design, and methods. In the southern

portion of the Dismal Swamp, Rose et al. (1990) used pitfall traps

set on a grid. We used only Sherman livetraps, and our collection

locations were selected to target habitats thought to be optional for

P. gossypinus without concern for determining density or other demo-

graphic parameters. Therefore, we were not confined to a grid, and

we were able to trap in areas, and place traps in sites, that would be

inappropriate to use with pitfall traps in a demographic study. Further-

more, our trapping was only conducted in the southernmost part of

128 James L. Boone and Joshua Laerm

the Swamp, an area more accessible to migrants from the Chowan

Swamp where P. gossypinus is abundant, whereas the majority of

Rose et al.'s (1990) effort was concentrated in the northern section

of the Swamp where P. gossypinus might be absent.

Although P. gossypinus is abundant in areas near the Dismal

Swamp, it is probably not currently abundant in the swamp proper.

Handley (1979) stated that P. gossypinus densities fluctuate widely in

the Swamp, and this population could simply be at a low point in

its cycle. This species now occurs in the Great Dismal Swamp, but

current management practices in the Great Dismal Swamp National

Wildlife Refuge that promote clearings and vegetational heterogeneity

might endanger it because we have observed that P. gossypinus occurs

in greatest density in mature, undisturbed riparian forests.

ACKNOWLEDGMENTS— -The Savannah River Ecology Laboratory

under contract DE-AC09-76SR00819 between the U. S. Department

of Energy and the University of Georgia's Institute of Ecology, the

University of Georgia Museum of Natural History, Sigma Xi, and the

Theodore Roosevelt Memorial Fund administered by the American

Museum of Natural History provided support for this work, and Kevin

Roe assisted with the trapping.

LITERATURE CITED

Boone, J. L. 1990. Reassessment of the taxonomic status of the cotton

mouse (Peromyscus gossypinus anastasea) on Cumberland Island,

Georgia, and implications of this information for conservation. M.S.

Thesis. University of Georgia, Athens.

Dice, L. R. 1940. Relationship between the wood-mouse and the cotton-

mouse in eastern Virginia. Journal of Mammalogy 21:14-23.

Handley, C. O., Jr. 1979. Mammals of the Dismal Swamp: a historical

account. Pages 297-357 in The Great Dismal Swamp (P. W. Kirk, Jr.,

editor) University Press of Virginia, Charlottesville.

Magnarelli, L. A., J. H. Oliver, H. J. Hutcheson, J. L. Boone, and J. F.

Anderson. 1992. Antibodies to Borrelia burgdorferi in rodents in the

eastern and southern United States. Journal of Clinical Microbiology

30:1449-1452.

Price, P. K, and M. L. Kennedy. 1980. Genetic relationships in the

white-footed mouse, Peromyscus leucopus, and the cotton mouse,

Peromyscus gossypinus. American Midland Naturalist 103:73-82.

Robbins, L. W., M. H. Smith, M. C. Wooten, and R. K. Selander. 1985.

Biochemical polymorphism and its relationship to chromosomal and

morphological variation in Peromyscus leucopus and Peromyscus

gossypinus. Journal of Mammalogy 66:498-510.

Great Dismal Swamp Cotton Mice 129

Rose, R. K., R. K. Everton, J. F. Stankavich, and J. W. Walke. 1990.

Small mammals of the Great Dismal Swamp of Virginia and North

Carolina. Brimleyana 16:87-101.

Selander, R. K., M. H. Smith, S. Y. Yang, W. E. Johnson, and J. B.

Gentry. 1971. Biochemical polymorphism and systematics in the genus

Peromyscus. I. Variation in the old-field mouse (Peromyscus polionotus).

Studies in Genetics VII. University of Texas Publication 7103:49-90.

Accepted 1 September 1992

130

ENDANGERED, THREATENED, AND

RARE FAUNA OF NORTH CAROLINA

PART II. A RE-EVALUATION OF THE MARINE

AND ESTUARINE FISHES

by

Steve W. Ross, Fred C. Rohde, and David G. Lindquist

This is the second in a series of reports by committees appointed in

1985 by the North Carolina State Museum of Natural Sciences to re-evaluate

the faunal lists presented in Endangered and Threatened Plants and Animals

of North Carolina (John E. Cooper, Sarah S. Robinson, and John B. Funderburg,

editors. N.C. State Mus. Nat. Hist., Raleigh, 1977), which is now out of

print. The report on marine and estuarine fishes by Ross, Rohde, and Linquist

treats one Endangered species, six Vulnerable species, and four anadromous

fishes that, while not formally listed, are of some concern. Five species

listed as being of Special Concern in 1977 no longer warrant formal status.

The publication includes six original drawings by Renaldo Kuhler.

1988 20 pages Softbound ISBN 0-917134-17-6

Price: $3 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: ETR MARINE FISHES, N.C. State Museum of Natural Sciences,

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

Observations Regarding the Diet of Florida Mice,

Podomys floridanus (Rodentia: Muridae)

Cheri A. Jones1

Mississippi Museum of Natural Science,

111 North Jefferson Street, Jackson, Mississippi 39201-2897

ABSTRACT — The diet and feeding behavior of the Florida mouse

(Podomys floridanus) were examined during an ecological study in

Putnam County, Florida. Field and laboratory observations pro-

vided additional evidence that Podomys takes a wide variety of

plant and animal foods. Preliminary preference tests with acorns

from six species of oaks suggest that acorns of the dominant

species (Quercus laevis) in the study area are less favored than

those of other species. The crash of a population where supple-

mental food was provided suggests that local populations are not

food limited.

Almost nothing is known about the natural diet of the Florida

mouse, Podomys floridanus (Chapman). Merriam (1890:53) reported

an observation that these mice ate seeds of "scrub-palmettoes" in

southeast Florida. The association of Podomys with turkey oaks

(Quercus laevis) and other oaks was noted by Merriam (1890) and

Bangs (1898); Layne (1970), Humphrey et al. (1985), and Packer

and Layne (1991) suggested that acorns were a major food during

masting years. Milstrey (1987) described Podomys eating engorged

soft ticks (Ornithodoros turicata americanus) that parasitize gopher

frogs (Rana capito) and gopher tortoises (Gopherus polyphemus).

Presumably other foods include insects, seeds, nuts, fungi, and other

plant material (Layne 1978, Jones and Layne In Press). The study by

Packer and Layne (1991) is the first to examine foraging behavior of

this species.

Effects of food supplies on local distributions of Podomys also

have been poorly studied. The only attempt to determine whether

populations are limited by food availability was the supplementation

experiment performed by Young (Young 1983, Young and Stout 1986)

on two grids in sand pine (Pinus clausa) scrub in Orange County,

Florida. Other rodents responded to the additional food, but Podomys

rarely appeared on grids and failed to establish a permanent population

during the experiment, although the species was abundant previously.

Young (1983) concluded that Podomys populations were limited by

factors other than food.

'Present address: Denver Museum of Natural History, 2001 Colorado Boulevard,

Denver, Colorado 80205-5798.

Brimleyana 18:131-140, June 1993 131

132 Cheri A. Jones

The purpose of this article is to report observations of feeding

behavior of wild and captive Podomys. Additionally, I performed

acorn preference tests and a supplemental feeding experiment to

determine whether local distributions were due to food supply.

MATERIALS AND METHODS

Field Studies — Florida mice were trapped on the Anderson-Cue

and Smith Lake sandhills on the Katharine Ordway Preserve-Swisher

Memorial Sanctuary in Putnam County, Florida. These xeric sandhills

are "high pine" communities dominated by longleaf pine (Pinus palustris)

and turkey oak (Q. laevis). Brand (1987), Eisenberg (1988), Franz

(1986, 1990), and Jones (1990) described the fauna of these sandhills.

Populations of Podomys occur on sandhills and old pastures on the

Ordway Preserve, where they are closely associated with burrows of

gopher tortoises. Cotton mice (Peromyscus gossypinus) and golden

mice (Ochrotomys nuttalli) inhabit lower, more mesic habitats on the

preserve.

Each tortoise burrow on the Anderson-Cue and Smith Lake

sandhills was flagged and marked with a unique number. Florida

mice were caught in Sherman traps placed at the mouths of tortoise

burrows on both sandhills. Animals were released near the burrow

entrance so that I could observe escape responses and foraging

behavior (Jones 1990). I used standard mark-and-recapture techniques,

in which individuals were toe-clipped, sexed, and weighed. I calculated

minimum trappability [(number of captures - 2)/(possible captures -

2)] for animals captured three or more times (Hilborn et al. 1976).

I used a breeding colony derived from animals captured at

Ashley Old Pasture and near Smith Lake to perform food preference

tests. Captive animals were housed in aquaria fitted with hardware-

cloth tops. The maintenance diet consisted of rodent chow (Wayne

Rodent Blox) and water provided ad libitum, supplemented with lettuce,

carrots, apples, strawberries, sunflower seeds, mixed bird seed, oatmeal,

mealworms, and crickets.

Preference Tests — I performed preference tests to determine whether

acorns of turkey oaks (Q. laevis) were selected over acorns of other

species present on the Ordway Preserve. At least 24 hours before

beginning a test, a mouse was placed in an aquarium with clean

kitty litter, nesting material, water, and rat chow. To start a trial I

removed the chow and added three bowls, each containing five acorns

of a single species, at about 1900 hours. Each acorn was marked,

weighed, and measured, but no effort was made to ensure that all

acorns in a bowl were identical in size. I controlled for location

effects by shifting relative positions of the three types of acorns in

Diet of Florida Mice 133

each trail. Acorns with weevil holes were not used, and in a single

trial all acorns either had caps or lacked them. Approximately 12

hours later I removed bowls, acorns, and acorn fragments. I recorded

whether acorns were removed from bowls and whether they were

opened and eaten, opened and evidently not eaten, gnawed, or appar-

ently untouched. After discovering that some acorns lacking external

holes were spoiled by insects or mold, I simplified analysis of results

by recording only whether acorns were opened, regardless of whether

any meat appeared to have been removed. Ranked data were subjected

to the Friedman test (Conover 1980) to test the null hypothesis that

species of acorns were opened in equal numbers.

Food Supplementation — The food supplementation experiment

consisted of trapping at three grids, two on Anderson-Cue (desig-

nated ACI and ACII) and one on Smith Lake (SL). Each grid consisted

of 10 columns and 10 rows 10-m apart, with a single Sherman trap

at each intersection (area = 10,000 m2). Prior trapping at burrows in

each area indicated that mice were present, and grids were set more

than 100-m apart to reduce movements of animals between grids. In

April, May, and June, 1987, I trapped SL for 800 trapnights and

determined that after three consecutive nights of trapping, no addi-

tional individuals were captured. Consequently, SL, ACI, and ACII

were trapped for three consecutive nights per month for a total of

300 trapnights/grid monthly. On ACI I provided a mixture of sun-

flower seeds, mixed bird seed, and oatmeal for 1 year in seven chick

feeders fitted with glass jars. To eliminate non-target species, plastic

buckets with two holes (2.5 cm) cut near the rims were upended

over the feeders and anchored with cinder blocks.

RESULTS

General Observations on Diet — Five feeding events were observed

at Smith Lake. An adult male caught and released on SL (5 May

1988) readily ate a cricket (Orthoptera: Gryllinae) offered to him.

On 15 June 1987 at approximately 0245 EST, an adult female (who

had been trapped and released) immediately caught a small moth

and consumed all but the wings. On 9 May, an adult female just

released from a trap ate a young shoot of wild bamboo (Smilax

auriculata). In July 1988, a female released from a trap at a burrow

hid in a hole at the base of a turkey oak approximately 5 m

northwest of the burrow. In a few minutes she left the hole, picked

up a small pawpaw (Asimina incarna), carried it back to the hole,

and ate it. Jones (1989) previously described consumption of a pawpaw

fruit (A. incarna) by a Florida mouse at Smith Lake. Predation and

dispersal of Asimina fruits have not been well studied, although

134 Cheri A. Jones

these fruits are eaten by opossums (Didelphis virginiana), humans,

and other mammals (Bartram 1791, Willson and Schemske 1980,

Norman and Clayton 1986). Although nutritional values and fruit set

have not been reported for A. incarna, it is the most abundant

pawpaw and the largest fruit produced in sandhills on Ordway and

probably represents a significant addition to the summer diet of

Podomys.

I offered captive animals the following fruits and seeds collected

at Ordway, all of which were eaten: acorns (Q. chapmanii, Q. geminata,

Q. hemisphaerica, Q. laevis, Q. myrtifolia, and Q. nigra), pine seeds

(P. elliottii and P. palustris), blueberries (Vaccinium myrsinites),

deerberries (V. stamineum), gall berries {Ilex galabra), blackberries

(Rubus argutus), gopher apples (Licania michauxii), pawpaw fruits (A.

incarna and A. pygmaea), flowers of queen's delight (Stillingia sylvatica),

and seed pods of legumes (Crotalaria rotundifolia and Galactia elliotti).

Captive mice also shredded seeds and stems of unidentified grasses

and incorporated them into the cotton nesting material in their cages;

the grasses probably were eaten as well.

The ready acceptance of a wide variety of foods in my study

implies that, like Peromyscus, P. floridanus is an opportunistic feeder.

In general, the feeding behavior I observed in captive P. floridanus

resembled that described for Peromyscus (Eisenberg 1968), in which

the animal picks up food with the paws or mouth, then crouches

and manipulates the food with the paws. Large items such as pawpaws

were propped against the substrate. Seed pods were opened by

grasping the pod vertically, resting one end on the substrate, chewing

off one end, and opening the pod longitudinally along a suture.

Larger foods, such as turkey oak acorns, were dragged with the

incisors; smaller items were carried in the mouth. Some captives

consistently cached acorns and sunflower seeds under kitty litter in

corners of the aquaria. Food items and remains also commonly were

found underneath nests.

Except for smaller acorns that might be split in half, acorns

were opened consistently at the hilum (basal scar). Although acorns

typically were carried by the point at the distal end, mice never

chewed open the hull there. For small, round acorns a neat incision

was made around the hilum; on more elongate nuts (Q. geminata

and Q. laevis) the hull was nibbled farther down the sides. Caps, if

present, were removed; there was no significant difference in the

numbers of capped and capless Q. laevis acorns opened.

Predation on vertebrates is probably rare. On Smith Lake in

1983, J. F. Eisenberg (University of Florida, personal observation)

trapped an adult male who detached and ate the posterior part (about

Diet of Florida Mice 135

3.5 cm) of a juvenile red snake (Elaphe guttata) that was caught

half way in the trap. An adult male caught on Anderson-Cue in

October 1987 ate the viscera of a juvenile Florida mouse also caught

in the trap. A litter of three young was eaten on one occasion

when a male and two lactating females were in one aquarium.

Acorn Preferences — Unpredictable acorn supplies made it impos-

sible to run tests with identical acorns in 1988 and 1989. Additionally,

the majority of acorns on the ground already contained weevil larvae

or were otherwise spoiled. In 1988, I concentrated on determining

which acorns were eaten by Podomys. I presented acorns from six

species of oaks — Chapman's (Q. chapmanii), live (Q. geminata), turkey

(Q. laevis), laurel (Q. hemisphaerica), myrtle (Q. myrtifolia), and water

oaks (Q. nigra) — to four captive animals. Five of these species belong

to the red oak group, which generally contains three or four times

more tannin (Briggs and Smith 1989) than species of the white oak

group (which includes Q. chapmanii). Each mouse was presented with

turkey oak acorns in combination with acorns from two other species;

two-four trials were run per animal for a total of 12 trials. Although

the sample was inadequate for statistical analysis, I noted that mice

opened acorns of all species, and in all but one trial turkey oak

acorns were opened in the smallest numbers.

In 1989 Chapman's and live oak acorns were not available, so

I gathered acorns from Q. laevis and two different trees of Q.

hemisphaerica, one from an old pasture near Ross Lake and the

second from a hammock past Anderson-Cue. I expected that acorns

of Q. laevis, the predominant oak on Smith Lake, would be preferred.

I tested nine animals, two trials each. I analyzed the results of the

first trial only, because there was no difference in ranks of first and

second trials. Results indicated that acorns were not opened in equal

numbers (Friedman test, T = 26.75, P = 0.01). For multiple compari-

sons at a significance level of P = 0.01 (Conover 1980), acorns from

Ross Lake (Q. hemisphaerica) were opened significantly more often;

differences between acorns from the hammock and from Q. laevis

were not significant. These results indicate not only a preference for

laurel oak acorns, but an ability to distinguish acorns from two indi-

viduals of Q. hemisphaerica.

Mice did not eat blackened nutmeats, but I did not test

preferences of sound acorns versus acorns with larvae. On one occasion

an adult female immediately ate a larva from an acorn opened but

not eaten by another female. Semel and Andersen (1988) suggested

that such differences in behavior might be due to mice being unable

to detect larvae in unopened acorns, or that larvae are detected but

avoided by some individuals. They also suggested that tooth marks

136

Cheri A. Jones

on hulls and movement of acorns might represent assessment. Of

the 450 acorns presented in 30 trials to 13 Podomys, 66% were

removed from bowls, whether opened or not.

Food Supplementation Experiment — Individual trappability of

Florida mice captured on the Ordway Preserve ranged from 14 to

100% (Jones 1990). The average trappability for 5 years of trapping

was 57%. According to Hilborn et al. (1976), estimates of minimum

numbers of individuals by direct enumeration become more reliable

as trappability exceeds 50%.

AC I GRID

AC II GRID

SL GRID

MONTH

Fig. 1. Minimum number of individuals known alive (MNI) on the

Anderson-Cue I, Anderson-Cue II, and Smith Lake grids in Putnam County,

Florida. Arrows indicate beginning and ending of food supplementation on

Anderson-Cue I.

Results of the food supplementation (Fig. 1) were similar to

those reported by Young (1983) in that Podomys disappeared during

the summer in spite of the additional food. Capture rates declined

sharply on ACI where extra food was provided, although mice persisted

in small numbers at burrows west and east of the grid. Maximum

trapping success for a single night was 9%, and was usually much

lower. Persistence (time between first and last captures) of mice on

the three grids was estimated (Table 1). It seems unlikely that the

decline on ACI was due to competition with immigrating rodents;

the only other rodents captured on these grids were flying squirrels

(Glaucomys volans).

Diet of Florida Mice

137

Table 1. Persistence data for Podomys on trap grids in Putnam County, 1987-88.

Data presented are minimum number known alive, mean + SD days present on

grid, and total number of trapnights.

DISCUSSION

Hulls of Q. laevis found in excavated burrows (Jones and Franz

1990) and vacuumed remains of Q. geminata and Q. laevis from

burrows on Roberts Ranch, Putnam County (E. G. Milstrey, University

of Florida, personal observation) were opened in a manner consistent

with that observed with captive Podomys. Small piles of similarly-

opened hulls occasionally were found at the base of turkey oaks

and near burrow entrances on Ordway.

Acorn selection might be based on chemical composition other

than tannin content. Acorns of Q. hemisphaerica contain more tannic

acid than Q. laevis and more fat and carbohydrates than Q. chapmanii

and Q. incana (Halls 1977, Harris and Skoog 1980). In their study

of acorn preference in Peromyscus, Briggs and Smith (1989) found

that five P. leucopus captured in habitats lacking oaks consumed

equal amounts of acorns from species of the red and white oak

groups, whereas mice from areas containing oaks selected acorns of

Quercus species found in their habitat, independent of fat, protein,

and tannin content.

Based on these preliminary results, I suggest that acorns of

oaks other than Q. laevis are preferred if available. Turkey oaks

provide an unreliable food supply. Umber (1975) noted low, variable

acorn production by Q. laevis in Citrus and Hernando counties, and

Kantola and Humphrey (1990) found that production by trees at

Ordway varied with slope and tree diameter. Layne (1990) correlated

the relatively greater abundance of Podomys in scrub and scrubby

flatwoods with higher and more consistent acorn production than in

sandhills. He also presented evidence that differences in morphology

and behavior in scrub and sandhill populations reflect differences in

vegetation structure and mast production in these habitats. Proximity

to Q. geminata and other oak species might partially explain the

relatively higher and more stable population of Podomys at Smith

Lake, although there is no evidence that populations are food limited.

138 Cheri A. Jones

If oak species in hammocks are more reliable producers, the distribution

of P. gossypinus and Ochrotomys nuttalli in hammocks at Ordway

might be partly due to the acorn supply. Competition could be one

factor restricting Podomys to sandhills when other rodents are present

in contiguous habitats. However, in sandhills and other habitats

elsewhere in Florida, P. floridanus P. gossypinus, and O. nuttalli are

sympatric (Packer and Layne 1991, Frank and Layne 1992).

ACKNOWLEDGMENTS— I thank the Florida Game and Fresh Water

Fish Commission for permits that allowed this work; the Florida

Museum of Natural History for space for the captive colony; and J.

F. Eisenberg, R. Franz, J. N. Layne, and an anonymous referee for

comments on an earlier draft. J. F. Anderson, L. S. Fink, W. H.

Kern, R. F. Labisky, C. A. Langtimm, P. Ryschkewitsch, and C. A.

Woods also provided assistance. The original version of this paper

appeared in a dissertation presented to the University of Florida.

LITERATURE CITED

Bangs, O. 1898. The land mammals of peninsular Florida and the coast

region of Georgia. Proceedings of the Boston Society of Natural

History 28:157-235.

Bartram, W. 1791. The travels of William Bartram. (1988 reprint) Penguin

Books, New York, New York.

Brand, S. B. 1987. Small mammal communities and vegetative structure

along a moisture gradient. MS Thesis, University of Florida, Gainesville.

Briggs, J. M., and K. G. Smith. 1989. Influence of habitat on acorn

selection by Peromyscus leucopus. Journal of Mammalogy 70:35-43.

Conover, W. J. 1980. Practical nonparametric statistics. John Wiley and

Sons, New York, New York.

Eisenberg, J. F. 1968. Behavior patterns. Pages 451-495 in Biology of

Peromyscus (Rodentia) (J. A. King, editor.) Special Publication 2,

American Society of Mammalogists.

Eisenberg, J. F. 1988. Mammalian species of the Ordway Preserve. A

reference for students. Ordway Preserve Research Series, Florida

Museum of Natural History Report No. 1, Gainesville.

Frank, P. A., and J. N. Layne. 1992. Nests and daytime refugia of cotton

mice {Peromyscus gossypinus) and golden mice {Ochrotomys nuttalli)

in south-central Florida. The American Midland Naturalist 127:21-30.

Franz, R. 1986. Gopherus polyphemus (gopher tortoise) burrow

commensals. Herpetological Review 17:64.

Franz, R. 1990. Annotated list of the vertebrates of the Katharine

Ordway Preserve-Swisher Memorial Sanctuary, Putnam County, Florida.

Ordway Preserve Research Series, Florida Museum of Natural

History Report Number 2, Gainesville.

Diet of Florida Mice 139

Halls, L. K., editor. 1977. Southern fruit-producing woody plants used by

wildlife. United States Department of Agriculture, Forest Service

General Technical Report SO-16.

Harris, L. D., and P. J. Skoog. 1980. Some wildlife habitat-forestry

relations in the southeastern Coastal Plain. Annual Forestry

Symposium Proceedings 29:103-119.

Hilborn, R., J. A. Redfield, and C. J. Krebs. 1976. On the reliability of

enumeration for mark and recapture census of voles. Canadian

Journal of Zoology 54:1019-1024.

Humphrey, S. R., J. F. Eisenberg, and R. Franz. 1985. Possibilities for

restoring wildlife of a longleaf pine savanna in an abandoned citrus

grove. Wildlife Society Bulletin 13:487-496.

Jones, C. A. 1989. First record of pawpaw consumption by the Florida

mouse. Florida Scientist 52:7.

Jones, C. A. 1990. Microhabitat use by Podomys floridanus in the high

pine lands of Putnam County, Florida. Ph.D. Dissertation, University

of Florida, Gainesville.

Jones, C. A., and R. Franz. 1990. Use of gopher tortoise burrows by

Florida mice {Podomys floridanus) in Putnam County, Florida. Florida

Field Naturalist 18:45-51.

Jones, C. A., and J. N. Layne. In Press. Podomys floridanus. Mammalian

Species.

Kantola, A. T., and S. R. Humphrey. 1990. Habitat use by Sherman's fox

squirrel (Sciurus niger shermani) in Florida. Journal of Mammalogy

71:411-419.

Layne, J. N. 1970. Climbing behavior of Peromyscus floridanus and

Peromyscus gossypinus. Journal of Mammalogy 51:580-591.

Layne, J. N. 1978. Florida mouse. Pages 21-22 in Rare and endangered

biota of Florida. Volume 1: Mammals (J. N. Layne, editor). University

Presses of Florida, Gainesville.

Layne, J. N. 1990. The Florida mouse. Pages 1-21 in Proceedings of the

eighth annual meeting gopher tortoise council (C. K. Dodd, Jr., R. E.

Ashton, Jr., R. Franz, and E. Wester, editors). Florida Museum of

Natural History, Gainesville.

Merriam, C. H. 1890. Descriptions of twenty-six new species of North

American mammals. North American Fauna 4:1-54.

Milstrey, E. G. 1987. Bionomics and ecology of Ornithodoros (P.)

turicata americanus (Marx) (Ixodoidea: Argasidae) and other

commensal invertebrates present in the burrows of the gopher tortoise,

Gopherus polyphemus Daudin. Ph.D. Dissertation, University of

Florida, Gainesville.

Norman, E. M., and D. Clayton. 1986. Reproductive biology of two

Florida pawpaws: Asimina obovata and A. pygmaea (Annonaceae).

Bulletin of the Torrey Botanical Club 113:16-22.

Packer, W. C, and J. N. Layne. 1991. Foraging site preferences and

relative arboreality of small rodents in Florida. American Midland

Naturalist 125:187-194.

140 Cheri A. Jones

Semel, B., and D. C. Andersen. 1988. Vulnerability of acorn weevils

(Coleoptera: Curculionidae) and attractiveness of weevils and infested

Quercus alba acorns to Peromyscus leucopus and Blarina brevicauda.

American Midland Naturalist 119:385-393.

Umber, R. W. 1975. Impact of pine plantation succession on sandhill

wildlife in central Florida. MS Thesis, University of Florida, Gainesville.

Willson, M. F., and D. W. Schemske. 1980. Pollinator limitation, fruit

production, and floral display in pawpaw (Asimina triloba). Bulletin

of the Torrey Botanical Club 107:401-408.

Young, B. L. 1983. Food supplementation of small rodents in the sand

pine scrub. MS Thesis, University of Central Florida, Orlando.

Young, B. L., and J. Stout. 1986. Effects of extra food on small rodents

in a south temperate zone habitat: demographic responses. Canadian

Journal of Zoology 64:1211-1217.

Accepted 1 September 1992

141

BRIMLEYANA

A Journal of Zoology of the Southeastern United States

INVITATION FOR PAPERS

The editorial staff of Brimleyana welcomes contributions. Research

papers based on studies conducted in the southeastern United States

are invited. Areas of interest include systematics, evolution, zoogeography,

ecology, behavior, and paleozoology. Brief communications, as well

as articles of more typical length, will be considered. Page charges

will be waived for those who lack institutional or grant support. Inquiries

should be directed to:

Editor, Brimleyana

North Carolina State Museum of Natural Sciences

P.O. Box 27647

Raleigh, NC 27611

NEW EDITORS

Richard A. Lancia, a professor in the Department of Forestry at

North Carolina State University in Raleigh, is the new Editor of

Brimleyana. A specialist in ecology and wildlife management, he is a

former editor-in-chief of The Journal of Wildlife Management.

Dr. Lancia holds a B.S. in Wildlife Management from the University

of Michigan, an M.A. in Zoology from Southern Illinois University,

and a Ph.D. in Wildlife Biology from the University of Massachusetts.

He is a member of the American Ornithologists' Union, the American

Society of Mammalogists, and The Wildlife Society. A charter member

of the North Carolina Chapter of The Wildlife Society, Dr. Lancia is

a past president of that organization. He has also served as president

of the N.C. State University Chapter of Sigma Xi. Dr. Lancia is the

principal author of a chapter on estimating animal population sizes in

a manual on wildlife management techniques soon to be published by

The Wildlife Society.

Suzanne Fischer is the Assistant Editor of Brimleyana. Ms. Fischer

holds a B.A. in English and an M.S. in Technical Communication,

both from North Carolina State University. She is a Communications

Specialist with CompuChem Laboratories, Inc., in the Research Triangle

Park. In addition to her work on Brimleyana, Ms. Fischer is serving

as editor for several parts of the Endangered, Threatened, and Rare

Fauna series of the Occasional Papers of the North Carolina Biological

Survey.

142

ENDANGERED, THREATENED, AND

RARE FAUNA OF NORTH CAROLINA

PART III. A RE-EVALUATION OF THE BIRDS

Edited by

David S. Lee and James F. Parnell

This book is a report prepared by a committee appointed in 1985 by

the North Carolina State Museum of Natural Sciences to re-evaluate the list

of birds presented in Endangered and Threatened Plants and Animals of

North Carolina (John E. Cooper, Sarah S. Robinson, and John B. Funderburg,

editors. N.C. State Mus. Nat. Hist., Raleigh, 1977), which is now out of

print. Committee members were David S. Lee, James F. Parnell, Allen

Boynton, Philip J. Crutchfield, Tom Howard, E. Wayne Irvin, Harry E.

LeGrand, Jr., Eloise F. Potter, and Jeffrey R. Walters. Detailed accounts

update the status of 21 North Carolina avian species that are Federally

Endangered, Federally Threatened, or in danger of being extirpated as breed-

ing birds within the state. Biological and environmental problems are dis-

cussed briefly for 43 species that are not of immediate concern.

1990 52 pages Softbound ISBN 0-917134-19-2

Price: $8 postpaid. North Carolina residents add 6% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: ETR BIRDS, N.C. State Museum of Natural Sciences,

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

143

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lewesi (Brimley) (Amphibia: Proteidae).

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citation is by the same author(s). For example:

Adams, J. J. 1977. Food habits of the masked shrew, Sorex cinereus

(Mammalia: Insectivora). Brimleyana 7:32-39.

Adams, J. J. 1988. Animals in North Carolina folklore. Second edition.

University of North Carolina Press, Chapel Hill.

Barnes, R. G. 1986. Range, food habits, and reproduction in Glaucomys

sabrinus in the southern Appalachian Mountains of North Carolina

and Tennessee. Ph.D. Thesis. North Carolina State University, Raleigh.

Barnes, R. G. 1989. Northern flying squirrel. Pages 203-230 in Mammals

of the southeastern United States (J. J. Adams and J. M. Smith, Jr.,

editors). Harper and Row, New York, New York.

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INFORMATION FOR CONTRIBUTORS (Continued)

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BRIMLEYANA NO. 18, JUNE 1993

CONTENTS

The Myriapod Types of Oscar Harger (Arthropoda: Diplopoda, Chilopoda).

Rowland M. Shelley 1

A Late Pleistocene Vertebrate Assemblage from the St. Marks River,

Wakulla County, Florida. Timothy S. Young and Joshua Laerm 15

No Decline in Salamander (Amphibia: Caudata) Populations:

A Twenty-Year Study in the Southern Appalachians.

Nelson G. Hairston, Sr., and R. Haven Wiley 59

On the Validity of the Name teyahalee as Applied to a Member of the Plethodon

glutinosus Complex (Caudata: Plethodontidae): A New Name.

Nelson G. Hairston, Sr 65

Differences in Variation in Egg Size for Several Species of Salamanders

(Amphibia: Caudata) That Use Different Larval Environments.

Christopher King Beachy 71

Helminth Parasites of the Eastern Box Turtle, Terrapene Carolina Carolina (L.)

(Testudines: Emydidae), in North Carolina.

Michael D. Stuart and Grover C. Miller 83

Notes on the Spiny Softshell, Apalone spinifera (Testudines: Trionychidae),

in Southeastern Virginia. Joseph C. Mitchell and Ronald Southwick 99

Range Expansion of the Tree Swallow, Tachycineta bicolor (Passeriformes:

Hirundinidae), in the Southeastern United States. David S. Lee 103

An Observation of Fea's Petrel, Pterodroma feae (Procellariiformes: Procellariidae),

Off the Southeastern United States, With Comments on the Taxonomy and

Conservation of Soft-plumaged and Related Petrels in the Atlantic Ocean.

J. Christopher Haney, Craig A. Faanes, and William R. P. Bourne 115

Cotton Mice, Peromyscus gossypinus LeConte (Rodentia: Cricetidae),

in the Great Dismal Swamp and Surrounding Areas.

James L. Boone and Joshua Laerm 125

Observations Regarding the Diet of Florida Mice, Podomys floridanus

(Rodentia: Muridae). Cheri A. Jones 131