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N.C. DOCUMENTS

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number 24

april 1997

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 29555, Raleigh,

NC 27626. 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 29555, Raleigh, NC 27626.

In citations please use the full name - Brimleyana.

North Carolina State Museum of Natural Sciences

Betsy Bennett, Director

North Carolina Department of Environment,

Health, and Natural Resources

James B. Hunt Jr., Governor

Jonathan B. Howes, Secretary

CODN BRIMD 7

ISSN 0193-4406

The Rock Shrew, Sorex dispar (Insectivora: Soricidae), in

Georgia with Comments on its Conservation Status in the

Southern Appalachians

Joshua Laerm, Charles H. Wharton,

Museum of Natural History and Institute of Ecology

University of Georgia, Athens, Georgia 30602

AND

William Mark Ford

Westvaco, Timberlands Division,

Box 577, Rupert, West Virginia 25984

ABSTRACT — The first state record of Sorex dispar is reported from

Georgia in a high elevation cliff and talus mixed-oak community

in Rabun County. New records from localities in Macon County,

North Carolina, are also reported. The conservation status of the

species is uncertain in the southern Appalachians where collection

records indicate it to be rare.

On 29 October 1995 one adult male rock shrew, Sorex dispar

Batchelder, was found in a sunken pitfall trap on the north face of

Rabun Bald, Rabun County, Georgia, at an elevation of 1,280 m. Pitfalls

were 946 cm3 plastic cups (11-cm lip diameter and 14-cm depth) set

flush to the ground adjacent to fallen logs, rocks, stumps, or other

forest floor debris. The specimen was captured under a protruding

gneiss boulder in a cliff and talus slope at the base of a massive

rock face which dominates the north face of Rabun Bald. Standard

body measurements were 129, 63, 15 mm. This is the first record

of the species from Georgia and represents an extension of its range

approximately 50 km south from its nearest reported locality in Jackson

County, North Carolina (Webster 1987).

The Rabun Bald locality is dominated by a chestnut oak (Quercus

prinus), northern red oak {Q. rubra), red maple {Acer rubrum), and

black birch (Betula lenta) overstory. Witch-hazel (Hamamelis virginiana),

rosebay rhododendron {Rhododendron maximum), sweet pepper bush

{Clethra acuminata), and fetter bush {Leucothoe recurva) dominate

the shrub layer. Other small mammals recovered in pitfalls at the

locality included Sorex cinereus, S. fumeus, Blarina brevicauda, Peromyscus

maniculatus, Microtus pinetorum, and Clethrionomys grapperi.

We previously collected three S. dispar specimens in Macon County,

North Carolina which, owing to the rarity of the species, we report

Brimleyana 24:1-5, April 1997

Joshua Laerm, C. H. Wharton, and W. M. Ford

on here. One (male; 124, 63, 15) was collected under a boulder on

5 February 1994 adjacent to Turtle Pond Road, 0.4 km east of Turtle

Pond Creek, 0.5 km west of US Highway 64. Rock outcrops dominate

this north facing slope at an elevation of 1,050 m about 50 m above

Turtle Pond Creek. The vegetational community consisted of hemlock

(Tsuga canadensis), white pine (Pinus strobus), and red maple with

a rosebay rhododendron understory. Two additional specimens (both

males; 120, 65, 16 and 124, 65, 16 mm) were taken on the same

date, approximately 3 km distance southwest at Turtle Pond Road,

1.4 km north of NC Highway 106. This community was markedly

more xeric, dominated by a white oak (Q. alba), chestnut oak, and

hemlock overstory with mountain laurel {Kalmia latifolia) and blueberry

(Vaccinium spp.) shrub layer. The site, approximately 100 m above

Turtle Pond Creek at an elevation of 1,120 m, was not markedly rocky,

and the shrew was taken in a pitfall trap set along a fallen tree. At

both of these localities S. dispar was taken in association with B.

brevicauda, S. cinereus, S. fumeus and C. gapperi. Specimens were

reposited in the University of Georgia Museum of Natural History.

Sorex dispar is endemic to the Appalachian Mountains and is

distributed from New Brunswick south. Regionally it is reported from

Maryland (Paradiso 1969, North Carolina State Museum records; S.

D. Lee, personal communication), Virginia (Handley 1956, 1979, 1991;

Holloway 1957; Pagels and Tate 1976; Pagels 1987, 1991; Kaldo and

Handley 1993), Kentucky (Caldwell 1980, Caldwell and Bryan 1982,

Bryan 1991), Tennessee (Conaway and Pfitzer 1952, Tuttle 1968, Linzey

and Linzey 1971, Smith et al. 1974, Kennedy and Harvey 1980, Harvey

et al. 1992), North Carolina (Schwartz 1956, Lee et al. 1982, Webster

1987), and now Georgia.

Once regarded as very rare in the central and southern Appalachian,

S. dispar is now believed to be more widely distributed and occurs

in a broader range of habitats than previously supposed (Kirkland et

al. 1976; Kirkland and Van Deusen 1979; Kirkland et al. 1979; Kennedy

and Harvey 1980; Handley 1979, 1991; Pagels 1987; Kalko and Handley

1993). Although no population estimates are available, published records,

available museum specimens, and trapping records suggest that it is

uncommon to rare throughout most of its range in the extreme southern

Appalachians, but that it may be locally abundant in the central Appala-

chians. For example, over a 15-year period at Mountain lake, Giles

County, Virginia, Kalko and Handley (1993) report S. dispar to comprise

10% of the total number of long-tailed shrews recovered and indicate

it is common in its preferred habitat (Handley 1979, 1991; C. O. Handley,

personal communication). Similarly, Pagels (1987) notes it to be more

Rock Shrew

locally abundant elsewhere in Virginia than previously believed. However,

recent survey data south of Virginia suggest it is rare. Harvey et al.

(1992) report only 11 individuals were recovered in 389,995 combined

pitfall and snap trap-nights of effort on the Northern District of the

Cherokee National Forest (Unicoi, Johnson, Carter, Greene, and Sullivan

counties) of eastern Tennessee. South of the Great Smoky Mountains

National Park, Harvey et al. (1991) reported none was recovered in

233,567 combined trap-nights in the Southern District of the Cherokee

National Forest (Polk, McMinn, and Monroe counties, Tennessee). Else-

where, in the southern Blue Ridge of western North Carolina, northern

Georgia, and northwestern South Carolina, we recovered only the four

individuals reported upon here based upon 175,000 combined pitfall

and snap trap-nights of effort. We conclude that in the extreme southern

Appalachians the species appears to be rare or extremely localized.

Additional survey efforts are required to determine the precise habitat

associations and status of the species at the southern limit of its range.

ACKNOWLEDGMENTS— This study was supported through a

cooperative funding agreement between the United States Department

of Agriculture Forest Service, Chattahoochee National Forest, the University

of Georgia Museum of Natural History, and National Science Foundation

grant BSR 9011661. Specimens were collected under authority of Georgia

Scientific Collecting Permit 29-000089 and North Carolina Scientific

Collecting Permit 95-ES-10.

LITERATURE CITED

Bryan, H. D. 1991. The distribution, habitat and ecology of shrews (Soricidae:

Blarina, Sorex and Cryptotis) in Kentucky. Journal of the Tennessee

Academy of Science 66:187-189.

Caldwell, R. S. 1980. First records of Sorex dispar and Microsorex thompsoni

in Kentucky with distributional notes on associated species. Transac-

tions of the Kentucky Academy of Science 41:46-47.

Caldwell, R. S., and H. Bryan. 1982. Notes on the distribution and hab-

its of Sorex and Microsorex (Insectivora: Soricidae) in Kentucky. Brimleyana

8:91-100.

Conaway, C. S., and D. W. Pfitzer. 1952. Sorex palustris and Sorex dispar

from the Great Smoky Mountains National Park. Journal of Mammal-

ogy 33:106-108.

French, T. W., and G. L. Kirkland, Jr. 1983. Taxonomy of the Gasp shrew,

Sorex gaspensis, and the rock shrew, Sorex dispar. Canadian Field-

Naturalist 97:75-78.

Hall, E. R. 1981. The mammals of North America. Second edition. John

Joshua Laerm, C. H. Wharton, and W. M. Ford

Wiley & Sons, New York, New York.

Handley, C. O., Jr. 1956. The shrew, Sorex dispar, in Virginia. Journal

of Mammalogy 37:435.

Handley, C. O., Jr. 1979. Sorex dispar dispar Batchelder. Pages 541-544

in Endangered and threatened plants and animals of Virginia (D. M.

Linzey, editor). Center for Environmental Studies, Virginia Polytechnical

Institute and State University, Blacksburg.

Handley, C. O., Jr. 1991. Mammals. Pages 539-613 in Virginia's endan-

gered species (K. Terwilliger, editor). McDonald and Woodward, Blacks-

burg, Virginia.

Harvey, M. J., C. S. Chaney, and M. D. McGimsey. 1991. Distribution,

status, and ecology of small mammals of the Cherokee National For-

est, Tennessee (Southern Districts). Report to the United States For-

est Service. Manuscript on file, Center for the Management, Utilization,

and Protection of Water Resources, Tennessee Technological Univer-

sity, Cookville.

Harvey, J. J., M. D. McGimsey, and C. S. Chaney. 1992. Distribution,

status, and ecology of small mammals of the Cherokee National For-

est, Tennessee (Northern Districts). Report to the United States For-

est Service. Manuscript on file, Center for Management, Utilization,

and Protection of Water Resources, Tennessee Technological Univer-

sity, Cookville.

Holloway, C. O. 1957. Sorex dispar at Mountain Lake Virginia. Journal

of Mammalogy 38:406.

Kalko, E. K. V., and C. O. Handley, Jr. 1993. Comparative studies of

small mammal populations with transects of snap traps and pitfall arrays

in southwestern Virginia. Virginia Journal of Science 44:3-18.

Kennedy, M. L., and M. J. Harvey. 1980. Mammals. Pages 1-50 in Ten-

nessee Rare Vertebrates (D. C. Eager and R. M. Hatcher, editors).

Tennessee Wildlife Resources Agency and Tennessee Department of

Conservation, Nashville.

Kirkland, G. L., Jr., C. R. Schloyer, and D. K. Hull. 1976. A novel habitat

record for the long-tailed shrew, Sorex dispar. Proceedings of the West

Virginia Academy of Science 48:77-79.

Kirkland, G. L., Jr., D. F. Schmidt, and C. J. Kirkland. 1979. A novel

habitat record for the long-tailed shrew {Sorex dispar) in New Brunswick.

Canadian Field-Naturalist 93:195-197.

Kirkland, G. L., Jr., and H. M. Van Deusen. 1979. The shrews of the

Sorex dispar group: Sorex dispar Batchelder and Sorex gaspensis An-

thony and Goodwin. American Museum Novitates 2675:1-21.

Lee, S. D., J. B. Funderburg, Jr., and M. K. Clark. 1982. A distribu-

tional survey of North Carolina mammals. Occasional Papers of the

North Carolina Biological Survey, North Carolina State Museum of Natural

Sciences, Raleigh.

Linzey, A. V., and D. W. Linzey. 1971. The mammals of the Great Smoky

Mountains National Park. University of Tennessee Press, Knoxville.

Rock Shrew

Pagels, J. F. 1987. The pygmy shrew, rock shrew and water shrew: Virginia's

rarest shrews (Mammalia: Soricidae). Virginia Journal of Science 38:364—368.

Pagels, J. F. 1991. A high elevation record for the least shrew, Cryptotis

parva (Say). Virginia Journal of Science 42:361-362.

Pagels, J. F., and C. M. Tate. 1976. Shrews (Insectivora: Soricidae) of

the Paddy Knob-Little Back Creek Area of western Virginia. Virginia

Journal of Science 27:202-203.

Paradiso, J. L. 1969. Mammals of Maryland. North American Fauna 66:1-193.

Schwartz, A. 1956. A new subspecies of the long-tailed shrew (Sorex dispar

Batchelder) from the southern Appalachian Mountains. Journal of the

Elisha Mitchell Scientific Society 72:24-30.

Smith, C. R., J. Giles, M. E. Richmond, J. Nagel, D. W. Lambert. 1974.

The mammals of northeastern Tennessee. Journal of the Tennessee Academy

of Science 49:88-94.

Tuttle, M. D. 1968. First Tennessee record of Mustela nivalis. Journal of

Mammalogy 49:133.

Webster, W. D. 1987. Sorex dispar. Pages 39-40 in Endangered, threat-

ened and rare fauna of North Carolina. Part 1. A reevaluation of the

mammals (M. K. Clark, editor). Occasional Papers of the North Carolina

Biological Survey, North Carolina State Museum of Natural Sciences,

Raleigh.

Received 7 December 1995

Accepted 6 March 1995

Digitized by the Internet Archive

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http://archive.org/details/brimleyana19nort_12

Dog Burials from the Eighteenth Century Cherokee Town

of Chattooga, South Carolina

Gerald F. Schroedl

Department of Anthropology, University of Tennessee,

Knoxville, Tennessee 37996-0720

AND

Paul W. Parmalee

Frank H. Mclung Museum, University of Tennessee

Knoxville, Tennessee 37996-3200

ABSTRACT — Archaeological excavations recovered the remains of three

dogs from two pit-features at the eighteenth century village site of

Chattooga in South Carolina. The three individuals were small to

medium-sized animals. Observations on one animal (Dog 3) indi-

cate extreme age at death, suggesting that the dog was given spe-

cial care during its life. These occurrences are consistent with ar-

chaeological and historical information about the role of dogs in

Cherokee society.

Archaeological investigations at the historic eighteenth century

town of Chattooga, Oconee County, South Carolina, recovered the remains

of three dogs which were deliberately interred in pit-features (Schroedl

1995). These burials are an example of a practice documented at other

historic Cherokee sites. Analysis of the skeletal remains shows that

some dogs were so incapacitated by old age that they must have received

special care for them to have lived so long. Intentional burial also

attests to the regard afforded these animal regardless of their age at

death.

Reported from two village sites in East Tennessee are three historic

period Cherokee dogs and four additional skeletons which may represent

historic Cherokee or late prehistoric Mississippian period (A.D. 1400

to 1600) associations (Parmalee and Bogan 1978; see also Bogan 1976,

1980, 1983; Bogan et al. 1986). Each animal was about the size of

a beagle and was deliberately buried. Significantly, Dog Burial 1 at

the Chota site was an older animal, which, because it was arthritic

and had a deformed right hind foot, must have received special care

during its life (Parmalee and Bogan 1978:105). Isolated elements of

domestic dogs are infrequent in late prehistoric and historic faunal

samples in East Tennessee, suggesting that the Cherokee seldom ate

dogs and infrequently discarded them with refuse. In other areas of

Brimleyana 24:7-14, April 1997

Gerald F. Schroedl and Paul W. Parmalee

the Southeast, dogs may have been used more frequently as a food

source (Mooney 1900:26) or consumed in ritual contexts (Swanton

1911:129). In general, however, wide-spread use of dogs for food or

ritual was uncommon in the Southeast.

Cherokee use of dogs is poorly documented, but the animals were

obviously kept as pets, sometimes eaten, and perhaps used in hunting

as recorded for other southeastern groups (Swanton 1946:345). Southeast

Indians once may have raised distinctive breeds of dogs, but soon

after historic contact most dogs were probably hybrids of European

and aboriginal animals (Parmalee and Bogan 1978:100-101). Most dogs

probably scavenged for food in village areas and received infrequent

handouts. The animals were tolerated but generally not provided great

care except for the occasional individual that was treated with some

respect or reverence such as the ones archaeologically represented by

intentional burial.

Despite their marginal role in Cherokee economic life, dogs also

are represented in myths and supernatural beliefs. The howling of a

family dog, for example, was an omen of sickness and death in the

family (Mooney and Olbrechts 1932:37). Dogs also played a prominent

role in myths about the great deluge and the creation of the Milky

Way (Mooney 1900:259, 261). Another story describes how dogs were

once wild, and how they replaced wolves who were once domesticated.

In Cherokee sacred formulas, dogs sometime occur as a metaphor for

spiritual healing. For example, spiritual deer chief, the cause of rheumatism,

is overcome by the spirit of the dog who is more powerful and the

natural enemy of the deer (Mooney 1886:346-347).

THE CHATTOOGA DOG REMAINS

In 1984, test excavations were made in the area of a domestic

structure and the deteriorated skulls of two animals (Dog Burial 1

and Dog Burial 2) were recovered from a pit-feature (Feature 3) associated

with the building. The pit measured 50 by 70 cm and 11 cm deep

(Elliot 1984:30). The position of the skulls on the pit floor and the

size of the pit suggest that the animals had been placed there together.

Decomposition of the postcranial skeletons of both animals was so

complete that none of these bones was observed or recovered for study.

In 1994, excavations at Chattooga in the vicinity of the village

council house or townhouse, approximately 500 m from the area studied

in 1984, revealed a second pit-feature containing the remains of a

single animal (Dog Burial 3) (Fig. 1). This pit, Feature 11, measured

86 cm long, 70 cm wide, and 42 cm deep. It may have been originally

dug for another purpose because it was much deeper than needed to

Cherokee Dog Burials

Fig. 1. Dog Burial 3 in situ, note extreme flexure of neck, view south (photograph

by Gerald F. Schroedl).

accommodate the dog, and approximately 20 cm of fill had accumulated

in the pit when the animal was interred. The position of the bones

indicates that the animal was laid on its right side so that its back

followed the curvature of the pit wall. The dog's head was bent forward

and under the animal's neck. It is impossible to determine whether

this was done after the dog was dead or whether this had caused its

death. Given the relationship between the skull and the pit wall, it

appears that this was done because the dog's neck was too long to

fit the burial pit.

The animal's bones were in poor condition, but most of the skull

was recovered and could be reconstructed (Figs. 2 and 3). Elements

of the postcranial skeleton were recovered, but none was well enough

preserved to obtain measurements or to identify any anomalies or bone

pathologies that might have been present. At some time after the dog's

interment, a prepared clay hearth was built partially covering the pit

outline. It is impossible to determine if this event was behaviorally

related to the dog's death and burial.

Dog Burial 1

The remains of this individual consisted of isolated teeth, cranial

fragments, and sections of both lower jaws with several teeth in place

10

Gerald F. Schroedl and Paul W. Parmalee

Table 1. Skull, mandible, and tooth measurements (mm) of dog burials re-

covered at the Chattooga site, South Carolina (format follows Hagg 1948).

Cherokee Dog Burials

11

Fig. 2. Occlusal view of the skull and lower mandible of Dog 3, showing

tooth loss and extreme cusp wear (photograph by Miles Wright).

o

L &,

Fig. 3. Right lateral view of the skull and mandible of Dog 3, showing tooth

wear and bone lesions around the roots of P34, M1, P , and M, 2 (photo-

graph by Miles Wright).

12 Gerald F. Schroedl and Paul W. Parmalee

in each. Although fragmentary, measurements on the upper as well

as lower premolar P4 and molars ( M1? M., and M3), and observations

of rounding and wear on the molar, indicate that this was a mature

individual (Table 1).

Dog Burial 2

The remains of this individual also consisted of fragmentary portions

of the cranium, incomplete and broken pieces of the upper and lower

jaws, and isolated teeth and tooth fragments. Measurements of the

teeth and alveoli indicate that this also was a mature dog with a skull

similar in size to Dog Burial 1 (Table 1).

Dog Burial 3

Except for the skull and mandibles, Dog Burial 3 also was poorly

preserved. None of the axial skeleton was complete enough to obtain

measurements, so the stature of the animal could not be determined.

Recovered fragments or sections of the postcranial skeleton included

seven cervical and two thoracic vertebrae, one scapula, both ulnae,

one radius, one humerus, one tibia, one femur (represented by the

head), acetabulum, and six elements from one foot.

The loss of many teeth and the wear pattern on those remaining

suggest the animal was quite old when it died. All incisors in both

upper and lower jaws, plus the right first premolar in both, were lost

and alveoli completely absorbed. Only the root of the left P1 remained.

Both right and left P2 3 were crowded and overlapped. Except for a

fragment of a root of the left P4 and the worn base of the hypocone

of the left M1, the left P4, M1, and M2 had been lost or worn away;

most of the alveoli of the molar roots had been absorbed. It is apparent

that with the loss of these teeth, important in tearing, crushing, and

chewing food, the dog was forced to use the right side for mastication.

All remaining cheek teeth on the right side in both upper and

lower jaws exhibit extreme wear (see Figs. 2 and 3). This is especially

noticeable when observing the greater degree of wear on all teeth on

the right side in both the upper and lower jaws compared with those

on the left side. The occlusal patterns of the right M12 were completely

worn away. Only the smooth base of the M2 hypocone remained, and

the surface wear on the M1 had been so intense as to not only erode

away the cusps but also to narrow the tooth in the hypocone/protocone

area. The right C1 also exhibited greater wear than the left C1, being

5.0 mm shorter. The right C1 had a pronounced groove on the lingual

surface, possibly resulting from continual pulling at or chewing of

Cherokee Dog Burials 13

food on the right side. Both canine teeth in the lower mandibles were

worn down to smooth nubs, exposing the nerve canals, and apparently

projected little beyond the gum line. In addition to tooth wear and

loss, the animal suffered from several gum lesions or abscesses, judging

by enlargement of alveoli of the right P3 4, M1, P , and M .

CONCLUSION

The cranial proportions of Dog Burial 3 are very similar to those

of a beagle, although the muzzle is slightly broader and the rami of

the mandibles are somewhat more massive. The dentition exhibits extreme

wear, loss, and abscessing, an indication of the animals advanced age.

Poor preservation of the postcranial skeleton prohibited determination

of stature. However, the most complete limb element, a 103.0 mm

section of the left ulna, including most of the semilunar notch, approximates

the proportion of a forelimb of a beagle-sized dog. This compares favorably

with the stature of the dogs recovered at Chota, especially Dog Burial

1 (see Parmalee and Bogan 1978: Table 1). This dog also was infirm

when it died, attesting to the care both must have received as they

aged. The fragmented skulls and dentition of Dog Burials 1 and 2

at Chattooga also represent mature individuals of comparable size. These

data suggest that there was little size variability in historic Cherokee

dogs.

Intentional burial of dogs by Native Americans in eastern North

America is well documented, this trait beginning over 7,000 years

ago (Morey and Wiant 1992). This implies that at least some individuals

attached a special meaning or feeling for a particular animal. Ethnographic

accounts, however, provide sparse information on the kinds of "breeds"

of dogs kept by southeastern groups, especially the Cherokee. Dogs

probably played a minimal role in the Cherokee economy, but they

were appropriately represented in social and ceremonial life as respected

spiritual forces. Intentional burial of dogs at Chattooga thus is consistent

with the archaeological and ethnographic occurrence of dogs in the

Southeast.

ACKNOWLEDGMENTS — Archaeological and ethnohistorical research

at the Chattooga Site were sponsored by the United States Forest Service

and the University of Tennessee, Knoxville. We thank Robert Morgan

and James Bates of the Forest Service and Sharon Perkul of the Institute

of Archaeology and Anthropology, University of South Carolina, for

making the dogs recovered in 1984 available for study.

14 Gerald F. Schroedl and Paul W. Parmalee

LITERATURE CITED

Bogan, A. E. 1976. A zooarchaeological analysis of vertebrate remains from

Chota-Tanasi, a historic Cherokee village in East Tennessee. M.A. Thesis.

University of Tennessee, Knoxville.

Bogan, A. E. 1980. A comparison of late prehistoric Dallas and Overhill

Cherokee subsistence strategies in the Little Tennessee River Valley.

Ph.D. Dissertation. University of Tennessee, Knoxville.

Bogan, A. E. 1983. Faunal remains from the historic Cherokee occupation

at Citico (40MR7), Monroe County, Tennessee. Tennessee Anthropologist

8: 28-49.

Bogan, A. E., L. LaValley, and G. F. Schroedl. 1986. Faunal remains. Pages

469-514 in Overhill Cherokee archaeology at Chota-Tanasee (G. F. Schroedl,

editor) Department of Anthropology, Report of Investigations 38, Uni-

versity of Tennessee, Knoxville.

Elliot, D. T. 1984. Archaeological Testing of Five Sites on the Sumter Na-

tional Forest, South Carolina. Cultural Resources Report 85-4, Francis

Marion and Sumter National Forests. United States Forest Service, Columbia,

South Carolina.

Haag, W. G. 1948. An osteometric analysis of some aboriginal dogs. Publi-

cations of the Department of Anthropology, University of Kentucky, Reports

in Anthropology 7:107-264.

Mooney, J. 1886. The sacred formulas of the Cherokees. Bureau of Ameri-

can Ethnology, Seventh Annual Report. Washington D.C.

Mooney, J. 1900. Myths of the Cherokee. Bureau of American Ethnology,

Nineteenth Annual Report, Washington D.C.

Mooney, J., and F. Olbrechts. 1932. The Swimmer manuscript: Cherokee sacred

formulas and medicinal prescriptions. Bureau of American Ethnology,

Bulletin 99, Washington D.C.

Morey, D. F., and M. Wiant. 1992. Early Holocene domestic dog burials

from the North American Midwest. Current Anthropology 33:224-229.

Parmalee, P. W., and A. E. Bogan. 1978. Cherokee and Dallas dog burials

from the Little Tennessee River Valley. Tennessee Anthropologist 3:100-

112.

Schroedl, G. F. 1995. A summary of archaeological studies conducted at the

Chattooga site, Oconee County, South Carolina, 1989-1994. Unpublished

report submitted to the United States Forest Service, Columbia, South

Carolina.

Swanton, J. R. 1911. Indian tribes of the Lower Mississippi Valley and ad-

jacent coast of the Gulf of Mexico. Bureau of American Ethnology Bulletin

43, Washington, D.C.

Swanton, J. R. 1946. The Indians of the southeastern United States. Bureau

of American Ethnology Bulletin 137, Washington, D.C.

Received 20 October 1995

Accepted 1 February 1996

Mensural Discrimination of Sorex longirostris and Sorex

cinereus (Insectivora: Soricidae) in the Southeastern

United States

Joshua Laerm, Michael A. Menzel, and James L. Boone

Museum of Natural History, Institute of Ecology, and

Daniel B. Warnell School of Forest Resources,

University of Georgia, Athens, Georgia 30602

ABSTRACT — The effectiveness of univariate and multivariate statistics

in distinguishing Sorex cinereus and S. longirostris from the southeastern

United States on the basis of standard body and cranial measure-

ments was assessed. Eleven of 15 characters in univariate comparisons

showed significant differences between species, but the range of measure-

ments overlapped. Bivariate comparisons permit identification us-

ing external measurements, cranial and external measurements combined,

and cranial measurements alone. Multivariate procedures permitted

maximum distinction of the species. A discriminant function model

is presented to permit identification on the basis of three cranial

characters.

The masked shrew (Sorex cinereus Kerr 1792) is distributed throughout

the transcontinental coniferous forests of North America from the Canadian

Arctic south into the extreme northern portions of the United States

with extension into the montane forests of the Rocky and Appalachian

mountains (Hall 1981, Junge and Hoffmann 1981, van Zyll de Jong

and Kirkland 1989, Laerm et al. 1995). The southeastern shrew (Sorex

longirostris Bachman 1837) ranges from northern Missouri east through

the southern portions of Illinois, Indiana, and Ohio to Maryland, and

southward from eastern Oklahoma to Florida (French 1980a, 19806;

Hall 1981; Junge and Hoffmann 1981; and Jones et al. 1991). The

two species have overlapping distributions in northcentral Missouri

(Mock and Kivett 1980, Schwartz and Schwartz 1981, Greer 1989),

southern Illinois (Hoffmeister 1989) and Indiana (Mumford and Whitaker

1982), and throughout much of the southern Appalachians from West

Virginia and Virginia south to Georgia and South Carolina (Hall 1981,

Pagels and Handley 1989, Jones et al. 1991, Ford et al. 1994, Laerm

et al. 1995).

Sorex cinereus and S. longirostris are morphologically remarkably

similar. The two are reported to differ in that cinereus is somewhat

larger, has a longer tail (usually more than 31 mm), a comparatively

longer and more slender rostrum, a higher braincase, and third unicuspids

Brimleyana 24:15-27 April 1997 15

16 Joshua Laerm, M. A. Menzel, and J. L. Boone

larger than the fourth (French 1980a, Junge and Hoffman 1981, Jones

et al. 1991). The latter character is frequently considered to be diagnostic

(e.g., Hall 1981). However, numerous authors (Miller 1895; Jackson

1928; Kellogg 1939; French 1980a, 1980/?, 1980c; Junge and Hoffmann

1981) point out that this is not always the case. French (1980a, 19806)

reported that 20% of S. longirostris examined in Alabama and Georgia

and 12% of those in Indiana were characterized by third and fourth

upper unicuspids that were equal or nearly equal in size. Similarly,

some populations of S. cinereus exhibit third unicuspids that are smaller

than the fourth. For example, Bole and Moulthrop (1942) described

S. c. ohioensis, in part, on the basis of the third unicuspid being

smaller than the fourth. Elsewhere, Kellogg (1939:251) suggested the

synonymy of S. fontinalis (now regarded as a subspecies of S. cinereus;

see van Zyll de Jong and Kirkland 1989) with S. longirostris concluding

that "...the supposed distinctions between S. longirostris and S. fontinalis

are nothing more than individual variation."

Qualitatively, S. longirostris and S. cinereus are not difficult

to distinguish; as Jones et al. (1991:265) point out, "...under visual

examination... skulls of the two species differed markedly, S. longirostris

has a strongly arched palate and shorter rostrum, and the first two

unicuspids are of larger diameter than the third and fourth. S. cinereus

has a flat long palate and unicuspids of relatively uniform diameter."

Unfortunately, qualitative comparisons are often frustratingly difficult

to apply in the absence of a good comparative series. Jones et al.

(1991) noted that S. cinereus and S. longirostris were so similar morpho-

logically that they were not able to use S. cinereus as an out-group

in their study of geographic variation of S. longirostris.

French (1980c) made quantitative comparisons between the two

species using a univariate statistical analysis of cranial measurements

of 162 S. cinereus and 110 5. longirostris from Virgo County, Indiana.

He concluded that S. cinereus and S. longirostris were morphologically

similar and that no single character was 100% diagnostic in distinguishing

them. Although 13 standard body and cranial measurements differed

significantly between S. cinereus and S. longirostris, none was characterized

by non-overlapping ranges. Univariate morphological comparisons in

Greer's (1989) study of seven cranial measurements indicated significant

differences between the two species for six out of seven characters

in Missouri; however, as in the French (1980c) study, there was considerable

overlap.

We are not familiar with a published study of a multivariate

morphometric comparison of the two species. The purpose of this paper

is to examine the effectiveness of both univariate and multivariate

Sorex Mensural Discrimination 17

statistical procedures in distinguishing S. cinereus and S. longirostris

from the southeastern United States, where the two species show a

broad area of sympatry, on the basis of standard body and cranial

measurements.

MATERIALS AND METHODS

We used univariate and multivariate statistics to examine 200

museum specimens for morphological variation. To provide for robustness

in our analysis and include any differences due to clinal variation,

we selected 50 specimens of each species from the southern portion

of its range in Georgia, North Carolina, and South Carolina and another

50 specimens from central and northern Virginia. A priori identifications

were based on specimen tag information. In addition, we used six

additional specimens of each species not used in the model building

process to test the model. These were measured to the nearest 0.01

mm with dial calipers under a disecting microscope. Specimens examined

are listed in the Appendix.

Menzel measured the cranial characters to the nearest 0.01 mm

with a Wild M400 Stereo microscope. Images were received by an

Optronics VA-470 video camera and transferred to a 486 PC utilizing

Analytical Imaging Concepts (Irvine, California) imaging software and

stored in the TIF format. To assess the repeatability of the video measure-

ment system, a set of 10 specimens were measured three times each.

The set of 10 specimens was measured, then the order was randomized,

and the set was measured again, and finally the order was again randomized

and remeasured. Although video images could be stored for re-examination,

each specimen was rescanned and the system was recalibrated prior

to each remeasurement.

Eleven cranial characters (Table 1) were measured on all individuals:

condylobasilar length (CBL), cranial breadth (CB), length of unicuspid

toothrow (LUT), length of 1st unicuspid (LU1), breadth of 1st unicuspid

(BUI), length of 3rd unicuspid (LU3), breadth of 3rd unicuspid (BU3),

length of 4th unicuspid (LU4), breadth of 4th unicuspid (BU4), length

of unicuspids 3 and 4 (LU34), and breadth across 2nd molars (BM2).

External body measurements (total length, tail length, and hind foot

length) and sex were recorded when available from specimen tags;

body length was calculated by subtracting tail length from total length.

Each specimen was assigned to one of 12 age classes based on the

criteria of Rudd (1955).

Statistical analyses were performed with Systat 5.1a (Wilkenson

1989) and SPSS 4.01 (Norusis 1990). Univariate normality and homogeneity

of variance were tested by inspection of plotted residuals and Bartlett's

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20 Joshua Laerm, M. A. Menzel, and J. L. Boone

test for homogeneity of group variances, respectively. Inspection of

residuals revealed that 12 of 2,400 measurements (200 specimens by

12 measurements) were found to be extreme (>5 standard deviations

from the mean). Five of these extreme measurements were attributed

to two individuals, and both individuals (USNM 75167, USNM 296566)

were deleted from the analysis. The other extreme measurements were

attributed to six different individuals. These six measurements and

11 other missing observations were replaced with the within-group

mean of the character in question so that these individuals could be

included in the multivariate analyses. After the extreme observations

were corrected, we assumed multivariate normality based on marginal

normality and multivariate homogeneity of variance based on failure

of rejection in the test of equality of group covariance matrices using

Box's M (P = 0.082).

Differences among repeated measures, adult age classes, and sexes

were tested with analysis of variance, and type-1 error rates were corrected

with the sequential Bonferrioni adjustment (Rice 1989) where necessary.

Taxa were classified using stepwise discriminant analysis. Variables

were included in the models based on minimizing Wilk's lambda, prior

probabilities were equal to sample size, and varimax rotation was employed.

Stepwise discriminant analysis will find an optimal solution based on

the data; however, depending on which variables enter the model first,

it may find a local optimum rather than the global optimum. To help

avoid this optimization problem, we removed variables that entered

the model in the first steps and repeated the analysis. All analyses

were performed on raw data without transformation and without removing

size (Rohlf and Bookstein 1987), because this produced the simplest

tool for future classification of new specimens consistent with a goal

of a high degree of group separation.

The model separating S. cinereus and S. longirostris was validated

in two ways. First it was validated internally by randomly selecting

subsets of the data (approximately 80% of the data selected without

regard to species), constructing the disciminate model, and using that

model to classify the remaining 20% of the specimens. This procedure

was repeated 200 times. Second, because the skulls were originally

measured utilizing a non-traditional approach, the model was validated

externally with additional specimens (six test specimens of each species)

measured with dial calipers under a disecting microscope.

RESULTS AND DISCUSSION

In the analysis of the repeated measures, no significant difference

was found among measurements for any of the 11 cranial characters.

Sorex Mensural Discrimination 21

Very little of the total variance could be attributed to the repeated

measures (range of 0 to 24%, x = 5.5%), suggesting that with careful

calibration the video system provided highly repeatable measurements.

In univariate comparisons of the sexes, only the length of unicuspids

3 and 4 (LU34) in S. longirostris differed significantly (P = 0.031).

Males and females averaged 0.71 mm (SE = 0.0064, n = 49) and

0.69 mm (SE = 0.0091, n = 20), respectively (Rice 1989). When this

character was examined within regions, sexes did not differ significantly

(southern sample, P = 0.23; northern sample, P = 0.10).

In a few cases, differences among age classes within species,

regions, and sexes were individually, but not collectively, significant

(a < 0.05). The only consistently significant (P < 0.01) difference

among age groups was length of first unicuspid (LU1) which tended

to decrease in magnitude with increasing age.

In univariate analysis of morphological variation, all characters

except body length, breadth of third unicuspid (BU3), and breadth

of fourth unicuspid (BU4) differed significantly (P < 0.001) between

species. For all characters that showed significant differences, except

breadth across second molars (BM2), S. cinereus was larger than S.

longirostris. For character BM2, the size of S. longirostris exceeded

S. cinereus. In all cases except tail length, however, the range of

measurements for both species overlapped (Table 1).

Multivariate analysis using cranial measurements was successful

in correctly identifying all specimens of the two species. However,

the geographic origin of only 79% of the northern and southern specimens

could identified correctly. For S. longirostris, 11 southern and 11 northern

specimens were incorrectly classified into the opposite geographic group;

for S. cinereus, 7 southern and 12 northern specimens were incorrectly

classified. Such a measure of geographic variation, perhaps clinal,

was expected.

Discriminant analysis using cranial measurements for the two

species correctly classified all specimens with three characters, LU34,

BM2, and CBL, in order of inclusion into the model. Standardized

canonical discriminant function coefficients were 0.60, 0.72, and

-0.73 for LU34, BM2, and CBL, respectively. Pooled within-groups

correlations between discriminating variables and canonical discriminant

functions, variables ordered by size of correlation within function, were

0.69, -0.25, and 0.53 for these characters, respectively.

Unknown specimens can be identified to species with this latter

model using unstandardized canonical discriminant function coefficients

for the three variables. To do so, measure the unknown specimen for

CBL, LU34, and BM2, then multiply each measurement by its coefficient

22

Joshua Laerm, M. A. Menzel, and J. L. Boone

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Sorex Mensural Discrimination 23

(1.58, 16.90, and -5.49, respectively), sum the three products, and

add a constant (-17.25). The resulting value is the specimen's discriminant

score. If the score is greater than zero, the specimens in assigned

to S. cinereus, otherwise it is assigned to S. longirostris. The average

discriminant score for S. cinereus is 3.06, and the average for S. longirostris

is -3.18.

Discriminant analysis using cranial and external measurements

for the two species correctly classified all specimens with two characters,

LU34 and tail length, and this bivariate comparison can be used to

identify new specimens without transformation (Fig. la). Similarly,

using only external characters, all specimens can be identified to species

with a bivariate comparison of body and tail lengths (Fig. lb). Using

only cranial characters, specimens can be correctly classified with a

high degree of probability (99.5%) with a bivariate comparison of LU34

and BM2 (Fig. lc).

VALIDATION

Validation of the model separating the species showed that the

results were stable, as 193 of 200 trial runs produced 100% correct

classification. The seven trails producing errors had one misclassification

each; therefore, of 7,579 individuals classified in the validation process,

only seven were classified incorrectly. All errors were the misclassification

of S. cinereus specimens.

We validated the utility of this model based upon six test specimens

of each species from localities not used in developing the model and

with a more conventionally available measuring device (i.e., dial calipers

and dissecting microscope). The discriminant analysis was sufficiently

robust that all specimens were correctly identified.

CONCLUSION

Sorex cinereus and S. longirostris can be distinguished on the

basis of any one of three bivariate plots using untransformed data

(Fig. 1) or by discriminant analysis. The results of our univariate compari-

sons are similar to those of French (1980c) and Greer (1989); we

observed a high degree of overlap in all but one mensural character

(tail length). Possibly, the separation of the two species by cranial

characters in our study is a reflection of the finer scale of measurement

permitted by computer assisted video imaging. We should note that

regional differences in the morphology of both S. cinereus and S. longirostris

might limit the effectiveness of the methods and characters used by

us in mensural discrimination of these two species in areas other than

the Southeast.

24 Joshua Laerm, M. A. Menzel, and J. L. Boone

ACKNOWLEDGMENTS'^^ thank the staff of the Electron Microscopy

Laboratory at the University of Georgia and, particularly, M. Farmer

for his patience in providing training in the use of the equipment.

M. E. McGhee, R. Fisher, and J. Pagels provided specimens under

their respective care. M. Kennedy, and M. Harvey examined several

specimens. A critical review by G. L. Kirkland, Jr. much improved

the manuscript. Support for this project was provided by the University

of Georgia Museum of Natural History and National Science Foundation

grant BSR 9011661 at Coweeta Hydrological Laboratory, North Carolina.

LITERATURE CITED

Bole, B. P., Jr., and P. N. Moulthrop. 1942. The Ohio Recent mammal

collection in the Cleveland Museum of Natural History 5:83-81.

Ford, W. M., J. Laerm, and D. C. Weinand. 1994. Abundance and distri-

bution of shrews and other small mammals in the Chattahoochee Na-

tional Forest of Georgia. Proceedings of the Annual Conference Southeastern

Association Fish and Wildlife Agencies 48:310-320.

French, T. W. 1980a. Sorex longirostris. Mammalian Species 143:1-3.

French, T. W. 1980b. Natural history of the southeastern shrew, Sorex

longirostris Bachman. The American Midland Naturalist 104:13-31.

French, T. W. 1980c. Ecological relationships between the southeastern

shrew {Sorex longirostris Bachman) and the masked shrew {Sorex cinereus

Kerr) in Virgo County, Indiana. Ph.D. Dissertation. Indiana State University,

Terre Haute.

Greer, K. S. 1989. Ecological and morphological comparison of two syntopic

species of soricids, Sorex cinereus and Sorex longirostris, in north central

Missouri. M. S. Thesis. Northeast Missouri State University, Kirksville.

Hall, E. R. 1981. The mammals of North America. Second edition. John

Wiley & Sons, New York, New York.

Hoffmeister, D. F. 1989. Mammals of Illinois. University of Illinois Press,

Urbana.

Jackson, H. H. T. 1928. A taxonomic review of the Americana long-tailed

shrews. North American Fauna 51:1-238.

Jones, C. A., S. R. Humphrey, T. M. Padgett, R. K. Rose, and J. H. Pagels.

1991. Geographic variation and taxonomy of the southeastern shrew

{Sorex longirostris). Journal of Mammalogy 72:263-272.

Junge, J. A., and R. S. Hoffmann. 1981. An annotated key to the long-

tailed shrews (genus Sorex) of the United States and Canada, with notes

on middle American Sorex. Occasional Papers of the Museum of Natural

History, University of Kansas 94:1-48.

Kellogg, R. 1939. Annotated list of -the mammals of Tennessee. Proceed-

ing of the United States National Museum 86:245-303.

Laerm, J., M. A. Menzel, E. Brown, A. Wotjalik, W. M. Ford, and M. Strayer.

1995. The masked shrew, Sorex cinereus Kerr (Insectivora: Soricidae),

Sorex Mensural Discrimination 25

and red-backed vole, Clethrionomys gapperi (Rodentia: Muridae) in the

Blue Ridge Province of South Carolina. Brimleyana 22:15-21.

Miller, G. S., Jr. 1895. The long-tailed shrews of the eastern United States.

North American Fauna 10:35-56.

Mock, O. B., and V. K. Kivett. 1980. The southeastern shrew, Sorex longirostris,

in northeastern Missouri. Transactions of the Missouri Academy of Science

14:67.

Mumford, R. E., and J. O. Whitaker, Jr. 1982. Mammals of Indiana. In-

diana University Press, Bloomington.

Norusis, N. J. 1990. SPSS base system user's guide. SPSS Inc., Chicago,

Illinois.

Pagels, J. F., and C. O. Handley, Jr. 1989. Distribution of the southeast-

ern shrew, Sorex longitrostris Bachman, in western Virginia. Brimleyana

15:123-131.

Rice, W. R. 1989. Analyzing tables of statistical tests. Evolution 43:223-

225.

Rohlf, F. J., and F. L. Bookstein. 1987. A comment on shearing as a method

for "size correction." Systematic Zoology 36:356-367.

Rudd, R. L. 1955. Age, sex, and weight comparisons in three species of

shrews. Journal of Mammalogy 36:323-339.

Schwartz, C. W., and E. R. Schwartz. 1981. The wild mammals of Mis-

souri. University of Missouri Press, Columbia.

Wilkenson, L. 1989. SYSTAT: The system for statistics. SYSTAT, Inc.,

Evanston, Illinois.

van Zyll de Jong, C. G., and G. L. Kirkland, Jr. 1989. A morphometric

analysis of the Sorex cinereus group in central and eastern North America.

Journal of Mammalogy 70:10-122.

Received 24 October 1995

Accepted 1 February 1996

26 Joshua Laerm, M. A. Menzel, and J. L. Boone

APPENDIX— SPECIMENS EXAMINED

For each species and each state, entries include county,

location, number of specimens from that location, and when

necessary, acronym of the museum housing the specimen (UGAMNH=

University of Georgia Museum of Natural History, USNM=National

Museum of Natural History, VCU=Virginia Commonwealth University).

Sorex longirostris

GEORGIA (all UGAMNH): Clarke Co.: Athens, Baldwin Avenue,

University of Georgia campus, 1. Fulton Co.: Long Island Creek

at Chattahoochee River, 10. Lumpkin Co.: Dockery Lake, 3.75

miles N Stone Pile Gap on GA 19, 2. Rabun Co.: Ann Gap Road

(FS 410), 2 miles W Low Gap Road, 2. Stephens Co.: Lake

Russell Wildlife Management Area, 2; Lake Russell Wildlife

Management Area, N of junction of FS Roads 62 and 62A,

1; Lake Russell Wildlife Management Area, Dike 5 Creek at

FS 87, 1; Davidson Creek, 220 m upstream from Panther Creek,

3. Union Co.: 1.9 miles WSW Suches, 1; 2.3 miles WSW Suches,

1; 2.0 miles W Suches, 3; GA 180, 0.25 miles North of Lake

Winfield Scott, 1.

SOUTH CAROLINA (all UGAMNH): Aiken Co.: Savannah

River Plant, Bullfrog Pond, 5; Savannah River Plant, F-Bay,

1; Savannah River Plant, Flamingo Bay, 1; Savannah River Plant,

Linda Pond, 1; Savannah River Plant, Pickerel Pond, 1; Savannah

River Plant, Rainbow Bay, 5; Savannah River Plant, Sun Bay,

1. Oconee Co.: Sumpter National Forest Road 709, 1.1 miles

west of Highway 107, 5. Picken Co.: van Clayton Memorial Highway,

0.9 M below summit of Sassafrass Mountain, 1.

VIRGINIA: Amelia Co.: Amelia Court House, 2 (USNM);

Burke, near Seward Forest, 1 (USNM); Falls Church, 1 (USNM);

Shenandoah National Park Headquarter, 3 (USNM); Triplett,

Seward Forest, 2 (USNM). Chesapeake Co.: Dismal Swamp,

Lake Drummond, 2 (USNM). Chesterfield Co.: 4 miles N Keswick

Farm, 1 (USNM). Culpepper Co.: 10 miles SE Legnum, 1 (USNM).

Cumberland Co.: Columbia (Goochland), 30 (VCU). Essex Co.:

3.5 miles SW Center Cross, 2 (USNM). Fairfax Co.: Fort Belvior,

Site 104, 1 (USNM); Fort Belvior, Site CA-5, 1 (USNM). Norfolk

Co.: Wallacetown, 4.7 miles NNE, near US 17, 1 (USNM).

Sorex cinereus

GEORGIA (All UGAMNH): Rabun Co.: Burnt Cabin Branch,

2 miles N Tate City at North Carolina State line, 4; Rabun

Bald, 1; Base of Rabun Bald at Beechgum Gap, 0.2 mile up

jeep trail from Gap, 2; FS 150, 4.0 miles S. Dillard at Thomas

Sorex Mensural Discrimination 27

Creek, 5; FS 150, 3.1 miles E Dillard at Thomas Creek, 3; FS

150, 2.5 miles E Dillard, 1. Towns Co.: Beech Creek at Tulula

River, 1; Swallow Creek Management Area, Fork Ridge, 1; FS

79, E of Mossy Creek Branch, N of Tray Mountain Gap, 9;

Swallows Creek Management Area, intersection of FS 698 and

FS 698A, 4. White Co.: FS 79, 0.4 miles South Tray Mountain

Gap, 4.

NORTH CAROLINA (all UGAMNH): Haywood Co.: Shining

Rock, 1. Macon Co.: Coweeta Hydrological Laboratory, 4; Coweeta

Hydrological Laboratory, Dryman's Fork, 1; Coweeta Hydrological

Laboratory, Lick Branch, 1.

SOUTH CAROLINA (all UGAMNH): Oconee Co.: USFWS Fish

Hatchery Visitor Center, 3; 1.0 mile up access road to Fish

Hatchery, 5.

VIRGINIA: Giles Co.: Mountain Lake, 1 (USNM); Mountain

Lake, 1.7 miles ENE Castle Rock, 2 (USNM); Mountain Lake,

1.8 miles NE Cross Trail, 1 (USNM); Mountain Lake, 2.5 miles

NW Ashby Flats, 1 (USNM); Mountain Lake, 2.6 miles NW

Ashby Bogs, 1 (USNM); Mountain Lake, 2.7 miles NE Warspur

Branch, 1 (USNM); Mountain Lake, 2.7 miles NW Ashby Flat,

3 (USNM); Mountain Lake, 2.7 miles NW Ashby Meadow, 3

(USNM); Mountain Lake, 4.3 miles NNE Castle Rock, 3 (USNM);

Mountain Lake, 4.5 miles NE Big Mountain, 1 (USNM); Mountain

Lake, 4.5 miles NE Big Soft Seep, 1 (USNM); Mountain Lake,

4.5 miles WNW area 4, 1 (USNM); Mountain Lake, 5 miles

NE Bob's Field, 1 (USNM); Mountain Lake, Ashby Bogs, 2

(USNM); Mountain Lake, Butt Mountain, Upper Field, 3 (USNM).

Highland Co.: Laurel Fork Area, 21 (VCU); Red Oak Knob, 5

(VCU).

Record of a Creek Chub, Semotilus atromaculatus

(Cypriniformes: Cyprinidae), Preying on a Jumping Mouse

(Zapodidae) in Bruffey Creek, Pocahontas County,

West Virginia

William J. Poly

Department of Zoology

Southern Illinois University

Carbondale, Illinois 62901

AND

Charles E. Boucher

Ohio Environmental Protection Agency

1685 Westbelt Drive

Columbus, Ohio 43228

ABSTRACT — In Bruffey Creek, West Virginia, we collected a creek

chub, Semotilus atromaculatus, that had consumed a jumping mouse

(Zapodidae). Small mammals have not been reported in the diet of

creek chubs.

On 30 September 1995, while capturing fishes in Bruffey Creek

as part of a study involving Bruffey-Hills Creek Cave fishes, we collected

a creek chub, Semotilus atromaculatus (Mitchill), excreting the remains

of a rodent. We preserved the creek chub and remains for later identification.

The creek chub measured 156.6 mm SL, 189.0 mm TL, 16.5 mm

gape width, 46.2 mm head length, and 37.8 mm body depth. The

skeleton of the prey was intact except for the skull. Fur and internal

organs were still present as well. Examination of the creek chub's

gut revealed no sign of the skull. Since the mouse was being excreted

tail first, we believe the head was severed and expelled during ingestion.

The feet were completely intact, and the tail appeared to be undigested

as well, but also apparently had been broken. Based on feet (24.9

and 25.1 mm) and tail (88.2 mm) lengths the mouse was a jumping

mouse (Zapodidae), either meadow jumping mouse, Zapus hudsonius

(Zimmerman) or woodland jumping mouse, Napaeozapus insignis (Miller).

A specific determination can not be made without the skull.

S. atromaculatus food habits have been studied by a number of

investigators, and food items include algae, plant material, terrestrial

and aquatic insects, Mollusca, Crustacea, fishes, and frogs (Forbes

1888; Hankinson 1910; Forbes and Richardson 1920; Leonard 1927;

Brimleyana 24:29-32, April 1997 29

30 William J. Poly and Charles E. Boucher

Greeley 1930; Sibley and Rimsky-Korsakoff 1931; Hubbs and Cooper

1938; Simpson 1941; Dobie et al. 1948; Starrett 1948; Dinsmore 1962;

Minckley 1963; Barber and Minckley 1971; Moshenko and Gee 1973;

Newsome and Gee 1978; Copes 1978; Johnson and Johnson 1982;

Magnan and FitzGerald 1982, 1984; Angermeier 1982, 1985; Keast

1985; Garman and Moring 1993); however, no mammalian remains

were reported in these studies. Dobie et al. (1948:91) remarked: "The

northern creek chub seems to eat anything that comes its way." We

found one record of cyprinids feeding on mammals: flathead chub

(Platygobio gracilis (Richardson)) eating a small rodent (McPhail and

Lindsey 1970). Larger predatory fishes such as northern pike (Esox

lucius Linnaeus) and muskellunge (E. masquinongy Mitchill) have been

known to consume small mammals or birds (Anderson 1948, Lawler

1965). In the early 1990s, one boreal red-backed vole (Clethrionomys

gapperi (Vigors)) was found in a smallmouth bass (Micropterus dolomieu

Lacepede) from Lake Saganaga, Minnesota/Ontario (David A. Etnier,

University of Tennessee, personal communication). A green sunfish

(Lepomis cyanellus Rafinesque), preyed on a Mexican free-tailed bat

(Tadarida mexicana (Saussure)) in a Texas cave (Jones and Hettler

1959), and goldeye (Hiodon alosoides (Rafinesque)) are known to consume

small mammals as well (Dymond and Hart 1927, Scott and Crossman

1973). Quimby (1951) reported E. lucius preying on a Z. hudsonius

in Minnesota. Z. hudsonius has been reported to swim and dive underwater

to avoid capture (Quimby 1951, Hoffmeister 1989, references in Krutzsch

1954) and is often found near water. Such an "affinity" for water

would explain why this mouse is occasionally preyed upon by fishes.

The creek chub and mouse remains have been catalogued in the Southern

Illinois University at Carbondale Fish Collection (SIUC 24849).

ACKNOWLEDGMENTS— Brady Porter provided an initial identification

of the mouse remains, and George Feldhamer confirmed that identification

by comparison with preserved specimens. Brooks Burr provided several

relevant publications, and David Etnier supplied the incident of red-

backed vole predation by a smallmouth bass.

LITERATURE CITED

Anderson, L. R. 1948. Unusual items in the diet of the northern muskel-

lunge (Esox masquinongy immaculatus). Copeia 1948:63.

Angermeier, P. L. 1982. Resource seasonality and fish diets in an Illi-

nois stream. Environmental Biology of Fishes 7:251-264.

Angermeier, P. L. 1985. Spatio-temporal patterns of foraging success for

Creek Chub 31

fishes in an Illinois stream. American Midland Naturalist 114:342-359.

Barber, W. E., and W. L. Minckley. 1971. Summer foods of the cyprinid

fish Semotilus atromaculatus. Transactions of the American Fisheries

Society 100:283-289.

Copes, F. C. 1978. Ecology of the creek chub. Reports on the fauna and

flora of Wisconsin, University of Wisconsin Museum of Natural His-

tory, Stevens Point, Number 12:1-21.

Dinsmore, J. J. 1962. Life history of the creek chub, with emphasis on

growth. Proceedings of the Iowa Academy of Science 69:296-301.

Dobie, J. R., O. L. Meehean, and G. N. Washburn. 1948. Propagation of

minnows and other bait species. United States Fish and Wildlife Service

Circular 12:1-113.

Dymond, J. R., and J. L. Hart. 1927. The fishes of Lake Abitibi (Ontario)

and adjacent waters. University of Toronto Studies, Biological Series,

Publications of the Ontario Fisheries Research Laboratory 28:3-19.

Forbes, S. A. 1888. On the food relations of fresh-water fishes: A sum-

mary and discussion. Bulletin of the Illinois State Laboratory of Natural

History 2:475-538.

Forbes, S. A., and R. E. Richardson. 1920. The fishes of Illinois. Second

edition. Illinois Natural History Survey, Champaign.

Garman, G. C, and J. R. Moring. 1993. Diet and annual production of

two boreal river fishes following clearcut logging. Environmental Bi-

ology of Fishes 36:301-311.

Greeley, J. R. 1930. A contribution to the biology of the horned dace,

Semotilus atromaculatus (Mitchill). Ph.D. Thesis. Cornell University,

Ithaca, New York.

Hankinson, T. L. 1910. An ecological study of the fish of a small stream.

Transactions of the Illinois State Academy of Science 3:23-31.

Hoffmeister, D. F. 1989. Mammals of Illinois. University of Illinois Press,

Urbana.

Hubbs, C. L., and G. P. Cooper. 1938. Minnows of Michigan. Cranbrook

Institute of Science Bulletin Number 8 (second printing, revised). Bloomfield

Hills, Michigan.

Johnson, J. H., and E. Z. Johnson. 1982. Diel foraging in relation to available

prey in an Adirondack mountain stream fish community. Hydrobiologia

96:97-104.

Jones, R. S., and W. F. Hettler. 1959. Bat feeding by green sunfish. Texas

Journal of Science 11:48.

Keast, A. 1985. The piscivore feeding guild of fishes in small freshwater

ecosystems. Environmental Biology of Fishes 12:119-129.

Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). Uni-

versity of Kansas Publications of the Museum of Natural History 7:349-

472.

Lawler, G. H. 1965. The food of the pike, Esox lucius, in Heming Lake,

Manitoba. Journal of the Fisheries Research Board of Canada 22:1357-

1377.

32 William J. Poly and Charles E. Boucher

Leonard, A. K. 1927. The rate of growth and the food of the horned dace

(Semotilus atromaculatus) in Quebec, with some data on the food of

the common shiner (Notropis cornutus) and of the brook trout (Salvelinus

fontinalis) from the same region. University of Toronto Studies, Bio-

logical Series, Publications of the Ontario Fisheries Research Labora-

tory 30:35-44.

Magnan, P., and G. J. FitzGerald. 1982. Resource partitioning between

brook trout (Salvelinus fontinalis Mitchill) and creek chub (Semotilus

atromaculatus Mitchill) in selected oligotrophic lakes of southern Quebec.

Canadian Journal of Zoology 60:1612-1617.

Magnan, P., and G. J. FitzGerald. 1984. Ontogenetic changes in diel ac-

tivity, food habits and spatial distribution of juvenile and adult creek

chub, Semotilus atromaculatus. Environmental Biology of Fishes 11:301—

307.

McPhail, J. D., and C. C. Lindsey. 1970. Freshwater fishes of northwest-

ern Canada and Alaska. Fisheries Research Board of Canada Bulletin

173.

Minckley, W. L. 1963. The ecology of a spring stream Doe Run, Meade

County, Kentucky. Wildlife Monographs 11:1-124.

Moshenko, R. W., and J. H. Gee. 1973. Diet, time and place of spawn-

ing, and environments occupied by creek chub (Semotilus atromaculatus)

in the Mink River, Manitoba. Journal of the Fisheries Research Board

of Canada 30:357-362.

Newsome, G. E. (Buck), and J. H. Gee. 1978. Preference and selection

of prey by creek chub (Semotilus atromaculatus) inhabiting the Mink

River, Manitoba. Canadian Journal of Zoology 56:2486-2497.

Quimby, D. C. 1951. The life history and ecology of the jumping mouse,

Zapus hudsonius. Ecological Monographs 21:61-95.

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada.

Fisheries Research Board of Canada Bulletin 184.

Sibley, C. K., and V. N. Rimsky-Korsakoff. 1931. Food of certain fishes

in the watershed. A biological survey of the St. Lawrence watershed.

Supplement to Twentieth Annual Report, New York State Conserva-

tion Department 1930 (1931):109-120.

Simpson, J. C. 1941. Food analysis of some important species of Wyo-

ming forage fishes. M.S. Thesis. University of Wyoming, Laramie.

Starrett, W. C. 1948. An ecological study of the minnows of the Des Moines

River, Boone County, Iowa. Ph.D. Thesis. Iowa State College, Ames.

Received 13 February 1996

Accepted 6 March 1996

Note added after typesetting: N. insignis and 5 other small mammals have

been reported in the diet of largemouth bass (Micropterus salmoides (Lacepede))

by J. R. Hodgson and M. J. Kinsella (1995. Small mannals in the diet of

largemouth bass, revisited. Journal of Freshwater Ecology 10:433-435).

Clam Siphon Tip Nipping by Fishes in the Estuarine Cape

Fear River, North Carolina

Frank J. Schwartz

Institute of Marine Sciences

University of North Carolina

Morehead City, North Carolina 28557

ABSTRACT— Over two million fishes within 57 families and 173

species were collected between 1973 and 1978 in the Cape Fear

River, North Carolina. Sampling consisted of repetitive six-year, 22

station gill net (2,362 sets) and otter trawl (8,284 tows) efforts.

Stomachs of 82 species contained fishes. Diets of 14 species rep-

resenting nine fish families were found to include clam siphon tips,

primarily Mercenaria mercenaria. The 14 species comprised 39.7%

of the total catch (798,607), and examining 21,732 stomachs found

siphon nipping had occurred 453 times by 889 individuals (4.1%).

Nipping was most intense in 1976 and 1977, years when river wa-

ter temperatures were historically lowest, and shoal areas were subjected

to large expanses of ice flows. Sampling daily, weekly, and monthly

revealed that clam populations were patchy. Most "nipping" fishes

were less than 126 mm in standard length (x = 90 mm SL). Most

siphon tip feeding fishes were caught in September, August, and

October, and least in December. Nipping behavior was dominated

by croakers, hogchokers, southern kingfish, spot, pinfish, and fringed

flounders.

Siphon tips of various molluscs have been noted in stomach contents

of bothid, coryphaenid, elasmobranch, gerreid, pholid, sciaenid, and

tetraodontid fishes (Joseph et al. 1982; McMichael and Ross 1983;

Modde and Ross 1983; Cyrus and Blaber 1983, 1984; Hughes 1985;

McMichael 1986; Cyrus 1988; Compagno 1990; Coen and Heck 1991).

Other animals (reviewed in Kamermans and Huiteman 1954) such as

crabs (Hines et al. 1990), shrimps (Kamermans and Huiteman 1994),

sea otters (Kvitek et al. 1991), walrus (Welsh and Martin-Bergmann

1990), and isopods and decapods (Bonsdorff et al. 1995) are also known

siphon tip nippers.

Most siphon nipping observations have been reported following

food content analyses of a variety of organisms. The importance and

impact of siphon nipping was discussed by Armitage and Alevizon

(1980) and Kamermans and Huiteman (1994), who commented on the

poor caloric value of siphon tips. Few efforts have attempted to describe

the frequency of siphon tip nipping, or its effects on mollusc growth

Brimleyana 24:33-45, April 1997 33

34 Frank J. Schwartz

with time (Coen and Heck 1991, Sutherland 1982, Peterson and Quammen

1982).

I document clam siphon tip nipping by 14 species of fishes captured

at 22 stations in the Cape Fear River system of North Carolina during

intensive gill net and otter trawl samplings (10,646 efforts) between

1973 and 1978 and discuss the impact by size of fish, station, month,

year, and species.

STUDY AREA AND METHODS

The Cape Fear River south of Wilmington, North Carolina, is

an estuarine system that lies entirely within, and is the largest river

drainage to the Atlantic Ocean, in North Carolina (Schwartz et al.

1982). A study area was a 7,854-ha portion of the river south of Wilmington

that varied 1.6-3.6 km wide, is 17 km long, and daily is subject to

±2 m tides that are affected by prevailing southeast or southwest winds

during nine months of the year. It included the main river from Buoy

42, just south of Wilmington, and near Campbell Island, southward

for 17 km to the ocean, and nearby Carolina Beach Inlet, Masonboro

Sound (Fig. 1). (See Schwartz et al. 1979a, b; 1982) for further habitat

and ecological details and descriptions.)

Yearly 22 stations (Fig. 1) were sampled, during daylight hours

of 1973 through 1978, 10,646 times (2,362 gill nets sets; 8,284 otter

trawl tows). Sampling occurred twice each January, weekly sampling

occurred February through May and September through November,

and half of December. Single monthly samplings occurred in June,

July, and August of each year. Each shoal and intake canal station

was sampled for pelagic species using 8.7-cm x 91.4-m gill nets set

12 hours. Semi-balloon 7.6-m (all shoal and intake canal stations)

and 12.6-m (all shoal, channel, intake canal, and ocean stations) 1.9-

cm-stretched mesh otter trawls towed 0.3 hour were used to capture

all other species. A total of 2,013,986 fishes, within 57 families and

173 species, were collected. Entire small catches were kept, whereas

large catches were subsampled using a 8.5-L pail. The resultant mixed

catches and subsamples were immediately preserved in 10% formalin

and later sorted in the lab. Remaining specimens of large catches

were further subsampled for total number and mass, and returned alive

to each original capture site. Eighty-four species and 798,607 specimens

(39.7%) of the total were measured (standard length in millimeters,

SL) weighed (0.1 gm), and examined- for food content (Table 1). These

had been obtained following sampling all stations, except Buoy 42

(80 times) and the ocean (1,056 times) (range 222-744 times/station;

Table 2).

Siphon Nipping by Fishes

3 5

Fig. 1. Locations of river or shoal gill net and otter trawl stations (■),

bouys (•), sampled in the Cape Fear River and adjacent areas between 1973

and 1978.

36

Frank J. Schwartz

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40 Frank J. Schwartz

Stomachs of 84 species of fishes were examined to note percent

fullness, percent volume, and frequency of occurrence of each food

item. Foods were present in the stomachs of 82 species. Stomach content

of each food item was estimated visually by percent volume and determined

to the lowest possible taxon.

Environmental features of salinity were noted with A/O refractometers,

air and water temperatures with Taylor portable field thermometers,

and oxygen content with YSI 51 or 57 units.

RESULTS

Siphon tips were found in the stomachs of 889 individual fishes

(4.1% of those examined with food) comprising 14 species and nine

fish families (Bothidae, 4 species; Sciaenidae, 3; Achiridae, 1; Soleidae,

1; Ephippidae, 1; Haemulidae, 1; Serranidae, 1; Sparidae, 1; and Triglidae,

1) collected 453 times between 1973-1978 (Table 1). Siphons tips had

been eaten by 514 males, 313 females, and 62 specimens whose sex

was undetermined (Table 1). Recognizable food was found in 21,732

of the 27,461 specimens (14 species) examined (Table 1).

Croaker (Micropogonias undulatus) stomachs (349 specimens) often

contained up to 80% of their stomach contents as siphon tips. Hogchokers

(Trinectes maculatus) (138 specimens) were the second most frequent

siphon tip browser, followed by southern kingfish {Menticirrus americanus)

(108), spot (Leiostomus xanthurus) (105), pigfish {Orthopristes chrysopterus)

(90), and fringed flounder (Etropus crossotus) (58) (Table 1).

Regardless of species caught, most specimens containing siphon

tips were less than 126 mm, average 90 mm SL (Table 1). Largest

specimens eating siphon tips were the pigfish (Orthopristis chrysoptera)

(270 mm SL), southern flounder (Paralichthys lethostigma) (270 mm

SL), and leopard searobin (Prinotus scitulus) (157 mm SL), respectively

(Table 1). Siphon tips were found most often in yearling fishes caught

in September (111 times), August, and October (59 each), with least

occurrences in December (1) (Table 2).

Sampling effort by station/month ranged between 80-1,056 (Table

3). Fishes with siphon tips in their stomach contents were caught

more often at Station 174 (67 times), Buoy 19 (43), 23 (42), and

the ocean (46) than at most other stations (Table 3), perhaps as a

result of clam patchiness. Most siphon tip nipping occurred in 1976

(155) and 1977 samples (137) (Table 4); the least in 1973. Although

sampling efforts were greater in 1976 and 1977 (Table 4), other factors

such as cold winter waters or ice cover were perhaps more important

in inducing nipping than sampling effort. Low field recorded water

temperatures were 7 C in January 1976 and 4 C in January 1977,

Siphon Nipping by Fishes 41

Table 4. Siphon tip nipping occurrences by family, species, year, and sampling

effort, Cape Fear River, North Carolina.

Year

Fishes 1973 1974 1975 1976 1977 1978

Bothidae

Fringed flounder

Bay whiff

Summer flounder

Southern flounder

Sciaenidae

Spot

Southern kingfish

Atlantic croaker

Achiridae

Hogchoker 0 4 4 22 23 7

Soleidae

Blackcheek tonguefish 0 10 0 10

Epippidae

Atlantic spadefish 0 0 0 1 10 3

Haemulidae

Pigfish 0 112 0 2

Serranidae

Black sea bass 0 0 0 0 10

Sparidae

Pinfish 0 3 1 32 16 2

Triglidae

Leopard searobin 0 10 0 0 0

Total occurrences 1 52 34 155 137 74 453

Sampling effort (stations) 1,240 1,417 1,972 2,139 2,116 1,762 10,646

42 Frank J. Schwartz

the latter causing ice flow development on the shoals of the river

and in the power plant intake canal. As a result, fish kills were common

each of the two years at several river stations. Highest water temperatures

were 30.5-32.0 C in July 1977. Oxygen levels associated near the

ice flows or high water temperatures were always high, yet were critical

for some species such as striped mullet, Mugil cephalus, grey trout,

Cynoscion regalis, and menhaden (Brevoortia tryannus). Salinities varied

by season, station, and after rainfall and runoff, thereby enchancing

or preventing greater range utilization of the river system than usual,

i.e., channel catfish Ictalurus punctatus, a species of the upper river

was often found as far down river as Buoy 18 (Fig 1).

DISCUSSION

Although sciaenids (croaker, spot, and southern kingfish) have

been reported eating Donax or other clam siphons (Modde and Ross

1983, Currin 1984, McMichael 1986, McMichael and Ross 1988, Irlandi

1993, Currin et al. 1994), my study adds 11 species to the list of

siphon tip nipping fishes. No attempt was made herein to note the

rate of siphon tip regeneration or length of siphon extension (Zwartz

et al. 1994). The most severe cold-winter-spring waters ever recorded

(1976 and 1977) may have caused increased siphon nipping (Table

4), and surface inhabiting invertebrates to vacate the area or influenced

their survival, even death. Clams on the other hand could simply withdraw

their siphons during the most severe water temperature extremes and

remain in the area. Thus, loss of other winter foods may have made

clams the only available food for bottom feeding fishes such as croakers,

spot, southern kingfish, etc.

Infrequent literature reports of siphon tips as stomach contents

of fishes should be viewed cautiously in light of pre-, during- and

post-capture factors. Prevailing environmental events should be factored

into the observations rather than simply assuming the presence of a

food item was preferred and expected, rather than unexpected (Bonsdorff

et al. 1995). Also efforts to interpret the effects of a fish's behavior,

such as siphon tip nipping, should consider whether a station was

sampled once or repeatedly to determine the long-term effects of fish

behavior (aversive or non-aversive; Kvitek 1991) and sampling effects

on the local clam population.

Siphon Nipping by Fishes 43

ACKNOWLEDGMENTS— Support for the long-term study was provided

by Carolina Power and Light Company, Raleigh, North Carolina. At

least 52 technicians helped collect and analyze the 1973-1978 collections

and the stomach contents of selected species. Institute laboratory facilities

were provided by Dr. A. F. Chestnut. Dr. S. T. Ross, Southern Mississippi

University, reviewed the manuscript and provided helpful comments.

R. Barnes produced Fig. 1. L. White typed the manuscript.

LITERATURE CITED

Armitage, T. ML, and W. S. Alevizon. 1980. The diet of the Florida pompano

(Trachinotus carolinus) along the east coast of central Florida. Florida

Science 43(l):19-26.

Bonsdorff, E., A. Norkko, and E. Sandbergy. 1995. Structuring zoobenthos:

the importance of predation, siphon cropping, and physical disturbance.

Journal of Experimental Marine Biology and Ecology 192:125-144.

Coen, L. D., and K. L. Heck. 1991. Interesting effects of siphon nipping

and habitat on bivalve {Mercenaria mercenaria (L.)) growth in a subtropical

seagrass {Halodule wrighti Askers) meadow. Journal of Experimental

Marine Biology and Ecology 145(1):1-14.

Compagno, L. J. V. 1990. Alternative life-history styles of cartilagenous

fishes in time and space. Environmental Biology of Fishes 28:33-75.

Currin, B. M. 1994. Food habits and food consumption of juvenile spot.

Leiostomus xanthurus, and croaker, Micropogonias undulatus, in their

nursery areas. M.S. Thesis. North Carolina State University, Raleigh.

Currin, B. M., J. P. Reed, and J. M. Miller. 1994. Growth, production,

food consumption, and mortality of juvenile spot and croaker: a com-

parison of tidal and nontidal nursery areas. Estuaries 7(4a):451-459.

Cyrus, D. P. 1988. Episodic events and estuaries: Effects of cyclonic flushing

in the benthic fauna and diet of Solea bleekeii (Teleostei) in lake St.

Lucia in the southeastern coast of Africa. Journal of Fish Biology 33

(Suppl. a):l-7.

Cyrus, D. P., and J. M. Blaber. 1983. The food and feeding ecology of

Gerreidae, Bleeker 1859, in the estuaries of Natal. Journal of Fish Biology

72:373-393.

Cyrus, D. P., and J. M. Blaber. 1984. The feeding ecology of Gerreidae

(Teleostei) in the Kosi system with special reference to their seasonal

diet. Lammanger 32:35-49.

Hines, A. N., A. M. Haddon, and L. A. Wiechert. 1990. Guild structure

and foraging impact of blue crabs and epibenthic fish in a subestuary

of Chesapeake Bay. Marine Ecology Progress Series 67:105-126.

Hughes, G. W. 1985. The comparative ecology and evidence for resource

partitioning in two pholid fishes (Pisces: Pholididae) from southern British

Columbia seagrass beds. Canadian Journal of Zoology 36(l):76-85.

Irlandi, E. A. 1993. Landscape ecology and the functions of barrier soft

44 Frank J. Schwartz

sediment habitats: How eelgrass landscapes influence growth survival

of a marine invertebrate in grass. Ph.D. Thesis. University of North

Carolina, Chapel Hill.

Joseph, D. C. 1962. Growth characteristics of two southern California surf

fishes, the California corbina and spotfin croaker, family Sciaenidae.

California Fish Game, Fish Bulletin 119:1-54.

Kammermans, P., and J. Huitema. 1994. Shrimp (Crangon crangon L).

browsing upon syphon tip inhibits feeding and growth in the bivalve

Macoma balthica (L.). Journal of Experimental Marine Biology and

Ecology 175:59-75.

Kvitek, R. G. 1991. Paralytic shellfish toxins sequestered by bivalves as

a defense against siphon-nipping fish. Marina Biology ll(3):369-374.

Kvitek, R. G., A. R. DeGange, and M. K. Bectler. 1991. Paralytic shell-

fish poisoning toxin mediate sea otter feeding behavior. Limnology and

Oceanography 36:393-404.

McMichael, R. H., and S. T. Ross. 1988. The relative abundance of feeding

habits of juvenile kingfish (Sciaenidae: Menticirrus) in a Gulf of Mexico

surf zone. Northeast Gulf Science 9(2): 109-123.

McMichael, S. 1986. The inflatable sharks, survival of the fattest. Sea

Frontiers 32(l):27-33.

Modde, T. C, and S. T. Ross. 1983. Trophic relationships of fishes oc-

curring within a surf zone habitat in the northern Gulf of Mexico. Northeast

Gulf Science 6(2):109-120.

Peterson, C. H., and M. L. Quammen. 1982. Siphon nipping: its impact

on growth of bivalve Protheca stamina (Conrad). Journal of Experi-

mental Marine Biology and Ecology 63:249-268.

Schwartz, F. J., W. T. Hogarth, and M. P. Weinstein. 1982. Marine and

freshwater fishes of the Cape Fear estuary, North Carolina, and their

distirbution in relation to environmental factors. Brimleyana 7:17-37.

Schwartz, F. J., P. Perschbacker, M. McAdams, L. Davidson, K. Sandoy, J.

Duncan, and D. Mason. 1979a. An ecological study of fishes and

invertebrate macrofauna utilizing the Cape Fear estuary, Carolina Beach,

and adjacent Atlantic Ocean. Summary Report 1973-1977. Institute Marine

Science, University of North Carolina, Morehead City.

Schwartz, F. J., P. Perschbacker, M. McAdams, L. Davidson, K. Sandoy, J.

Duncan, D. Mason, and J. Tate. 1979/?. An ecological study of fishes

and invertebrate macrofauna utilizing the Cape Fear estuary, Carolina

Beach, and adjacent Atlantic Ocean. Annual Report for 1978. Insti-

tute Marine Sciences, University of North Carolina, Morehead City.

Sutherland, W. C. 1982. Growth of two species of Macoma bivalvaes as

affected by predation on their siphons by juvenile spot, Leiostomus xanthurus,

and croaker, Micropogonias undulatus. M.S. Thesis. North Carolina

State University, Raleigh.

Welch, H. E., and K. Martin-Bergmann. 1990. Does the clam Mya truncata

regenerate its siphon after predation by walrus? An experimental ap-

proach. Arctic 43(2):157-158.

Siphon Nipping by Fishes 45

Zwarts, L., A. M. Blomest, P. Spaak, and B. Devier. 1994. Feeding ra-

dius, burying length, and siphon size of Macoma baltica and Scrobicularia

plana. Journal Experimental Marine Biology Ecology 183(2):193-212.

Received 2 August 1995

Accepted 15 February 1996

Condylura cristata (Insectivora: Talpidae)

in the Blue Ridge Province

of Western South Carolina

Joshua Laerm

Museum of Natural History, University of Georgia,

Athens, Georgia 30602

Gayle Livingston, Christine Spencer, and Bryan Stuart

Highlands Biological Station

Highlands, North Carolina 28741

The occurrence of Condylura cristata in western South Carolina

is questionable. Hall and Kelson (1959) and Hall (1981) mapped the

distribution of C. cristata to include northwestern South Carolina, provid-

ing as a marginal record a reference to Penny (1950:83) who, in turn,

citing Burnett (1851), places it on record from "Upper South Carolina."

Golley (1966:48), on the other hand, did not concur with Hall and

Kelson (1959). He cites "Pickens (1928) [who] states Burnett (1851)

lists the species from Aiken County." Lee (1987:57), in a very thorough

review of all distributional records of C. cristata in the southeastern

United States, alludes to the "Upper South Carolina" record as "probably

from Burnett (1851) who wrote on the fauna of the Pine Barrens of

Upper South Carolina. Thus, the record is from the Aiken County

area." Curiously, Penny (1950), Pickens (1928), and Lee (1987) all

misinterpret Burnett (1851). Pickens (1928:157) actually comments "Burnett

says he observed it at Aiken just below the fall line, the southern

boundary of the Piedmont." A careful reading of the Burnett reference

indicates this not the case. In fact, there is no geographic reference

whatsoever in the 1851 Proceedings of the Boston Society of Natural

History other than the introductory sentence (page 115) "Dr. Burnett

read some notes on the Fauna of the Pine Barrens of upper South

Carolina." Immediately following was a list of mammals observed,

including C. cristata. In the text Burnett does refer twice to the "pine

barren region" (Burnett, 1851:115, 116), but makes no further allusion

to locality. Even an approximate locality for the Burnett observation

is impossible, given the considerable extent of the upper South Carolina

Pine Barrens in 1851.

Thus, until now, no records of C. cristata are known from the

mountains of western South Carolina. We report here on the capture

of a single male specimen taken in a 5-gallon pitfall trap at the United

States Department of Agriculture Walhalla Fish Hatchery, Oconee County,

at 1.5 road miles north along the Fish Hatchery access road from

Brimleyana 24:46-49, April 1997 46

Condylura cristata in Western South Carolina 47

U.S. 107 (34° 12' 00" N, 83° 04' 11" W). The collection site was

a fern glade located in a relatively narrow, steep-walled gorge of the

East Fork of the Chattooga River dominated by an eastern hemlock

(Tsuga canadensis), white pine {Pinus strobus), and rhododendron

(Rhododendron maximum) streamside community which grades upslope

into yellow poplar (Liriodendron tulipifera), mixed oak (Quercus sp.),

and hickory (Carya spp.). Elevation was approximately 760 m. In addition

to the C. cristata specimen, six Sorex fumeus and four S. cinereus

were recovered in the pitfall. This collection site is the locality from

which the first state records of S. cinereus (Laerm et al. 1995) and

Clethrionomys gapperi (Pivorun et al. 1984) were reported. Additional

small mammals reported by Laerm et al. (In press) and Pivorun et

al. (1987) include S. hoyi, Blarina brevicauda, Peromyscus leucopus,

and P. maniculatus.

The star-nosed mole is documented in nearby areas of North Carolina

including the mountains "near the border of South Carolina" (Audubon

and Bachman 1851) and Clay, Henderson, Macon, Polk, and Transylvania

counties (Brimley 1945, Odum 1949, Johnston 1967, Lee et al. 1982,

Lee 1987, Webster 1987, Beane 1995). The nearest Blue Ridge locality

in Georgia is Union County (Laerm 1981).

Condylura cristata is apparently rare in the Blue Ridge or exceedingly

difficult to trap (Clark et al. 1985). Approximately 140,000 pitfall

and snap trap nights in western South Carolina by Laerm et al. (1995)

and Laerm et al. (In press) have failed to yield another specimen.

Lee (1982) noted the absence of C. cristata from the Piedmont of

North Carolina, South Carolina, and Georgia (see also Laerm 1981)

indicating populations in the Blue Ridge to be disjunct from those

on the Coastal Plain. However, Beane (1995) mapped undocumented

records of C. cristata from the Piedmont suggesting a possible continuous

distribution from the Coastal Plain to the Blue Ridge. We would not

concur that undocumented records are sufficient to justify such a sup-

position. Golley's (1966:49) map for C. cristata indicates a record

from the South Carolina Piedmont (Marion County) without supporting

comment or documentation, but neither Lee (1982) nor we have been

able to confirm this record. In so far as we have been able to ascertain,

there are no documented records from the Piedmont of South Carolina.

Laerm (1981) noted an unconfirmed report from the Piedmont (Jackson

County) of Georgia. The possible occurrence of C. cristata in the Piedmont

of these states is questionable. We concur with Beane (1995) that

efforts be made to provide documentation for the species in areas from

which it has not been confirmed.

48 Laerm, Livingston, Spencer, and Stuart

ACKNOWLEDGMENTS — We gratefully acknowledge support provided

by the U.S. Department of Agriculture Forest Service and support of

the University of Georgia Museum of Natural History.

LITERATURE CITED

Audubon, J. J., and J. Bachman. 1851. The quadrupeds of North America.

Volume 2. V.G. Audubon, New York, New York.

Beane, J. C. 1995. New distributional records for the star-nosed mole,

Condylura cristata (Insectivora: Talpidae), in North Carolina, with comments

on its occurrence in the Piedmont region. Brimleyana 22:77-86.

Brimley, C. S. 1944-1946. The mammals of North Carolina. Carolina Tips.

Carolina Biological Supply Company, Elon College, North Carolina.

Burnett, W. 1851. Notes on the pine barrens of upper South Carolina. Proceedings

of the Boston Society of Natural History 4:115-118.

Clark, M. K., D. S. Lee, and J. B. Funderburg, Jr. 1985. The mammal

fauna of Carolina bays, pocosins, and associated communities in North

Carolina: an overview. Brimleyana 11:1-38.

Golley, F. B. 1966. The mammals of South Carolina. Contributions from

the Charleston Museum XV, Charleston, South Carolina.

Hall, E. R. 1981. The mammals of North America. Second edition. John

Wiley & Sons, New York, New York.

Hall, E. R., and K. R. Kelson. 1959. The mammals of North America.

Ronald Press, New York, New York.

Johnston, D. W. 1967. Ecology and distribution of mammals at Highlands,

North Carolina. Journal of the Elisha Mitchell Scientific Society 83:

88-98.

Laerm, J. 1981. A survey of the status, distribution, and abundance of

potentially threatened and endangered vertebrates in Georgia. Part 4.

The mammals. University of Georgia Museum of Natural History Technical

Report, Athens.

Laerm, J., M. A. Menzel, E. Brown, A. Wotjalik, W. M. Ford, and M. Strayer.

The masked shrew, Sorex cinereus Kerr (Insectivora: Soricidae), and

red-backed vole, Clethrionomys gapperi (Rodentia: Muridae) in the Blue

Ridge Province of South Carolina. Brimleyana 22:15-21.

Laerm, J., W. M. Ford, T. S. McCay, M. A. Menzel, and J. L. Boone. In

press. Soricid communities in the southern Appalachians. Southern Appala-

chian Biogeorgraphy Symposium. Virginia Museum of Natural History,

Charlottesville.

Lee, D. S. 1987. The star-nosed mole on the Delmarva Peninsula: zoogeo-

graphic and systematic problems of a boreal species in the South. The

Maryland Naturalist 31:44-57.

Lee, D. S., J. B. Funderburg, Jr., and M. K. Clark. 1982. A distribu-

tional survey of North Carolina mammals. Occasional papers of the North

Carolina Biological Survey, Raleigh.

Condylura cristata in Western South Carolina 49

Penny, J. T. 1950. Distribution and bibliography of the mammals of South

Carolina. Journal of Mammalogy 31:81-89.

Pickins, A. L. 1928. Mammals of upper South Carolina. Journal of Mam-

malogy 9:155-157.

Pivorun, E. B., D. H. Allen, and D. T. Sawyer. 1984. First record of

Clethrionomys gapperi (Mammalia: Rodentia) in South Carolina. Jour-

nal of the Elisha Mitchell Society 100:33.

Odum, E. P. 1949. Small mammals of the Highlands (North Carolina) Plateau.

Journal of Mammalogy 30:179-192.

Webster, W. D. 1987. Condylura cristata parva Paradiso. Pages 42-43

in Endangered, threatened and rare fauna of North Carolina. Part 1.

A reevaluation of the mammals (M. K. Clark, editor) Occasional Pa-

pers of the North Carolina Biological Survey, Raleigh.

Received 24 October 1995

Accepted 15 Feburary 1996

Frontispiece. Theatops posticus. Photograph of live individual on substrate

at Black Mountain Campground near Mt. Mitchell, Yancey County, North

Carolina (courtesy of R. L. Hoffman).

The Holarctic Centipede Subfamily Plutoniuminae

(Chilopoda: Scolopendromorpha: Cryptopidae) (Nomen

Correctum Ex Subfamily Plutoniinae Bollman, 1893)

Rowland M. Shelley

North Carolina State Museum of Natural Sciences,

P. O. Box 29555, Raleigh, North Carolina 27626-0555, U.S.A.

ABSTRACT — The Holarctic chilopod subfamily Plutoniuminae Bollman,

a corrected name for Plutoniinae, consists of two genera, Plutonium

Cavanna and Theatops Newport, and six species; synapomorphies

between them show that the subfamily is a monophyletic group and

that the different number of spiracles, 19 pairs in Plutonium and

9 pairs in Theatops, is only a generic-level character. Plutonium

and P. zwierleini Cavanna occur in Sicily, Sardinia, Napoli and Sorrento

provinces in mainland Italy, and Granada Province, Spain. Theatops

erythrocephalus (C. L. Koch) occurs along the eastern side of the

Adriatic Sea in the Balkan Peninsula and in coastal Spain and Portugal.

The other four species — T. posticus (Say), T. spinicaudus (Wood),

T. phanus Chamberlin, and T. californiensis Chamberlin — occur in

the United States and northwestern Mexico. Theatops posticus oc-

cupies a broad area east of the Central Plains from Connecticut and

southern New York to the south Florida keys and eastern Texas;

an allopatric western population extends from southwestern New Mexico

and western Chihuahua to the southern Great Basin, the Califor-

nia desert east of the Sierra Nevada, the Pacific Ocean in Baja California

Norte, the Channel Islands off the southern California coast, and

the eastern slope of the Coast Range near the latitude of San Francisco

Bay. Theatops spinicaudus occurs sympatrically with T. posticus in

two areas of the east; the inner surfaces of its caudal legs possess

variable series of ridges and teeth. Theatops phanus occurs in epigean

and subterranean environments in southern Texas and extends from

east of highway 1-35 to west of the Pecos River; the inner surfaces

of its caudal legs also possess variable series of ridges and teeth.

The distribution of T. californiensis, anatomically convergent with

T. erythrocephalus, is as described previously, but locality infor-

mation is detailed, as only one site, the type locality, is currently

known. Relationships among the plutoniuminine species are postu-

lated as P. zwierleini + (T. spinicau-dus + (T. phanus + (T erythrocephalus

+ (T. posticus + T. californiensis)))). The Plutoniuminae and Cryptopinae

logically share ancestry, and the Scolopocryptopinae may warrant

elevation to family status.

Brimleyana 24:51-113, April 1997 51

52 Rowland M. Shelley

GENERAL INTRODUCTION

Introduction

Among the more readily identified scolopendromorph centipedes

in North America and Europe, ones with 21 or 23 pairs of legs and

pedal segments, are the representatives of the cryptopid subfamily Pluto-

niuminae Bollman,1 a senior subjective synonym of Theatopinae Verhoeff,

occasionally misspelled as "Theatopsinae."2 Recognizable to the unaided

eye by their extremely robust ultimate legs (Fig. 1; see also Attems

1926, Fig. 433, and Shelley 1990a, Fig. 1), they also feature 21 leg

pairs, a long caudal segment, roughly twice as long as the penultimate,

Fig. 1. Theatops posticus, dorsal view. Scale line = 1.00 cm.

and pale, lightly pigmented patches in the ocellar positions, lateral to

the bases of the antennae. Crabill (1977) referred to the patches as

"eyespots," an unfortunate term because these blind chilopods lack photoreceptors.

The heavily sclerotized, forcipulate caudal legs are the most visible diagnostic

feature, and according to Cloudsley-Thompson (1958) and Manton (1965),

function to hold food. The plutoniuminines are thus convergent in this

regard with the scolopendrid genus Cupipes Kohlrausch (compare Figs.

2-5, with Figs. 6-9).

The Plutoniuminae is comprised of two genera — Plutonium Cavanna,

monotypic with P. zwierleini Cavanna occurring in Spain and mainland

Italy, Sicily, and Sardinia, and Theatops Newport, with one Palearctic

!As noted by Wurmli (1975), Plutoniinae Bollman (1893) has priority by 13 years over

Theatopinae Verhoeff (1906) as the senior name for this family-group taxon. However,

Plutoniinae Bollman is preoccupied by the senior homonym, Plutoniinae Cockerell (1893)

(Mollusca: Gastropoda), which has priority by one month. Shelley and Backeljau (1995)

petitioned the International Commission on Zoological Nomenclature to remove the homonymy

but incorrectly believed Plutoniinae Bollman was the older name. By agreement of all

parties, Plutoniinae Bollman is being emended to "Plutoniuminae"; this decision will be

announced in a forthcoming issue of Bulletin of Zoological Nomenclature. The present

contribution is the first taxonomic usage of the corrected name.

2Confusion has existed as to formation of family-group names from genera with the

"-ops" suffix, whether the "s" is retained or dropped. The genitive of this ending is "opis",

of which the "is" is dropped to form family-group names, so the correct spelling is "Theatopinae"

rather than "Theatopsinae."

Centipede Subfamily Plutoniuminae

5 3

Figs. 2-5. Cupipes spp. 2, ultimate legs and segment of species from Trinidad

taken in cargo at Honolulu, Hawaii (NMNH), dorsal view. 3, the same, ventral

view. 4, ultimate legs and segment of species from Isle of Palms, Cuba (NMNH),

dorsal view. 5, the same, ventral view. Scale lines = 1.00 mm for each figure.

54 Rowland M. Shelley

and four Nearctic species. Both the subfamily and the genus Theatops

thus demonstrate Holarctic/Laurasian distribution patterns, and their

biogeographies are intriguing. In addition to the North American/Euro-

pean disjunction in Theatops, allopatric populations exist in three species.

In Europe, T erythrocephalus3 (C. L. Koch) occurs in Croatia, Montenegro,

Bosnia-Hercegovina,4 and the Iberian peninsula, thus exhibiting a hiatus

of some 992 km (620 mi) that is partly occupied by P. zwierleini.

In North America, T. posticus (Say) occurs east of the Great Plains

and in the desert southwest, with an intervening gap of some 1200

km (750 mi) (Shelley 1990a), and T. spinicaudus (Wood) occupies

two regions in the eastern states segregated by a lacuna ranging from

368-688 km (230-430 mi) (Fig. 31). Because of the extensiveness of

past collecting, these lacunae are undoubtedly real and are unlikely

to change significantly with future discoveries.

A third genus, Tonkinodentus Schileyko (1992), monotypic from

Viet Nam, was assigned to the Theatopinae, but the type and only

specimen of its species, Tonkinodentus lestes Schileyko, is missing

the last three leg pairs, the caudalmost of which possesses most of

the taxonomically critical characters in this subfamily. This genus occurs

some 11,200 km (7,000 mi) east-southeast of the most proximate locality

of Theatops erythrocephalus in Montenegro and some 11,920 km (7,450

mi) west-southwest of that of Theatops posticus in Mexico, and is

hence implausible for the Plutoniuminae, which is otherwise geographically

coherent. I therefore remove Tonkinodentus from the subfamily and

leave it unassigned; proper placement awaits the discovery of fresh

material, preferably several individuals, possessing all 21 leg pairs.

It should be emphasized in this regard that the proposal of a new

taxon is a serious action involving the placement of a new entry on

the roster of available scientific names. Future students will have to

consider this taxon and address shortcomings in the original account,

completely rediagnosing it if necessary, and the proliferation of poorly

conceived taxa and substandard accounts by past authors is a major

reason for the nascency of myriapodology. It is therefore imperative

3Like most chilopods with the -ops generic suffix, confusion has existed over the gender

of Theatops and whether it requires the feminine or masculine form of the species-group

name. Thus, Kraepelin (1903) reported T. erythrocephalus whereas Attems (1930) cited

T. erythrocephala. I (Shelley 1987) reviewed this situation in footnote 2 and noted that

article 30 (a) (ii) of the 1984 edition of the International Code of Zoological Nomen-

clature supercedes past recommendations and declares that genus-group names ending in

-ops are to be considered masculine regardless of derivation or treatment by the author.

4The countries of Slovenia, Croatia, Macedonia, Bosnia-Hercegovina, Montenegro, and Serbia

comprise the former country of Yugoslavia.

Centipede Subfamily Plutoniuminae 55

that modern myriapod taxonomy be soundly based and not recapitulate

this heritage, and the ultimate, and often penultimate, legs hold taxonomic

importance in many scolopendromorph genera. The erection of new

genera for anatomically incomplete chilopods, taxa that future workers

will be compelled to reconceptualize, cannot be too strongly discouraged.

Another intriguing aspect of Theatops is the nearly identical external

structures of T. erythrocephalus, in Europe, and T californiensis Chamberlin,

in California and Oregon, which are separated by around 11,520 km

(7,200 mi). This resemblance was interpreted as convergence by Shelley

(1990a), and it has produced especially similar external anatomies;

the species are so nearly identical that confusion reigned for 88 years

over the correct name for the latter. Past authors labeled it T californiensis

(Chamberlin 1902), T erythrocephalus californiensis (Chamberlin 1911),

and T. erythrocephala (Attems 1930, Chamberlin 1951a). Crabill (1960)

and Kevan (1983) called it T. californiensis but suggested synonymy

with T. erythrocephala. This enigma was resolved by Shelley (1990a),

who deduced from variation in southwestern forms of T. posticus that

T californiensis is a valid species and hence that the name T erythrocephalus

applies exclusively to the European representative. Many of the geographi-

cally intermediate southwestern variants of T. posticus display anatomical

conditions that are intermediate between those of its eastern population

and T. californiensis, showing that these taxa were once united in a

single species spanning the breadth of North America through the

southern United States and the adjacent part of Mexico (Shelley 1990a).

This knowledge indicates that the phenotypic resemblance between T.

californiensis and T. erythrocephalus represents convergence, but it

is an especially perfect example with no clear differences in their

external anatomies. At present, I can only use geography to distinguish

them in the key and assign specific names, and the search for differences

requires substantially more critical information as might derive from

comparative biochemistry. Theatops californiensis and T. erythrocephalus

are thus prime candidates for investigation by immunological techniques

and .electrophoresis of haemolymph proteins.

Still another fascinating aspect of Theatops is the adaptability

of T. phanus Chamberlin, which is known only from caves in Sutton

and Menard counties, Texas, where it displays troglobitic adaptations.

These include pallid color and long, slender appendages, the antennae

reaching back to tergites 6-7, instead of to tergites 3-4, the antennomeres

being three to five times longer than wide, instead of about twice

as long, and the podomeres on the penultimate legs being four to

five times longer than wide, as opposed to only two to three times

longer (Weaver 1982). Some cave specimens are quite large, and the

56 Rowland M. Shelley

anatomical modifications are visibly striking. However, while examining

museum collections, I discovered individuals from epigean populations

that lack these features; the relative proportions of their podomeres

and antennomeres are similar to those of T. spinicaudus and T posticus.

These surface forms are geographically proximate to, and clearly con-

specific with, cave populations of T. phanus, which is consequently

a highly variable species demonstrating a level of genetic plasticity

that is not apparent in other Nearctic cryptopids. Occasional specimens

of Scolopocryptops sexspinosus (Say) have been discovered in caves,

but they are identical anatomically to epigean individuals and are not

modified by the subterranean environment.

This study of the Plutoniuminae, the first monographic treatment

of a supra-generic chilopod taxon since Attems' (1930) ordinal treatise

on the Scolopendromorpha, derives from an ongoing survey of North

American scolopendromorphs. The discovery of the widespread, anatomi-

cally variable southwestern population of T. posticus and the resultant

deduction of the correct binomial for T. californiensis (Shelley 1990a)

focused attention on Theatops and led to the deletions of T. spinicaudus

from Hawaii, Mexico, and Canada (Shelley 1990a, 1991). These works

indicated that T. erythrocephalus should be examined in its proper

context, along with the American congeners, and that P. zwierleini

should be included to consolidate knowledge of the subfamily. The

Plutoniuminae is thus one of the few chilopod family-group taxa that

is amenable to a modern, all-inclusive treatment, because of its limited

composition and occurrence in only two biogeographic regions. Because

of the difference in the number of spiracles, Verhoeff (1906, 1907)

and Schileyko (1992, 1996) believed that separate families were required

for Plutonium and Theatops, and Schileyko (1996) even suggested that

Plutonium deserved a separate superfamily as an "absolutely different

group." However, most authors, including myself, subscribe to the system

originated by Attems (1930), in which Theatops and Plutonium are

assigned to the same subfamily. Such features as the depigmented patches

in the ocellar positions, the elongated ultimate tergite, and the heavily

sclerotized, forcipulate caudal legs constitute strong synapomorphies

that unite Theatops and Plutonium in a monophyletic group. Although

unique to the Scolopendromorpha, the autapomorphic 19 pairs of spiracles

in Plutonium is only a generic feature; using it as the basis for a

separate family or superfamily overemphasizes the character's signifi-

cance in relation to the several attributes that are shared with Theatops,

which indicate common ancestry. Consequently, I believe that the present

concept of the Plutoniuminae represents a natural assemblage of related

taxa.

Centipede Subfamily Plutoniuminae 57

This contribution presents diagnoses of all components of the

Plutoniuminae, complete synonymies at all levels, detailed range descrip-

tions with locality data for each species, and discussions of anatomical

variation, ecology, and relationships. Complete citations for all available

samples of T. posticus and T. spinicaudus would be prohibitively long,

so these are summarized for certain states as indicated in each account.

Acronyms of repositories of preserved study material are as follows:

AAW — Private collection of A. A. Weaver, Wooster, Ohio.

AMNH — American Museum of Natural History, New York, New York.

ANSP — Academy of Natural Sciences, Philadelphia, Pennsylvania.

AU — Entomology Department, Auburn University, Auburn, Alabama.

CDFA — California Department of Food and Agriculture, Sacramento.

CAS — California Academy of Sciences, San Francisco.

DC — Natural Science Division, Dixie College, St. George, Utah.

EIU — Zoology Department, Eastern Illinois University, Charleston.

FMNH — Field Museum of Natural History, Chicago, Illinois.

FSCA — Florida State Collection of Arthropods, Gainesville.

INHS — Illinois Natural History Survey, Urbana.

LACMNH — Los Angeles County Museum of Natural History, Los Angeles,

California.

LEM — Lyman Entomological Museum, MacDonald College, McGill

University, Ste. Anne de Bellevue, Quebec, Canada.

MCZ — Museum of Comparative Zoology, Harvard University, Cambridge,

Massachusetts.

MEM — Mississippi Entomological Museum, Mississippi State University,

Starkville.

MPM — Milwaukee Public Museum, Milwaukee, Wisconsin.

NCSM — North Carolina State Museum of Natural Sciences, Raleigh.

NHM — The Natural History Museum, London, England.

NMNH — National Museum of Natural History, Smithsonian Institution,

Washington, DC.

OPCNM — Organ Pipe Cactus National Monument, Arizona.

SEM — Snow Entomological Museum, University of Kansas, Lawrence.

SREL — Savannah River Ecological Laboratory, Aiken, South Carolina.

SWRS — Southwest Research Station, Portal, Arizona.

TMM — Texas Memorial Museum, University of Texas, Austin.

UAR — University of Arkansas Arthropod Museum, Fayetteville.

UAZ — Entomology Department, University of Arizona, Tucson.

UCB — Essig Museum of Entomology, University of California at Berkeley.

UCD — Bohart Entomological Museum, University of California at Davis.

UCR — Entomology Department, University of California at Riverside.

58 Rowland M. Shelley

UCT — Zoology Department, University of Connecticut, Storrs.

UGA — Zoology Department, University of Georgia, Athens.

UL — Biology Department, University of Louisville, Kentucky.

UMMZ — University of Michigan Museum of Zoology, Ann Arbor.

UMO — Enns Entomological Museum, University of Missouri, Columbia.

VMNH — Virginia Museum of Natural History, Martinsville.

WAS — Private collection of W. A. Shear, Hampden-Sydney, Virginia.

WVDA — West Virginia Department of Agriculture, Charleston.

ZMH — Zoologisches Institut und Museum, Universtitat Hamburg, Germany.

ZMUC — Zoological Museum, University of Copenhagen, Denmark.

LITERATURE REVIEW

The literature of the Plutoniuminae is relatively orderly; difficulties

arose primarily from the tendency of early authors to inconsistently

cite species under more than one genus, and an erroneous observation

by Newport (1844) in his proposal of the oldest genus-group name,

Theatops. Newport mistook the unpigmented spots in the ocellar positions

as eyes, thus stating in the original and subsequent generic accounts

(Newport 1844, 1845, 1856) "ocelli distincti." However, he contradicted

this statement in the accompanying species accounts with the phrase,

"ocellis inconspicuis lateralibus." Confusion thus developed as to whether

Theatops and its type species, Cryptops postica Say, did or did not

possess ocelli, which was partly responsible for Wood's proposal (1862)

of the genus Opisthemega. Because the specimen of C. postica on

which Newport's proposal was based was sent to him at the NHM

by Say, no one else had seen it and could unequivocally resolve the

question of eyes. Underwood (1887) reviewed the confusion in footnote

8 and concluded that Theatops "may as well be consigned to oblivion"

and "at least it is not necessary to include it in future lists." These

statements concerned R. I. Pocock, who was at the NHM and in a

position to settle the issue by reexamining Say's specimen. He did

so; reported (Pocock 1888) that it lacked eyes and that Newport was

mistaken; and synonymized Opisthemega with Theatops. Pocock 's analy-

sis was accepted by subsequent authors, and future problems in the

Plutoniuminae chiefly involved misidentifications, a few ill-conceived

proposals of synonyms, and disagreement about the name and taxonomic

status of T. calif orniensis.

The history of the Plutoniuminae begins with the description of

C. postica for a specimen from Georgia or east Florida by Say (1821),

a binomial subsequently cited by Newport (1844), Kohlrausch (1881),

and, in the masculine gender, by Lucas (1840) and Bollman (1893).

One of the first dozen or so centipedes to be described from North

Centipede Subfamily Plutoniuminae 59

America, it became the type species of Theatops (Newport 1844), based

on the aforementioned specimen from Say. Gervais (1847) placed Theatops

under Cryptops, though indicating that it might refer to a form of

Scolopendra. Koch (1847) described C. erythrocephalus from Pula,

Croatia, on the Istrian peninsula, and enhanced the description in

an expanded account (1863) with a full-length illustration. He did

not connect this centipede with Say's species and said nothing about

Theatops, if he was even aware of Newport's taxon. Wood (1862)

proposed Opisthemega for two ostensibly new species, O. postica and

O. spinicauda, from North Carolina and Illinois, respectively, without

designating either as type species. Wood stated that O. postica lacked

eyes; questioned whether it was identical to Say's species because it

agreed with T. postica except for the eyes and teeth; and suggested

that Newport might be mistaken about the presence of ocelli. However,

he followed these accounts with others on Theatops and T. postica

stating, "We have never seen a specimen of this species." Wood (1865)

repeated his previous accounts of all these taxa, adding west Pennsylvania

to the localities of O. spinicaudus. Cope (1869) cited O. postica from

the mountains of southwestern Virginia, misspelling Wood's genus

as "Opisthomega" and Saussure and Humbert (1872) repeated the previous

names and localities.

In the ensuing decade, Latzel (1880) tentatively placed Theatops

in synonymy under Scolopendra. He recognized Opisthemega and trans-

ferred Koch's species into this genus, forming the new combination,

O. erythrocephalum. Kohlrausch (1881) recognized both Theatops and

Opisthemega; included postica under both names and Cryptops; cited

erythrocephalus under Cryptops and Opisthemega; but listed spinicauda

under Opisthemega only. Cavanna (1881) erected Plutonium for a new

centipede from Sicily, P. zwierleini, possessing spiracles on segments

2-20. Meinert (1886) recorded O. spinicauda from Acapulco, Mexico;

proposed the synonym, O. insulare, for specimens ostensibly from Hawaii,

then called the Sandwich Islands; and erected O. crassipes for specimens

from Florida, Virginia, and Kentucky. Though he questioned its distinction

from O. postica, McNeill (1887, 1888)5 recorded O. crassipes from

Indiana and Escambia County, Florida. In addition to synonymizing

Opisthemega with Theatops, Pocock (1888) also placed O. postica and

O. crassipes under T. postica. The remaining publications of this decade

belong to Bollman (1888a-e), who cited the new combination T crassipes,

placed postica under Cryptops, Theatops, and Opisthemega, and reported

5To conserve space, subsequent publications that merely provide new localities are summarized

in the species listings at the conclusion of this section.

Rowland M. Shelley

several new localities for this species and for T. spinicaudus.

In the 1890's, Bollman (1893) established the Plutoniinae and

attempted the first general range descriptions for T. posticus and T.

spinicauda,6 cited in the ensuing listings. He recorded the latter from

the southwestern United States in general, a citation not justified by

any collection or previous records and the probable source for future

erroneous listings from California and the southwest (Chamberlin 1902,

Crabill 1960). Verhoeff (1896) proposed O. lusitanum for a centipede

from Portugal and attempted to contrast it with O. erythrocephalus.

The twentieth century began with the proposal of T. californiensis

for a form from Quincy, Plumas County, California, by Chamberlin

(1902). He also summarized the synonymies and distributions of T.

posticus and T. spinicaudus, largely repeating the ranges reported by

Bollman (1893), and provided a key to the then three American species.

Kraepelin (1903) assigned Chamberlin's species to synonymy under

T. erythrocephalus and recorded it from Oregon and California; he

also provided a key to T. posticus, T. spinicaudus, and T. erythrocephalus,

along with synoptic accounts to these species, P. zwierleini, and the

genera Theatops and Plutonium. Other new synonymies proposed by

Kraepelin (1903) include O. insulare under T. spinicaudus and O.

lusitanum under T. erythrocephalus, the last binomial being a new

combination. His and subsequent listings of T. erythrocephalus from

Italy are erroneous and refer instead to P. zwierleini (Minelli, in litt.),

and one also wonders about the source for Kraepelin's record from

Oregon. Theatops californiensis had only been described the previous

year, and Quincy, the only locality Chamberlin (1902) listed, is too

distant from Oregon (ca. 224 km [140 mi]) to imply occurrence in

that state. I know of no pre-1903 Oregon specimens, and if geographic

proximity were the basis for Kraepelin's citation, one would expect

him to choose Nevada, since Quincy is only about 80 km (50 mi)

from this state. The basis for the sudden Oregon citation is thus a

mystery, but it is nevertheless correct as shown by recent samples

from Douglas and Josephine counties. Three years later, Verhoeff (1906)

proposed the family "Theatopsidae."

In ensuing decades, Chamberlin (1911) reduced his species to a

race of T. erythrocephalus. Gunthorp (1920) reviewed Wood's papers

and authorships, and credited him with Opisthemega, O. postica, and

6Bollman's inconsistencies are noteworthy. He (1888a, c) cited spinicaudus, suddenly

changing (1893) without explanation to the feminine termination. However, he simulta-

neously and consistently employed the masculine suffix for posticus (1888c-e, 1893),

but he suddenly and without explanation cited it (1893) under both Cryptops and Theatops,

whereas he previously (1888c-e) used only the latter genus.

Centipede Subfamily Plutoniuminae 61

O. spinicauda without alluding to the confusion surrounding Opisthemega

and postica or the synonymy with Theatops proposed by Pocock (1888).

Attems (1930) published the last comprehensive work on the Scolopendro-

morpha, recognizing Plutonium, P. zwierleini, Theatops, T. postica,

T. erythrocephala, and T. spinicauda. He provided a key to the species

of Theatops and summarized locality information since Kraepelin's

work (1903). Attems (1938) included T. spinicauda among the Hawaiian

fauna, a record deleted by Shelley (1991). Chamberlin (1951/?) described

the fourth American and fifth total species of Theatops, T. phanus,

from a cave in Sonora County, Texas, and presented a key to species.

He recognized only four species, implying that he then considered

T. calif or niensis to be a synonym of T. erythrocephala. Crabill (1957)

reviewed Newport's chilopod genera, credited him with authoring Theatops,

and indicated that the type species, C. postica Say, was fixed by subsequent

monotypy. Matic (1960) proposed T erythrocephala breuili for a specimen

from a Spanish cave. Crabill (1960) included Theatops and Theatopinae

in a key to North American scolopendromorph genera, subfamilies,

and families, and characterized the ranges of the four North American

species. He considered T. californiensis to be a species, but suggested

that it might be a synonym of T. erythrocephala.

In recent years, Wiirmli (1975) reported authentic localities for

P. zwierleini and provided a distribution map. In concluding paragraphs,

he reviewed the distribution of T. erythrocephalus, which he considered

as including California and Oregon, thus implying synonymy of T.

californiensis, placed T. e. breuili in synonymy, and noted that Plutoniinae

Bollman antedates Theatopidae Verhoeff. Crabill (1977) included Theatops

and Theatopinae in a key to North American and Mexican cryptopid

taxa, and Summers (1979) included T. posticus, T. spinicaudus, and

the Theatopinae in a key and taxonomic listing to centipedes of the

north-central United States.

In a definitive text on centipede biology, Lewis (1981) reported

the results of Baerg (1924) on the effect of centipede bites, noting

that T. spinicauda had little to no effect on rats and caused sharp

pain in humans that disappeared in 30 minutes. Lewis recognized the

subfamily Theatopsinae with Plutonium, in Sicily, Sardinia, and Campania,

and Theatops, in North America, the Mediterranean region, and Hawaii.

He noted that Plutonium has 19 pairs of spiracles, one each on segments

2-20, or all leg bearing segments except the first and last, instead

of the nine pairs typically found in scolopendromorphs with 21 segments.

Kevan (1983) reported the northernmost records for T postica, T spinicauda,

and T. californiensis and questioned whether the last name was a synonym

of T. erythrocephala. Shelley and Edwards (1987) reported T. posticus

62 Rowland M. Shelley

statewide from Florida and presented a distribution map, and Shelley

(1987) reported general distributions of T. posticus and T. spinicaudus

in North Carolina, showing counties of occurrence on a locality map.

In the latest four papers on the Plutoniuminae, Shelley (1990a,

1991) deleted T. spinicaudus from Hawaii, Mexico, and Canada; reported

the first Mexican localities for T. posticus', demonstrated broad occurrence

of this species in the southwestern United States; and deduced that

T. calif or niensis is the correct binomial for the congener in northern

California and Oregon. Hence, he restricted the name T. erythrocephalus

to the European species. In a paper on Yugoslavian centipedes Kos

(1992) characterized T. erythrocephalus as a mediterranean to sub-

mediterranean species and recorded it from Croatia, Bosnia-Hercegovina,

and Montenegro. Finally, Hoffman (1995) reported T. posticus from

14 counties in Virginia. Thus at this writing, plutoniuminine species

have been reported from the following states in Mexico, counties and

states in the United States, and islands and general regions of European

countries. The published overall range statements are summarized first

for T. posticus, T. spinicaudus, and T. calif orniensis, and where appropriate,

the general range statements for each state are presented first under

each state, for the American species, and under each country or island,

for the European representatives.

Theatops posticus

USA: General range statements — East of the Mississippi River,

north to Virginia, Indiana, and Illinois (Bollman 1893); the eastern

United States south of Virginia, Indiana, and Illinois (Chamberlin 1902);

sporadic in Utah and Arizona, and ranging from southern Illinois,

Ohio, and northern Virginia south to the Gulf States (Crabill 1960);

and the eastern United States generally south of the Great Lakes and

east of the Central Plains, extending along the Gulf of Mexico to

San Patricio County, and inland to Limestone County, Texas (Shelley

1990a).

Indiana— Bloomington, Monroe Co. (McNeill 1887, Bollman 1888e);

Dublin, Wayne Co., and Brookville, Franklin Co. (McNeill 1887); New

Providence, Clark Co., and Wyandotte, Crawford Co. (Bollman 1888e).

Illinois — Gallatin, Hardin, Pope, Jackson, and Pulaski counties

(Summers et al. 1980, 1981).

Ohio — southern Ohio in general (Kevan 1983); southeastern Ohio

in general (Williams and Hefner 1928); Vinton, Gallia Co. (Morse

1902).

Virginia — Virginia in general (Underwood 1887); southwestern

Virginia in general (Cope 1869); Lee Co. (Meinert 1886); Luray, Page

Centipede Subfamily Plutoniuminae 63

Co., and Natural Bridge, Rockbridge Co. (Bollman 1888d); Albemarle,

Alleghany, Botetourt, Buchanan, Floyd, Greenville, Henry, Lee, Mont-

gomery, Page, Patrick, Pittsylvania, Rockbridge, and Rockingham cos.

(Hoffman 1995).

Kentucky — Kentucky in general (Underwood 1887, McNeill 1888);

Jessamine Co. (Crabill 1955a); Bee Spring, Edmonds Co. (Meinert

1886, Crabill 1955a); Pine Ridge, Wolfe Co. (Crabill 1955a, Branson

and Batch 1967); Louisville, Jefferson Co., Campbellsville, Taylor Co.,

Cumberland Falls State Park, Whitley Co., near Livingston, Rockcastle

Co., and near Irvine, Estill Co. (Crabill 1955a); and along Red R.,

Powell Co. (Branson and Batch 1967).

Tennessee — Beaver Cr., Jefferson Co. (Bollman 1888/?); Mossy

Creek, Jefferson Co. (Bollman 1888c); Glendale Hills, Davidson Co.

(Chamberlin 1918a).

Carolina in general without specifying the state (Kraepelin 1903,

Attems 1930).

North Carolina — North Carolina in general (Wood 1865, Saussure

and Humbert 1872, Kohlrausch 1881, Underwood 1887, Brolemann

1896, Kraepelin 1903); North Carolina in general, but rare in mountains,

specific counties shown in Fig. 13 (Shelley 1987); Goldsboro, Wayne

Co. (Wood 1862, Bollman 1888o\ Brimley 1938, Wray 1950, 1967);

Hendersonville, Henderson Co., and Duke Forest, Durham Co. (Brimley

1938, Wray 1950, 1967); Greensboro, Guilford Co. (Causey 1940).

South Carolina— Clemson, Pickens Co. (Crabill 1950).

Georgia — Georgia in general (Say 1821, Lucas 1840, Gervais 1847,

Saussure and Humbert 1872, Kohlrausch 1881, Pocock 1888, Attems

1930); Macon, Bibb Co. (Bollman 1888rf); Okefenokee Swamp, Ware

Co. (Chamberlin 1918b); near Sylvania, Screven Co., Savannah, Chatham

Co., and near Pendergrass, Jackson Co. (Chamberlin 1944a); and Camilla,

Mitchell Co. (Chamberlin 1945).

Florida — Statewide (Shelley and Edwards 1987); Florida in general

(Underwood 1887, McNeill 1888, Kraepelin 1903, Attems 1930); East

Florida (Say 1821, Lucas 1840, Gervais 1847, Saussure and Humbert

1872, Kohlrausch 1881, Pocock 1888). Jacksonville, Duval Co., and

St. Johns River, county uncertain (Meinert 1886); Escambia Co. in

general (McNeill 1887); Archbold Biological Station, Lake Placid, High-

lands Co., and Gainesville, Alachua Co. (Chamberlin 1951a).

Louisiana — Louisiana in general (Brolemann 1896, Attems 1930).

Creston, Natchitoches Par. (Chamberlin 19186); Covington, St. Tammany

Par. (Viosca 1918).

Texas — Houston, Harris Co. (Chamberlin 1943).

New Mexico — Peloncillo Mountains, Hidalgo Co. (Shelley 1990a).

64 Rowland M. Shelley

Arizona — Rincon Mountain, Pima Co. (Chamberlin 1944b); numerous

localities in Coconino, Yavapai, Gila, Maricopa, Graham, Pinal, Yuma,

Pima, Santa Cruz, and Cochise cos. (Shelley 1990a).

Utah — Washington Co. in general (Chamberlin 1925, Shelley 1990a).

St. George, Washington Co. (Shelley 1990a).

Nevada — Nelson, Clark Co., and Nuclear Testing Site, Pahute

Mesa, and Rocky Valley, Nye Co. (Shelley 1990a).

California — Santa Cruz Island, Channel Islands National Park,

and 48 km (30 mi) SW Palm Desert, Riverside Co. (Shelley 1990a).

MEXICO: Chihuahua— 51 km (31.8 mi) S Minaca (Shelley 1990a).

Sonora— 5 km (3.1 mi) NW Huicoche (Shelley 1990a).

Baja California Norte — La Turquesa, 23.2 km (14.5 mi) N

Ensenada, and 22.4 km (14 mi) S US border (Shelley 1990a).

Theatops spinicaudus

General range statements — Southwestern United States in general,

north to Chicago, east in Tennessee to the mountains, and north to

Allegheny Co., Pennsylvania (Bollman 1893); southwestern United States

in general, Tennessee to Pennsylvania (Chamberlin 1902); Hawaiian

Islands and southeastern United States (Chamberlin 1920); America

north of Mexico (Biicherl 1942); western Pennsylvania to the Gulf

Coast west to Missouri and Arkansas (Crabill 1955b); Mexico and

California without further specification, northern Missouri and Illinois

to the Gulf States north through the Carolinas and up the Coastal

Plain possibly to southern Pennsylvania (Crabill 1960); the eastern

United States generally south of the Great Lakes and east of the Central

Plains, being unknown beyond southwestern Arkansas, eastern Oklahoma,

and central Iowa (Shelley 1990a).

Illinois — Illinois in general (Underwood 1887); southern Illinois

(Wood 1862, 1865, Saussure and Humbert 1872, Kohlrausch 1881,

Kraepelin 1903, Attems 1930); Chicago and Cook Co. (Wood 1862,

1865, Auerbach 1951a, b); Alto Pass, Union Co. (Chamberlin 1944b);

and Cook, Champaign, McLean, Greene, Union, Gallatin, Johnson,

Pope, Union, Jackson, Williamson, Randolph, and Pulaski cos. (Summers

et al. 1980, 1981).

Missouri— St. Charles, St. Charles Co. (Chamberlin 1928, Crabill

1955); Libertyville, St. Francois Co. (Chamberlin 1944b); Ranken, St.

Louis, Chesterfield, and Glencoe, St. Louis Co.; Sullivan, Franklin

Co.; and High Ridge and Vaugirard, Jefferson Co. (Crabill 1955c).

Arkansas — Near Oliphant, Jackson Co. (Chamberlin 1942); near

Hot Springs, Garland Co.; Ben Lomond, Sevier Co.; Mt. Magazine,

Logan Co.; Arkadelphia, Clark Co.; and Delight, Pike Co. (Chamberlin

Centipede Subfamily Plutoniuminae 65

19446); Little Rock, Pulaski Co., and Ultima Thule, county unknown

(Bollman 1888a, 1893); and Arkadelphia and Okolona, Clark Co.,

and Muddy Fork, county unknown (Bollman 1893).

Pennsylvania — Pennsylvania in general (Underwood 1887); western

Pennsylvania (Wood 1862, 1865, Saussure and Humbert 1872, Kevan

1983).

Tennessee — Mossy Cr., Jefferson Co. (Bollman 1888c); Gatlinburg

and Great Smoky Mountains National Park, Sevier Co. (Chamberlin

19446).

North Carolina — North Carolina in general (Brolemann 1896,

Kraepelin 1903, Attems 1930); west of the central Piedmont Plateau,

specific counties shown in Fig. 13 (Shelley 1987); Mt. Pisgah, Haywood/

Buncombe cos. (Wray 1950, 1967).

Georgia— Lula, Hall Co. (Chamberlin 1944a).

Theatops calif orniensis

General range statement — along the western slope of the Sierra

Nevada and Cascade Mountains from Tulare County, California, to

Douglas County, Oregon, extending to San Francisco Bay and the Pacific

Ocean from Marin to Mendocino cos., California (Shelley 1990a).

California — California in general (Kraepelin 1903, Attems 1930,

Crabill 1960); Quincy, Plumas Co. (Chamberlin 1902, Shelley 1990a).

Oregon — Oregon in general (Kraepelin 1903, Chamberlin 1911,

Attems 1930, Crabill 1960, Kevan 1983).

Theatops phanus

Texas — Texas in general (Crabill 1960); Sonora, Sutton Co. (Cham-

berlin 19516, Reddell 1965, Shelley 1990a); Powell's Cave, Menard

Co. (Reddell 1965, Shelley 1990a).

Theatops erythrocephalus

Croatia — Dalmatia (= littoral Croatia) (Kraepelin 1903, Attems

1930). Istrian peninsula in general (Attems 1930), Pula (Koch 1847,

1863, Kohlrausch 1881, Attems 1929). Sipan Island (Shelley 1990a).

Hungary — Hungary in general (Kraepelin 1930); south Hungary

(Attems 1930). In modern Europe these reports probably refer to parts

of Croatia.

Montenegro — Montenegro in general (Attems 1930).

Spain — Cueva del Cerro de la Pileta (Matic 1960).

Portugal — Portugal in general (Verhoeff 1896, Kraepelin 1903,

Attems 1930).

Plutonium zwierleini

66 Rowland M. Shelley

Sicily — Sicily in general (Cavanna 1881, Kraepelin 1903, Wiirmli

1975); Ficuzza and Taormina (Attems 1930).

Sardinia — Sardinia in general (Kraepelin 1903, Attems 1930, Wiirmli

1975).

Italy — South Italy, Cave of Tirreni and Calabria (Attems 1930);

Campania Region (Wiirmli 1975, Lewis 1981).

TAXONOMIC CHARACTERS

Aside from the segmental spiracles that characterize Plutonium,

the taxonomic features in the Plutoniuminae are located primarily on

the ultimate legs and segment. The dorsal and ventral surfaces are

both important in distinguishing species of Theatops, and the head

and first 20 segments hold little taxonomic utility. The massive caudal

legs instantly identify the centipede as a plutoniuminine, and if these

are lost, Theatops can be confused with the species of Scolopocryptops

Newport that exhibit complete paramedian dorsal sutures. In Theatops,

tergites 2-20, or every tergite except the first and last, exhibit two

such grooves, which lie on both sides of the midline, run the complete

length of the plate, and divide it into nearly equal thirds. Three species

of Scolopocryptops also demonstrate complete paramedian sutures on

all except the anteriormost tergites — S. gracilis Wood in California,

occurring sympatrically with T. californiensis; S. rubiginosus L. Koch

in the midwest, occurring sympatrically with T. spinicaudus; and S.

peregrinator (Crabill) in the northern Appalachians, occurring sympatrically

with T. posticus. In these areas, care is needed to avoid misidentifying

a cryptopid lacking the caudal legs, or the legs and caudal segment,

and confusing Theatops and Scolopocryptops. If such a specimen is

found, one should rely on secondary features to distinguish the genera,

as listed in Table 1.

Ultimate segment. Dorsal surface. The ultimate/caudal tergite

is noticeably longer than the preceding ones and lacks paramedian

sutures. It either lacks sutures altogether, as in most specimens of

T. spinicaudus (Figs. 7-9) (occasional specimens have a short sutural

remnant anteriad), or has a complete or nearly complete one in the

midline, as in the other species (Fig. 6). The suture is complete in

T. posticus and T. phanus, running the entire length of the tergite

from the anterior to caudal margins; it is rarely complete in T. californiensis

and T. erythrocephalus, as it usually disappears just short of the caudal

margin. Thus, in the absence of the caudal legs, the presence or absence

of the median suture on the ultimate tergite distinguishes T spinicaudus

and T. posticus in their areas of overlap in the eastern United States.

The latter species was incorrectly illustrated without this suture by Shelley

Centipede Subfamily Plutoniuminae

6 7

Table 1. Comparisons of Theatops and species of Scolopocryptops

with complete paramedian sutures.

Scolopocryptops

Theatops

Ultimate legs

Ultimate tergite

Caudal coxopleurae

Number of segments

and leg pairs

Color

Cephalic plate

Anteriormost tergites

subequal in breadth and

degree of sclerotization

to other legs, not

noticeably enlarged

subequal in length and

with same suture pattern

as other tergites

not extended and without

spurs

23

usually orange, reddish-

orange, or brown,

excepting yellowish

variant of S.

peregrinator

depigmented "eyespots"

absent

lst-3rd tergites with

incomplete paramedian

sutures except in S.

gracilis

heavily sclerotized and

greatly enlarged, much

larger than other legs

noticeably longer than

other tergites, either

without sutures or with

single one in midline

extended and/or with

distal spur except in

eastern population of

T. posticus

21

yellowish

pale, depigmented "eye-

spots" present

complete paramedian

sutures begin on 2nd

tergite

(1987, Fig. 3) and Shelley and Edwards (1987, Fig. 8); the errors were

corrected by Shelley (1990a, Fig. 3).

Ventral surface. The caudal coxopleurae, flat, not extended, and

apically rounded in the eastern population of T. posticus (Shelley 1990a,

Fig. 6), are modified to some degree in the other species and most

individuals in the southwestern population of T. posticus. Conditions

vary widely, but the structures are usually elevated, produced slightly

beyond the caudal segmental margin, and typically have a variable apical

Rowland M. Shelley

spur or a black dot; in the residual southwestern intergrades, the structures

are elevated and often produced, but lack an apical spur or black dot

(Shelley 1990a, Figs. 7-11). The strongest, most pronounced coxopleurae

are in T. californiensis and T. erythrocephalus (Figs. 13-17, 40-41).

In the absence of the caudal legs, this coxopleural feature also distinguishes

T. spinicaudus from T. posticus in their areas of sympatry.

Ultimate legs. The robust, heavily sclerotized, forcipulate caudal

legs are the most obvious diagnostic trait of the Plutoniuminae and

readily identify Theatops in the United States. Occasionally distended

(Figs. 7, 19), the legs typically extend directly caudad basally then

curve towards each other and converge such that the tips either meet

or cross. They are believed to hold prey for feeding, and this can

probably occur either apically, as the sharply pointed tips could puncture

most prey organisms, or basally, with the prey being squeezed or pinched

by the legs. The caudal legs are more heavily sclerotized than the

tergites or the other appendages, but the inner or medial surfaces in

many individuals are particularly hard, much more so than the outer

surfaces. Perhaps the caudal legs serve to kill prey by crushing or

puncturing as well as to hold it.

The outer surfaces of these legs (dorsal, ventral, and lateral)

are usually gently curved to flattened, but the medial surfaces are

often strongly flattened, such that there are sharp ventro- and dorsomedial

corners that are often elevated above the ventral and dorsal surfaces,

forming a ridge. The ridge sometimes points mediad rather than ventrad

or dorsad, such that the medial surface is slightly recessed, in some

individuals being slightly concave rather than flat. These ridges or

corners may be linear or irregularly and lightly scalloped, and are

often ornamented with variable teeth or spurs. Ventrally, the number,

size, and arrangement of the teeth vary greatly in T. spinicaudus, T.

phanus, and the southwestern population of T. posticus (Figs. 19-30,

35-38; Shelley 1990a, Figs. 7-11), whereas there is usually a single

distal tooth in T. californiensis and T. erythrocephalus (Figs. 13-17,

40-41). Dorsally, there is less ornamentation, but occasional individuals

show a serrated margin with several fine teeth (Figs. 7-9, 33-34).

The distomedial prefemoral spurs in T. spinicaudus and T. phanus

are really at the distal corner of this ridge; consequently, the dorsal

surfaces of the legs hold taxonomic utility in distinguishing these species

from T. posticus, particularly in areas where T. posticus and T. spinicaudus

are sympatric. Although differing in the presence (T. posticus) and

absence (T. spinicaudus) of the median suture on the ultimate tergite,

they are most readily distinguished by the presence or absence of the

distomedial prefemoral spur. Variation on the ventral surface holds little

Centipede Subfamily Plutoniuminae

6 9

Figs. 6-9. Caudal tergite and legs of T. posticus and T. spinicaudus, dorsal

views. 6, T. posticus from Jefferson County, Florida. 7-9, T. spinicaudus

from selected localities. 7, McDowell County, North Carolina. 8, Polk County,

Arkansas. 9, Haywood County, North Carolina. Scale lines = 1.00 mm for

each figure.

70 Rowland M. Shelley

taxonomic utility, but the presence of, usually, four large distal spurs,

one on each prefemur and femur, distinguishes T. californiensis from

T. posticus in California; the spurs are smaller, more on the order of

teeth, and there are consistently less than four total in the southwestern

population of the latter. To be certain of determinations of California

material, one should also examine the caudal coxopleurae, which almost

always have a sizable terminal spur in T. californiensis (Figs. 13-17)

and no adornment in T. posticus (Shelley 1990a, Figs. 7-11).

TAXONOMY

Order Scolopendromorpha

Key to Families

With four ocelli on each side of cephalic plate lateral to bases

of antennae Scolopendridae Leach

Without ocelli Cryptopidae Kohlrausch

Family Cryptopidae Kohlrausch

Diagnosis (adapted from Hoffman (1982)). Scolopendromorpha

with 21 or 23 leg pairs and pedal segments; ocelli absent; tarsi of

leg pairs 1-19 single-segmented; sternites usually with median and

transverse grooves, without paramedian grooves; intercalary tergites

usually strongly developed.

Key to Subfamilies

1. With 23 leg pairs and pedal segments Scolopocryptopinae

Pocock

With 21 leg pairs and pedal segments 2

2. Ultimate legs strongly incrassate and forcipulate; ultimate tergite

much longer than preceding; cephalic plate with conspicuous,

lightly pigmented patches in ocellar positions lateral to bases

of antennae Plutoniuminae Bollman

Ultimate legs at most only slightly crassate, relatively long and

slender, not forcipulate; ultimate tergite at most only slightly

longer than preceding; cephalic plate uniformly pigmented,

without lighter patches in ocellar positions Cryptopinae

Kohlrausch

Subfamily Plutoniuminae Bollman (Nomen Correctum)

Plutoniinae Bollman, 1893:165, 168.

Theatopsidae Verhoeff, 1906:432; 1907:253. Ribaut, 1915:338. Attems,

1930:249-250.

Plutoniidae: Verhoeff, 1906:433.

Centipede Subfamily Plutoniuminae 71

Theatopsini: Attems, 1926:376.

Theatopinae: Biicherl, 1942:325. Matic, 1960:446. Crabill, 1960:9. Summers

1979:696. Summers, Beatty, and Magnuson, 1980:245; 1981:59.

Kevan, 1983:2945. Shelley, 1987:505.

Theatopsinae: Lewis, 1981:427. Schileyko, 1992:13.

Components. Theatops Newport, 1844, and Plutonium Cavanna,

1881. Tonkinodentus Schileyko, 1992, assigned to the Theatopinae in

its original proposal, is removed from subfamily and left unassigned.

Diagnosis (adapted from Attems 1926). A subfamily of moderate-

size to large cryptopids with 21 pairs of legs and pedal segments,

color generally yellowish, usually with lighter, pale or unpigmented,

patches lateral to bases of antennae in ocellar positions; complete paramedian

dorsal sutures present on tergites 2-20 (see Attems 1926, Fig. 433,

p. 369); caudal tergite nearly twice as long as penultimate, with or

without median suture; ultimate legs short, massive, and forcipulate,

heavily sclerotized, basal podomeres nearly as wide as long, with or

without dorsal and ventral teeth and spurs.

Distribution. Southern Europe — coastal Croatia and Montenegro,

Bosnia-Hercegovina, southern Italy (Campania), Sardinia, Sicily, Spain,

and Portugal (Fig. 44); eastern North America generally east of the

Central Plains and south of the Great Lakes, from western New England

and the vicinity of Chicago, Illinois, to the south Florida Keys and

central, southern, and western Texas; and western North America from

western Chihuahua and northern Baja California Norte, Mexico, to

the southern Basin and Range Province in southern Nevada and south-

western Utah, westward to the Pacific Ocean and southwestern Oregon

(Fig. 10).

Genus Theatops Newport

Theatops Newport, 1844:409; 1845:60; 1856:60. Wood, 1862:36; 1865:171.

Saussure and Humbert, 1872:200. Kohlrausch, 1881:93. McNeill,

1888:16. Pocock, 1888:285, 287; 1895:28. Bollman, 1893:127,

142, 169-170. Brolemann, 1896:50; 1904:244. Kraepelin, 1903:64-

65. Verhoeff, 1906:385; 1907:253. Chamberlin, 1911:472. Attems,

1930:250-251. Biicherl, 1942:325-326. Crabill, 1957:345. Matic,

1960:446. Kevan, 1983:2945. Foddai et al., 1995:8.

Opisthemega Wood, 1862:35; 1865:169. Latzel, 1880:145-147. Kohlrausch,

1881:130. Meinert, 1886:207-208. Underwood, 1887:64-65. Daday,

1889:92.

72

Rowland M. Shelley

Fig. 10. Approximate probable distributions of Theatops and the Plutoniuminae

in the Western Hemisphere. A smooth line is drawn around range extremes

in all directions.

Opisthomega: Saussure and Humbert, 1872:200. Haase, 1887:78.

Type species. Of Theatops, Cryptops postica Say, 1821, by subsequent

monotypy of Newport (1845). As explained by Crabill (1957), Newport

(1844) originally proposed Theatops without mention of included species,

but the next year, he (Newport 1845) cited C. postica under it, thereby

fixing this as the type species. Opisthemega was proposed for two

ostensibly new species, O. postica and O. spinicauda. Wood (1862)

did not designate either as the type species, nor did he (Wood 1865)

do so in his work on North American myriapods. No one else has

done so either, and this action would be redundant now with Opisthemega

in synonymy under Theatops. According to Article 12(b)(5) of the

International Code of Zoological Nomenclature, genus-group names

published before 1931 are available if one or more available species-

group names are published in combination with it, or clearly included

under it, or clearly referred to it by bibliographic reference. As both

postica and spinicauda are valid species-group names, published in

accordance with the Code, Opisthemega meets the requirements of

Centipede Subfamily Plutoniuminae 73

this article and is an available genus-group name even though it lacks

a type species. It is one of the rare genus-group names lacking a

genotype, the only one I know of in myriapods.

Diagnosis. With nine pairs of spiracles, on segments 3, 5, 8,

10, 12, 14, 16, 18, and 20.

Distribution. Coastal Croatia and Montenegro, Bosnia-Hercegovina,

Spain, and Portugal; eastern North America from western New England

and the vicinity of Chicago, Illinois, to the south Florida Keys and

nearly to Corpus Christi, Texas, ranging westward to the eastern border

of the Central Plains in Oklahoma and onto the Edwards Plateau in

Texas; and western North America from western Chihuahua and southern

Sonora, Mexico, to the Pacific Ocean in California and northern Baja

California Norte, north to southern Nevada, southwestern Utah, and

southwestern Oregon (Fig. 10).

Species. Five, as distinguished in the following key, adapted from

those in Attems (1930), Chamberlin (1951/?), and Weaver (1982).

Remarks. Sutural differences on the ultimate tergite deserve emphasis.

Every specimen of T. posticus and T. phanus that I examined displayed

a complete median suture running the entire length of the tergite from

the anterior to the posterior edges; conversely nearly every individual

of T. spinicaudus lacked the suture, but occasional specimens have

minute, barely detectable remnants from the anterior margins. For practical

purposes, T. spinicaudus can be characterized as lacking the suture

because these occasional remnants extend no more than 1/32-1/16 of

the tergal length. The other species, T. californiensis and T. erythro-

cephalus, typically possess incomplete median sutures that extend caudad

from the anterior margin but terminate or fade out just before the

caudal edge. Rarely in these species does the suture extend the entire

length of the tergite.

Key to Species of Theatops

1. European species, occurring in Portugal, Spain, Croatia, Bosnia-

Hercegovina and Montenegro ...erythrocephalus (C. L. Koch)

North American species 2

2. Ultimate prefemora with dorsal distomedial spurs (Figs. 7-9) ... 3

Ultimate prefemora without dorsal spurs (Fig. 6) 4

3. Ultimate tergite with complete longitudinal, midline suture dorsally

(Figs. 33-34); central and western Texas phanus

Chamberlin

Ultimate tergite without midline suture or with only minute remnant

anteriad (Figs. 7-9); eastern United States from northern Illinois

and central Iowa to southwestern Arkansas, and from northwestern

74 Rowland M. Shelley

North Carolina, northeastern Tennessee and adjacent periphery

of Virginia to central North and South Carolina, and eastcentral

and northwestern Alabama spinicaudus (Wood)

4. Caudal coxopleurae distinctly elongate and apically acuminate,

usually with blackened terminal spurs; ventral surfaces of ultimate

prefemora and femora usually with one spur each (four spurs

total) distal to midlength (Figs. 13-17); Kern County, California,

to Douglas County, Oregon californiensis Chamberlin

Caudal coxopleurae at most only slightly prolonged, apically rounded,

without blackened spurs; ventral surfaces of ultimate prefemora

and femora with or without spurs, rarely with one on each

podomere, total almost always less than four (Shelley 1990a,

Figs. 6-11); eastern United States from eastern Connecticut

to eastern and southern coastal Texas, eastcentral Oklahoma,

and south Florida Keys, and western United States and northwestern

Mexico from western Chihuahua, southern Sonora, and northern

Baja California Norte to southwestern Utah, southern Nevada,

and southern California posticus (Say)

Theatops posticus (Say)

Figs. 1, 6, 11-12, 18, 32

Cryptops postica Say, 1821:112-113. Newport, 1844:100. Kohlrausch,

1881:130.

Cryptops posticus: Lucas, 1840:547. Bollman, 1893:147-148.

Theatops postica: Newport, 1845:410. Wood, 1862:36-37; 1865:171.

Saussure and Humbert, 1872:200. Kohlrausch, 1881:93. Attems,

1930:251. Crabill, 1955:259. Branson and Batch, 1967:83-84.

Kevan, 1983:2945.

Cryptops prolonge Gervais, 1847:294.

Opisthemega postica: Wood, 1862:35; 1865:169-170, pi. I., fig. 4.

Cope, 1869:179. Kohlrausch, 1881:130.

Opisthomega postica: Saussure and Humbert, 1872:200.

Opisthemega crassipes Meinert, 1886:209. McNeill, 1887:326; 1888:16.

Theatops crassipes: Bollman, 18886:110.

Theatops posticus: Bollman, 1888c:342; 1888J:346; 1888e:408; 1893:170.

Brolemann, 1896:50; 1904:244. Chamberlin, 1902:41; 1918^:23;

19186:375; 1925:57; 1942:184-185; 1943:97; 1944a:33; 19446:178;

1945:215; 1951a:33. Kraepelin, 1903:65-66, fig. 25. Williams

and Hefner, 1928:137. Brimley, 1938:501, in part. Crabill,

1950:201. Wray, 1950:156, in part; 1967:156, in part. Summers

et al., 1980:245; 1981:59. Shelley, 1987:505, figs. 3, 13. Shelley

and Edwards, 1987:Fig. 8.

Centipede Subfamily Plutoniuminae 75

Type specimen. Holotype (NHM) collected by T. Say on an unknown

date in the winter of 1818, possibly near Picolata, St. Johns County,

Florida. According to Weiss and Ziegler (1931), Say and friends visited

Florida in the winter of 1818, traveling overland by carriage to Charleston,

then by boat to Savannah, then by smaller boat through the "sea islands"

of Georgia. After stopping at Fernandina, on Amelia Island, Florida,

they proceeded up the St. Johns River to Picolata, where they disembarked

and crossed by foot to St. Augustine to present their papers to the

Spanish governor. Because of hostile indians, he advised against traveling

farther upriver, so they returned to Picolata, sailed back to the coast

and, eventually, Charleston. The type of C. postica was collected on

this trip somewhere in Georgia or Florida, and although the party

stopped repeatedly on the islands of Georgia, including a few days

on Cumberland Island, they were ashore for the longest time in the

area of Picolata/St. Augustine, the most likely site for the collection.

Picolata exists today as a small community on the river along St.

Johns County Highway 13, ca. 25.3 km (15.8 mi) west of St. Augustine.

According to Underwood (1887) and Pocock (1888), Say sent some

or all of the type specimens of his myriapods to Leach in Britain,

who deposited them in the NHM, and according to Newport (1845)

and Pocock (1888), there was only one specimen of C. postica in

this shipment, making it the holotype.

Diagnosis. Ultimate tergite with complete, median suture; ultimate

legs dorsally without distomedial prefemoral spurs (Fig. 6); ultimate

prefemora and femora with or without relatively short, "weak," ventral

spurs, when present, usually less than four spurs total or one per podomere;

caudal coxopleurae with borders apically rounded, at most only slightly

elevated and extended, without apical teeth (Fig. 2; Shelley 1990a,

Figs. 5-12).

Variation. The eastern population lacks ventral spurs on the caudal

legs, and the coxopleurae are flat and not extended caudad. Variation

of the coxopleurae and the ventral surfaces of the ultimate legs in

the southwestern population is discussed by Shelley (1990a).

Ecology. In the east, I have found T. posticus primarily in moist

deciduous litter, occasionally in pine litter, and rarely under rocks

and logs (Shelley 1987, 1990a); specimens may also be encoundered

under large rocks in rather dry sites (Hoffman, in lift.). The southwestern

population, which inhabits deserts and arid biotopes, has been encountered

under rocks, logs, wood debris, and cattle dung (Shelley 1990a). Even

the Stanislaus County, California, locality is arid, for it is on the eastern

slope of the Coast Range and in the rain shadow of these mountains.

76

Rowland M. Shelley

Distribution. Theatops posticus consists of allopatric populations

in eastern and southwestern North America segregated by around 1200

km (750 miles) (Shelley 1990a, Fig. 3). The eastern population extends,

north-south, from eastern Connecticut, the Catskill Mountains of New

York, southwestern Pennsylvania, and southern Illinois to the south

Florida keys and San Patricio County, Texas, and, east-west, from

the Atlantic Ocean to the eastern periphery of the Central Plains in

Seminole County, Oklahoma, and Limestone County, Texas. It encompasses

all of West Virginia, Kentucky, Tennessee, South Carolina, Georgia,

Florida, Alabama, Mississippi, Arkansas, and Louisiana. The southwestern

population, centered in the Sonoran Desert, occurs from western Chihuahua,

Mexico, to the Channel Islands in the Pacific Ocean off the southern

California coast, and from the eastern slope of the Coast Range in

Fig. 11. Distribution of T. posticus in eastern North America.

Centipede Subfamily Plutoniuminae 77

Stanislaus County, California, and the southern Great Basin in southern

Nevada and southwestern Utah to southern Sonora and northern Baja

California Norte, Mexico. The eastern area represents a slight modification

of that showed previously (Shelley 1990a, Fig. 4), which did not include

Connecticut and New York (Figs. 11-12), and the Stanislaus County

record extends the range significantly northward in western California.

In the following locality listing, general range statements, cited

from the literature where appropriate, are presented for each state before

detailed data. Counties of occurrence of the eastern population are

listed alphabetically for states where the centipede has been taken from

five or more counties; full locality data are presented for states where

less than five counties are represented. The most peripheral record(s)

are cited in detail for states forming range boundaries, even when

the centipede is common and only counties are listed. Locality data

for the southwestern population presented in Shelley (1990a) are also

summarized by county; new sites for this population obtained since

that paper was published are detailed.

EASTERN POPULATION

CONNECTICUT: Expected in the western 2/3 of the state, but

only one record. Tolland Co., Mansfield, 1 spmn., June 1965, collector

unknown (UCT).

NEW YORK: Expected in the southernmost section, south of Catskills,

but only one record; may be absent from Long Island. Sullivan Co.,

4.8 km (3 mi) N Bruce, 1 spmn., 22 May 1968, S. B. Peck (NCSM).

NEW JERSEY: Expected in the northern 1/3 of state, but no

available records.

PENNSYLVANIA: Expected in the east and south, but only, one

definite record. Allegheny Co., Sewickly, 3 spmns., date unknown,

W. L. Walker (NMNH).

INDIANA: Apparently common in the southern third with one

northern record, from the northeastern corner, that is somewhat disjunct

and needs confirmation. Brown, Crawford, Franklin, Jefferson, Monroe,

and Noble counties (ANSP, NMNH). Northernmost record: Noble Co.,

Indian Village Lake, 1 spmn., date unknown, B. G. Owens (NMNH).

ILLINOIS: Known only from the southern periphery; the northernmost

record is from Pulaski County (Summers et al. 1980). Gallatin Co.,

Shawneetown, 2 spmns., 23 June 1950, M. W. Sanderson (INHS). Hardin

Co., Cave in Rock, 1 spmn., 2 May 1956, L. J. Stannard (INHS). Pope

Co., Eddyville, 1 spmn., 1 May 1953, L. J. Stannard (INHS); and 3.2

km (2 mi) N Dixon Springs, 1 spmn., 1 May 1969, collector unknown

(EIU). Johnson Co., Bellsmith Springs E of Ozark, 1 spmn., 24 June

Rowland M. Shelley

1958, H. S. Dybas (FMNH).

MARYLAND: Expected west of Chesapeake Bay, but only one

record. Allegany Co., Cumberland, 1 spmn., 29 March 1966, collector

unknown (NCSM).

WEST VIRGINIA: Statewide. Berkeley, Dickenson, Greenbrier,

Mercer, Raleigh, and Summers counties (AAW, NCSM, WAS, WVDA).

VIRGINIA: Expected throughout most of the state except for the

eastern shore (Accomack and Northampton counties) and possibly the

southeastern corner of the mainland, around Norfolk. Albemarle, Alle-

ghany, Appomattox, Augusta, Bedford, Botetourt, Buchanan, Dickenson,

Fauquier, Floyd, Frederick, Giles, Greenville, Henry, Lee, Madison,

Montgomery, Page, Patrick, Pittsylvania, Pulaski, Rappahannock, Rock-

bridge, Rockingham, and Surry counties (AMNH, FSCA, LEM, MCZ,

NCSM, NMNH, VMNH). Easternmost record: Surry Co.,Chipokes Plantation

St. Pk., 2 spmns., 12 September 1988, R. M. Shelley (NCSM).

KENTUCKY: Statewide. Bell, Caldwell, Clark, Edmonson, Estill,

Grant, Grayson, Hardin, Harlan, Jefferson, Jessamine, Oldham, Shelby,

Todd, and Wolfe counties (FMNH, INHS, MPM, NCSM, NMNH, TMM,

UL, ZMUC).

TENNESSEE: Statewide. Anderson, Bledsoe, Davidson, Franklin,

Hamilton, Jefferson, Knox, Lake, Madison, Marshall, Morgan, Overton,

Roane, and Sevier counties (AMNH, FMNH, FSCA, INHS, MCZ, NCSM,

NMNH, TMM).

NORTH CAROLINA: Statewide except for the Outer Banks and

the eastern extremity of the mainland; more prominent east of the

Blue Ridge Province (Shelley 1987). Bladen, Burke, Chatham, Cherokee,

Clay, Cumberland, Davidson, Durham, Franklin, Gaston, Harnett, Iredell,

Jackson, Johnston, Lee, Madison, Pitt, Richmond, Stanly, Stokes, Surry,

Wake, Wayne, and Wilkes counties (FSCA, MCZ, NCSM, NMNH).

Easternmost record: Pitt Co., 2.1 km (1.3 mi) W Greenville, along

NC hwy. 43, 0.2 km (0.1 mi) W jet. NC hwy 903, 1 spmn., 19 October

1979, R. M. Shelley, P. T. Hertl (NCSM).

SOUTH CAROLINA: Expected statewide, but known from only

four counties. Chester Co., 18.6 km (11.6 mi) W Chester, 1 spmn.,

30 April 1977, R. M. Shelley (NCSM). Greenville Co., Greenville,

1 spmn., 29 July 1961, S. & D. Mulaik (NMNH). Aiken Co., Savannah

River Plant, Sunshine Bay, 16 April 1969, collector unknown (SREL).

Jasper Co., Ridgeland, 1 spmn., 6 April 1975, D. Brady (AMNH);

and 12.8 km (8 mi) S Hardeeville, along US hwy. 17A, 0.6 km (0.4

mi) W jet. SC hwy. 170, 1 spmn., 9 November 1977, R. M. Shelley

(NCSM).

GEORGIA: Statewide. Bacon, Berrien, Bibb, Camden, Charlton,

Centipede Subfamily Plutoniuminae 79

Chatham, Clarke, Decatur, Habersham, Jackson, Jefferson, Jenkins, Lanier,

Mcintosh, Polk, Rabun, Screven, Thomas, and Ware counties (FMNH,

FSCA, MCZ, NCSM, NMNH, UGA).

FLORIDA: Statewide. Alachua, Baker, Bay, Charlotte, Clay, Collier,

Columbia, Dade, Duval, Escambia, Gadsden, Glades, Hamilton, Hernando,

Highlands, Hillsborough, Jackson, Jefferson, Lake, Leon, Liberty, Madison,

Manatee, Marion, Martin, Monroe, Nassau, Pinellas, Polk, Putnam,

Santa Rosa, and Sarasota counties (AMNH, EIU, FMNH, FSCA, INHS,

MCZ, NCSM, NMNH). Southernmost record: Monroe Co., Sugarloaf

Key, 1 spmn., March 1898, O. F. Cook (NMNH).

ALABAMA: Statewide. Baldwin, Butler, Chilton, Choctaw, Clarke,

Cullman, Dallas, DeKalb, Lee, Marengo, Marshall, Marion, Mobile,

Morgan, Wilcox, and Winston counties (AU, FMNH, FSCA, NCSM,

NMNH).

MISSISSIPPI: Statewide. Alcorn, Forrest, Hancock, Harrison, Jackson,

Jones, Lafayette, Lamar, Lee, Noxubee, Oktibbeha, Pontotoc, Prentiss,

Rankin, Scott, Smith, Stone, Tishomingo, Wayne, Webster, and Winston

counties (FMNH, FSCA, INHS, MCZ, MEM, NMNH).

LOUISIANA: Statewide. East Baton Rouge, Evangeline, Grant,

Jefferson, Lincoln, Natchitoches, Orleans, Ouachita, Rapides, St. Tam-

many, Washington, West Baton Rouge, and Winn parishes (FMNH,

FSCA, MCZ, NMNH).

ARKANSAS: Expected statewide, but known from only three counties.

Baxter Co., Lake Norfolk, 1 spmn., 2 August 1952, N. B. Causey

(NMNH). Washington Co., locality unknown, 1 spmn., October 1958,

G. Ogden (NMNH). Columbia Co., Magnolia, 2 spmns., 24 December

1949, N. B. Causey (NMNH).

OKLAHOMA: Expected in the eastern 1/3 of the state, but only

one record. Seminole Co., locality unknown, 5 spmns., May 1931,

P. Newport (NMNH).

TEXAS: Expected throughout the forested, eastern 1/4 of the

state, generally east of interstate highway 45, extending southward

along the coast nearly to Corpus Christi, Nueces County. Angelina,

Brazos, Chambers, Galveston, Harris, Hunt, Jasper, Limestone, Nacog-

doches, Sabine, San Patricio, Tyler, Van Zandt, and Walker counties

(AMNH, CAS, FSCA, NMNH). Westernmost records: Limestone Co.,

Mexia, 1 spmn., 3 December 1961, B. E. Oberholtzer (NMNH) and

Brazos Co., 8 km (5 mi) S College Station, 3 spmns., 21 April 1936,

L. Hubricht (NMNH). Southernmost record: San Patricio Co., 11.2

km (7 mi) N Sinton, Welder Wildlife Refuge, 1 spmn., 5 July 1965,

R. O. Albert (FSCA).

The following literature records of the eastern population are

80 Rowland M. Shelley

considered valid and are incorporated into fig. 11.

OHIO: Gallia Co., Vinton (Morse 1902); southeastern Ohio in

general (Williams and Hefner 1928). Morse's is the only definite Ohio

record, but T. posticus probably occurs widely in the southern half

of the state.

ILLINOIS: Jackson and Pulaski cos. (Summers et al., 1980, 1981).

INDIANA: Wayne Co., Dublin (McNeill 1888); Clark Co. New

Providence (Bollman 1888e).

KENTUCKY: Jefferson Co., Louisville, and Whitley Co., Cumberland

Falls St. Pk. (Crabill 1955a); and Powell Co., Middle Fork of Red

R. near Slade (Branson and Batch 1967).

NORTH CAROLINA: Guilford Co., Greensboro (Causey 1940);

and Wayne Co., Goldsboro (Wood 1862, Bollman 1888J, Brimley 1938,

Wray 1950, 1967)).

SOUTH CAROLINA: Pickens Co., Clemson (Crabill 1950).

GEORGIA: Mitchell Co., Camilla (Chamberlin 1945).

LOUISIANA: St. Helena Par., Greenburg (Chamberlin 1942).

SOUTHWESTERN POPULATION

MEXICO:

BAJA CALIFORNIA NORTE, SONORA, and CHIHUAHUA (AMNH,

NCSM, NMNH, UCB). Expected in the western periphery of Chihuahua,

all but the southern tip of Sonora, and an equivalent distance down

the Baja peninsula, including all of Baja California (B. C.) Norte

and the northern extremity of B. C. Sur. It is currently unknown from

the last state and the southern half of B. C. Norte. New Record: SONORA:

6.4 km (4 mi) SW Los Vidrios, W of Sonoita, 2 spmns., 11 February

1960, V. Roth (AMNH).

USA:

NEW MEXICO: Expected only in the southwestern corner. Hidalgo

County (NMNH).

ARIZONA: Expected throughout all but the northeastern 1/4 of

the state, or most of Navajo and Apache counties. Cochise, Coconino,

Gila, Graham, Maricopa, Pima, Pinal, Santa Cruz, Yavapai, and Yuma

counties (AMNH, CAS, FSCA, NMNH, OPCNM, SWRS, UAZ). New

Records: Coconino Co., Schnebly Hill Vista, 1 spmn., 7 October 1987,

B. Hebert (LACMNH). Pima Co., Elkhorn Ranch, E slope of Baboquivari

Mts., 1 spmn., H. B. Leech, J. W. Green (CAS). Navajo Co., nr.

Showlow, 1 spmn., 10 August 1948, G. E. Ball, H. E. Evans (NMNH).

Santa Cruz Co., Yane Springs, Sycamore Cyn., 1 spmn., 21 March 1967,

V. F. Lee, T. S. Briggs (CAS); and Blanca L. nr Pena, 1 spmn., 21

Centipede Subfamily Plutoniuminae

81

V

•7>

12

\

Fig. 12. Distribution of T. posticus in Mexico and the southwestern United

States.

March 1967, K. Horn, P. S. Sum (CAS).

UTAH: Expected only in the southwestern corner. Washington County

(NMNH). New Record: Washington Co., Warner Valley, 1 spmn., 5

April 1975, A. H. Barnum (DC).

NEVADA: Expected only in the southern corner. Clark and Nye

counties (FSCA, NMNH).

CALIFORNIA: Expected in desert areas east of the Sierra Nevada

82

Rowland M. Shelley

Figs. 13-17. Variation of the ventral surfaces of the caudal legs and coxopleurae

of T. californiensis from selected localities. 13, Mariposa County, Califor-

nia. 14, Butte County, California. 15, Tuolumne County, California. 16, El

Dorado County, California. 17, Douglas County, Oregon. Scale lines = 1.00

mm for each figure.

Centipede Subfamily Plutoniuminae 83

and southward to the Mexican border, possibly extending along the

coast to Ventura County. Riverside County and Santa Cruz Island, Channel

Islands National Park (UCR). New Records: Inyo Co., 4.8 km (3 mi)

E Big Pine, Saline Valley Rd., 1 spmn., 11 June 1967, W. J. Ball,

H. E. Evans (NMNH); Saline Valley, Grapevine Rd. Sta. 32, 1 spmn.,

7 May 1960, B. Banta (CAS); and 40 km (25 mi) S Saline Valley, 1

spmn, 29 April 1975, A. R. Hardy (CDFA). Riverside Co., Whitewater

Cyn., 1 spmn., 15 February 1959, I. Newell (AMNH); and Palm Springs,

nr. Taquitz Cyn., 4 spmns., 23 March 1965, D. Yang (CAS). Stanislaus

Co., Del Puerto Cyn., ca. 28.8 km (18 mi) W Patterson, 2 spmns., 10-

11 April 1990, E. I. Schlinger (UCB).

Remarks. Theatops posticus was collected at Appomattox Court

House, Virginia, on the very day of the surrender there of Gen. Robert

E. Lee's Confederate army, 9 April 1865, by an unknown member

of the surrender parties (Shelley 19906).

Apparently, T. posticus and T. spinicaudus can occur syntopically,

because some preserved samples, ostensibly collected at one place on

one date, contain both species.

Theatops californiensis Chamberlin

Figs. 13-18

Theatops californiensis Chamberlin, 1902:41. Kevan, 1983:2945.

Theatops erythrocephalus (nee C. L. Koch): Kraepelin, 1903:66-67,

Fig. 26.

Theatops erythrocephalus californiensis: Chamberlin, 1911:472.

Theatops erythrocephala (nee C. L. Koch): Attems, 1930:251-252, Figs.

331-335. Kevan, 1983:2945.

Type specimens. Three syntypes (NMNH) collected by E. Garner

in the summer of 1901 at Quincy, Plumas County, California.

Diagnosis. Ultimate tergite usually with incomplete, median suture,

running from anterior margin to just short of caudal edge; ultimate

legs without dorsal distomedial prefemoral spurs; ultimate prefemora

and femora usually with four strong, distinct ventral spurs, one on

each podomere, rarely with three or fewer spurs; caudal coxopleurae

with medial borders strongly elevated and extended caudad, usually

apically acuminate with blackened terminal spurs (Figs. 13-18).

Variation. Occasional exceptions exist to the typical pattern of

four ventral spurs, one on each caudal prefemur and femur (Fig. 13).

A specimen from Brush Creek, Butte County, California, exhibits three

spurs, the right prefemur lacking the structure (Fig. 14). An individual

from 19.2 km (12 mi) E Buck Meadows, Tuolumne County, has two

spurs, lacking those on the left prefemur and femur (Fig. 15), and

84

Rowland M. Shelley

Fig. 18. Distributions of T. calif orniensis (dots) and T. posticus (stars) in

California, Oregon, and Nevada.

Centipede Subfamily Plutoniuminae

one from 16.8 km (10.5 mi) SW Bucks Lake, Plumas County, lacks

spurs on the right leg, which is considerably smaller than the left

and apparently regenerating. No individuals are available with only

one spur, but four are devoid of the structures, one from Oroville,

Butte County, and three from Canyonville, Douglas County, Oregon

(Fig. 17). More rarely, a specimen will lack a coxopleural spur, as

for example an individual from El Dorado County, which lacks that

on the right coxopleura (Fig. 16).

Ecology. Labels with samples indicate that specimens were found

under logs and the bark of decaying logs or stumps. Most, however,

were encountered in litter, which was my experience during field work

in California. In June 1990 and April 1991, I found T. californiensis

to be abundant in litter in Yosemite Valley, Yosemite National Park,

Mariposa County, but I did not find a single specimen under a log.

Distribution. The only specific published locality is the type locality.

Several authors have reported this species from California and Oregon

in general (Kraepelin 1903, Attems 1930, Crabill 1960, Kevan 1983),

but Shelley (1990a, Fig. 4) first delineated regions — along the western

slope of the Sierra Nevada and southern Cascade Mountains from Tulare

County, California, to Douglas County, Oregon, extending to San Francisco

Bay and the Pacific Ocean from Marin to Mendocino counties. The

northern limit, in southern Douglas County, Oregon, is unchanged,

but I have examined more southerly material from northern Kern County,

and the southern limit is thus in this county, in the southern part

of the Sequoia National Forest and the Sierra Nevada Mountains. In

May 1993, I spent a day searching unsuccessfully for T. californiensis

in the Toiyabe National Forest, on the eastern side of Lake Tahoe

in Nevada. The centipede may eventually be found in this area, but

it is currently known only from the California side of the lake. The

Oregon localities may be disjunct and represent a small, allopatric,

northern population, as there are no records between Josephine County

and Mendocino County, California. Because the type locality is the

only specific recorded site, I list below all records of T. californiensis

(Fig. 18).

OREGON: Occurring only in the southwestern interior. Douglas

Co., Susan Cr. E of Glide, 1 spmn., 23 July 1962, V. Roth (AMNH);

and Canyonville, 3 spmns., 13 February 1946, S. & D. Mulaik (NMNH)

and 5 spmns., 12 July 1946, S. & D. Mulaik (NMNH). Josephine

Co., 14.4 km (9 mi) W Sunny Valley, 2 spmns., 22 July 1962, V.

Roth (AMNH).

CALIFORNIA: Widespread from the northwestern interior to the

southern Sierra Nevada, extending primarily along the western slope

86 Rowland M. Shelley

of this range and the Cascades, traversing the crest to the vicinity of

Lake Tahoe, and expanding westward to San Francisco Bay and northward

along the Pacific Ocean. Mendocino Co., 3.2 km (2 mi) W Piercy, 1

spmn., 19 August 1959, W. J. Gertsch, V. Roth (AMNH). Marin Co.,

Mill Valley, 1 spmn., 28 February 1954, H. B. Leech (CAS) and 1

spmn., 13 November 1958, H. B. Leech (CAS). Contra Costa Co., W

of Pittsburg, 1 spmn., 21 March 1957, J. Russell (UCB). Plumas Co.,

Quincy, 3 spmns., summer 1901, E. Garner (NMNH) and 2 spmns., 7

July 1946, S. & D. Mulaik (NMNH) TYPE LOCALITY; and 16.8 km

(10.5 mi) SW Bucks Lake, 1 spmn., 14 September 1983, M. E. Bugler

(UCB). Butte Co., 11.2 km (7 mi) E Chico, Bidwell Park, 1 spmn., 2

April 1965, H. B. Leech (CAS), 6.4 km (4 mi) SW Stirling City, Toadtown,

2 spmns., 11 April 1979, C. L. Hogue (LACMNH); Brush Cr., 2 spmns.,

30 May 1955, K. W. Haller (AMNH); Forest Ranch, 1 spmn., 27 April

1991, R. M. Shelley (NCSM); and Oroville, 1 spmn., 10 April 1911,

collector unknown (NMNH). Yuba Co., Campton-ville, 1 spmn., 7 September

1959, V. Roth (AMNH). Nevada Co., Grass Valley, 1 spmn., 12 September

1966, J. S. Buckett, M. R. Gardner (UCD); and 8 km (5 mi) S Washington,

2 spmns., 8 October 1967, V. F. Lee, K. Horn (CAS). Placer Co., E

end of Bear Valley, 7 spmns., 1 April and 1 June 1964, P. H. Arnaud

(CAS); Colfax, 2 spmns., June 1888, collector unknown (NMNH); Emigrant

Gap, 3 spmns., 16 July 1937, R. V. Chamberlin (NMNH); 4.8 km (3

mi) E Auburn, 1 spmn., 28 March 1941, S. & D. Mulaik (NMNH);

and 7.5 km (4.7 mi) W Foresthill, 11 spmns., 27 November 1965

and 25 April 1966, H. B. Leech (CAS). El Dorado Co., Lake Tahoe,

25 July 1915, 1 spmn., 25 July 1915, collector unknown (NMNH)

and 1 spmn., 11 July 1952, W. J. Gertsch (NMNH); Fallen Leaf Lake,

1 spmn., 9 September 1959, W. J. Gertsch (AMNH); Glen Alpine

Springs, 1 spmn., 28 June 1915, collector unknown (NMNH); Echo

Pass S Meyers, 1 spmn., 30 June 1955, M. Cazier (AMNH) and 1

spmn., 19 September 1963, W. J. Gertsch (AMNH); Echo Lake, 1

spmn., July 1934, L. W. Saylor (NMNH); Kyburz, 1 spmn., 29 June

1977, C. E. Griswold (UCB); 6.4 km (4 mi) W Kyburz, 2 spmns.,

15 September 1959, W. J. Gertsch, V. Roth (AMNH); Pollock Pines,

2 spmns., 14 July 1948, J. W. MacSwain (NMNH); Blodgett For.,

20.8 km (13 mi) E Georgetown, 1 spmn., 27 May 1972, J.B. Heppner

(FSCA); Garden Valley, 1 spmn., 2 July 1965, S. G. Shepa (LACMNH);

and Sly Park, 2 spmns., 6 July 1958, W. J. Gertsch, V. Roth (AMNH).

Amador Co., Pine Grove, 1 spmn., 7 July 1958, W. J. Gertsch, V. Roth

(AMNH). Calaveras Co., Dorrington, 8 spmns., 12 and 29 September

1952, C. R. Snick (NMNH), 4 spmns., 11 June 1956, H. Ruckes, B.

J. Andelson (UCB), 1 spmn., 7 May 1961, W. B. Simonds (CDFA),

Centipede Subfamily Plutoniuminae 87

and 5 spmns., 27 May 1966, V. F. Lee, A. Jung (CAS). Tuolumne Co.,

Emigrant Pass, 2 spmns., 1937, M. Bocker (CAS); Strawberry, 1 spmn.,

15 June 1957, D. D. Linsdale (UCB); Pinecrest, 1 spmn., 29 June 1946,

Pearce (NMNH) and 2 spmns., 1 July 1947, P. H. Arnaud (CAS, NMNH);

Twain Harte, 2 spmns., 11 and 22 October 1948, Linsley & Smith (NMNH);

19.2 km (12 mi) E Buck Meadows, 1 spmn., 11 September 1959, W.

J. Gertsch, V. Roth (AMNH); Groveland, 1 spmn., 15 August 1957,

R. H. Goodwin (UCB); and Yosemite Nat. Pk., Aspen Valley, 2 spmns.,

4 September 1958, V. Roth (AMNH), and "14.4 km (9 mi) E Smoky

Jack," nr. Yosemite Cr. cpgd., 1 spmn., 5 July 1946, collector unknown

(AMNH)7. Mariposa Co., Miami Ranger Sta., Stanislaus Nat. For., exact

location unknown, 10 spmns., 19 and 25 July 1946, B. A. Maina (FMNH,

NMNH); Yosemite Nat. Pk., location unknown, 6 spmns., 20 May 1934,

O. Bryant (CAS) and following known sites in Park — Merced Sequoia

Gr., 1 spmn, 22 June 1990, R. M. Shelley (NCSM); Glacier Pt. Rd.,

along Ostrander Tr., 1 spmn., 4 July 1946, S. & D. Mulaik (NMNH);

Yosemite Val., 5 spmns., 24 July 1947, Lafferty (NMNH); nr. Vernal

Falls, 1 spmn., 1 July 1964, M. Kosztarab (NMNH); Bridal Veil Falls

pkg. area, 4 spmns., 21 June 1990, R. M. Shelley (NCSM); Happy

Isles, 2 spmns., 24 April 1991, R. M. Shelley (NCSM); and Mirror

Lake Loop Tr., 1 spmn., 22 June 1990, R. M. Shelley (NCSM) —

11.2 km (7 mi) NW Fish Camp, 1 spmn., 16 July 1946, H. P. Chandler

(CAS); and Fish Camp, 1 spmn., 24 March 1941, S. & D. Mulaik (NMNH).

Madera Co., Nelder Sequoia Gr., 2 spmns., 4 July 1946, R. L. Usinger,

T. O. Thatcher (NMNH); and 9.6 km (6 mi) NE Coarsegold, 1 spmn.,

24 March 1941, S. & D. Mulaik (NMNH). Fresno Co., 11.2 km (7

mi) N Badger, 8 spmns., 23 March 1941, S. & D. Mulaik (NMNH).

Tulare Co., Sequoia Nat. Pk., Crystal Cave Rd. at Marble Fork Cyn.,

1 spmn., 16 June 1990, R. M. Shelley (NCSM); 16 km (10 mi) E Three

Rivers, along Mineral King Rd., 1 spmn., 4 July 1956, W. J. Gertsch,

V. Roth (AMNH); and 16 km (10 mi) W Johnsondale, 1 spmn., 15

September 1959, W. J. Gertsch, V. Roth (AMNH). Kern Co., 17.6 km

(11 mi) E Glenville, 1 spmn., 19 March 1941, S. & D. Mulaik (NMNH).

Theatops spinicaudus (Wood)

Figs. 7-9, 19-32

Opisthemega spinicauda Wood, 1862:36, Figs. 7-8; 1865:170-171, Figs.

7Smoky Jack is an abandoned campground on the Tioga Road (California highway 120)

about 4.8 km (3 mi) west of the turnoff to White Wolf campground. On the highway

that existed in 1946, 14.4 km (9 mi) east of Smoky Jack would be near today's Yosemite

Creek Campground.

Rowland M. Shelley

Centipede Subfamily Plutoniuminae

89

Figs. 19-30. Variation of the ventral surfaces of the caudal legs and coxopleurae

of T. spinicaudus from selected localities. 19, McDowell County, North Carolina.

20, Union County, Illinois. 21, Stone County, Missouri. 22, Pike County,

Arkansas. 23, Lee County, Alabama. 24, Cobb County, Georgia. 25, Cleve-

land County, North Carolina. 26, Wilkes County, North Carolina. 27, Gra-

ham County, North Carolina. 28, Polk County, Arkansas. 29, Edgefield County,

South Carolina. 30, Mongtomery County, North Carolina. Scale lines = 1.00

mm for each figure.

90 Rowland M. Shelley

8-11. Kohlrausch, 1881:130. Meinert, 1886:208-209.

Opisthomega spinicauda: Saussure and Humbert, 1872:200.

Opisthemega insulare Meinert, 1886:209-210. Haase, 1887:79.

Theatops spinicaudus: Bollman, 1888a:6; 1888c:341. Chamberlin, 1902:41;

1920:10; 1928:153. 1942:185; 1944^:33; 19446:177-178. Kraepelin,

1903:65. Brolemann, 1904:244. Crabill, 1950:201; 19556:39.

Summers, 1979, Figs, 7-8. Shelley, 1987:505-506, Figs. 4, 13.

Summers et al., 1980:245; 1981:59.

Theatops spinicauda: Bollman, 1893:170. Pocock, 1895:28. Brolemann,

1896:50-51. Attems, 1930:253. Bucherl, 1942:326. Crabill, 1955c:157.

Kevan, 1983:2945.

Theatops posticus (nee Say): Brimley, 1938:50, in part. Wray, 1950:156,

in part; 1967:156, in part.

Type specimens. Neotype (NMNH) collected by an unknown person

on an unknown date in Chicago, Cook County, Illinois. A vial at

the ANSP, supposedly containing a paratype taken by R. Kennicott

in southern Illinois, is empty, and the holotype is not known to exist.

Diagnosis. Ultimate tergite without median suture or with only

minute vestige anteriad; ultimate legs dorsally with a distomedial spur

on each prefemur (Figs. 7-9).

Variation. I examined over 250 specimens and observed unreported

variation along the inner surfaces of the caudal legs. This surface is

generally flattened, particularly on the prefemur and femur, thus forming

a ridge along its dorsal and ventral edges. These ridges are highly

variable, and the dorsal surfaces vary from unadorned, as in an individual

from McDowell County, North Carolina (Fig. 7), to scalloped, as in

a specimen from Polk County, Arkansas (Fig. 8), to scalloped with

minute teeth, as in one from Haywood County, North Carolina (Fig.

9). The ventral ridges are more variable and display conditions with

one or more small, fine teeth on the prefemora and, in a few specimens,

the femora. Moreover, these teeth also vary in size from sharply acuminate

spurs to minute denticles. The most common condition, with no ventral

spurs or teeth is shown by an individual from McDowell County, North

Carolina (Fig. 19); eleven variants are depicted in Figures 20-30. This

variation does not conform to an observable geographic pattern; it

occurs sporadically in both areas occupied by T. spinicaudus. The medial

borders of the caudal coxopleurae are slightly elevated and prolonged

caudad, but there is a darkly pigmented subapical spot and a suggestion

of a tooth on nearly all specimens. Individuals from McDowell County,

North Carolina, and Polk County, Arkansas, have two distinct teeth

at this position (Figs. 19, 28), those in the former being larger and

Centipede Subfamily Plutoniuminae

91

resembling the conditions in T. californiensis and T. erythrocephalus

(compare Fig. 19 with Figs. 13 and 40).

Ecology. As with T. posticus, T. spinicaudus occurs primarily

in moist deciduous litter. It is occasionally found in predominantly

pine litter, and rarely under logs, loose wood debris, and large rocks.

Distribution. As shown in Fig. 32, the distribution of T. spinicaudus

is more restricted than that of the eastern population of T. posticus,

and the available records cluster into two segregated areas: the southern

Blue Ridge Province of North Carolina and the southern periphery

of Virginia, extending eastward and southward into the Piedmont Plateau

and Fall Zone of the Carolinas, Georgia, and Alabama, and westward

onto the Cumberland Plateau of Tennessee and Alabama; and from

the Central Lowlands of northeastern Illinois and central Iowa southwestward

to the Ozark and Ouachita provinces of southwestern Arkansas and

eastern Oklahoma, extending onto the Coastal Plain in southeastern

Arkansas (Fig. 31). Specimens were examined as follows; counties

are listed alphabetically for states where the species is known from

more than five counties, and complete data are provided for states

where T. spinicaudus is known from five or fewer counties. Each state

listing begins with a general description of anticipated occurrence.

Fig. 31. Distribution of T. spinicaudus.

92

Rowland M. Shelley

Fig. 32. Comparison of the distributions of T. spinicaudus (vertical shad-

ing) and the eastern population of T. posticus (horizontal shading).

ILLINOIS: Expected throughout most of the state except for the

northern and eastern peripheries adjoining Wisconsin and Indiana.

Cook, Greene, Jackson, Johnson, Monroe, Pope, and Union counties

(EIU, FMNH, INHS, NMNH, UMO). The northern- and easternmost

record is the neotype.

IOWA: Expected in the southeastern 1/3 of the state. Storey Co.,

Ames, 1 spmn., 1949, collector unknown (NMNH). Henry Co., Mt.

Pleasant, 2 spmns., date and collector unknown (NMNH).

Centipede Subfamily Plutoniuminae 93

MISSOURI: Expected statewide except for the northwestern corner.

Barry, Camden, Oregon, Reynolds, Shannon, St. Charles, St. Louis,

Stone, and Wayne counties (EIU, FSCA, INHS, NMNH, UMO).

ARKANSAS: Expected statewide except for the southern tier of

counties. Baxter, Benton, Carroll, Clark, Cleburne, Drew, Franklin,

Garland, Hot Spring, Howard, Independence, Jackson, Lawrence, Lincoln,

Madison, Montgomery, Pike, Polk, Pulaski, Saline, Searcy, Stone, Wash-

ington, and Yell counties (CAS, FSCA, INHS, MCZ, MPM, NMNH,

UAAM, UGA). Southernmost record: Drew Co., locality unknown, 6

spmns., 22 July, 21 August, and 5 November 1990, L. Thompson (UAR).

KANSAS: Expected only in the southeastern corner. Cherokee

Co., 9.6 km (6 mi) E Baxter Springs, 1 spmn., 7 April 1955, R.

W. Frederickson (SEM).

OKLAHOMA: Expected in the eastern periphery. Muskogee Co.,

locality unknown, 2 spmns., April 1957, H. Gibson (FSCA).

VIRGINIA: Expected only in the southern fringe of the Blue

Ridge Province and possibly to the west in the Ridge and Valley Province.

Scott Co., 2.4 km (1.5 mi) E Shelleys, 1 spmn., 2 May 1989, C. A.

Pague (VMNH). Carroll Co., New River Trail St. Pk., nr end of VA

hwy. 737, 3.2 km (2 mi) NNE Fries, 1 spmn., 18 September 1988,

R. L. Hoffman (VMNH); and 2.4 km (1.5 mi) NNW Lambsburg, along

Stewart's Cr., 1 spmn., 23 May 1993, R. L. Hoffman (VMNH).

TENNESSEE: Expected in the eastern 1/4 of the state, extending

westward onto the Cumberland Plateau. Greene, Hamilton, Hawkins,

Jefferson, Knox, Morgan, Sevier, Unicoi, Warren, and Washington counties

(AMNH, FMNH, MCZ, NMNH, TMM, UMMZ). Westernmost record:

Warren Co., S slope of Cardwell Mtn., exact location unknown but

probably in southeastern corner of county SE of McMinnville, 1 spmn.,

27 September 1958, T. C. Barr (NMNH).

NORTH CAROLINA: Common in the mountains and foothills,

ranging eastward into the central Piedmont Plateau (Shelley 1987).

Ashe, Avery, Buncombe, Burke, Caldwell, Cherokee, Cleveland, Gaston,

Graham, Haywood, Henderson, Jackson, Macon, Mitchell, Montgomery,

Polk, Randolph, Rockingham, Swain, Transylvania, and Wilkes counties

(AMNH, FSCA, INHS, MCZ, NCSM, NMNH, TMM). Easternmost

record: Rockingham Co., 1.1 km (4.8 mi) SW Wentworth, along co.

rd. 2192, 1.0 mi (1.6 km) N NC hwy. 704, 1 spmn., 18 April 1973,

R. M. Shelley (NCSM).

SOUTH CAROLINA: Expected in the western half of the state,

from the central Piedmont Plateau westward. Spartanburg Co., Landrum,

3 spmns., 4 August 1910, R. V. Chamberlin (NMNH). Pickens Co.,

Table Rock St. Pk., 1 spmn., 12 August 1976, R. M. Shelley (NCSM);

94 Rowland M. Shelley

and Clemson, 1 spmn., 23 May 1962, J. A. Payne (NMNH). Oconee

Co., 27.2 km (17 mi) S Highlands, NC, along SC hwy. 28, 1 spmn.,

3 June 1931, S. & D. Mulaik (NMNH); and Seneca, 4 spmns., 2 August

1910, R. V. Chamberlin (NMNH). Newberry Co., 15 km (9.4 mi) NW

Newberry, along SC hwy. 32, 1 spmn., 5 August 1976, R. M. Shelley

(NCSM). Saluda Co., 8.5 km (5.3 mi) NE Saluda, along SC hwy.

39, 1.4 km (0.9 mi) N jet. SC hwy. 450, 1 spmn., 4 May 1977, R.

M. Shelley (NCSM). Edgefield Co., ca. 14.4 km (9 mi) N of Edgefield,

along US hwy. 378, 1 spmn., 13 June 1958, collector unknown (FSCA).

GEORGIA: The northern half of the state, from the Fall Zone

northward. Bartow, Cobb, Coweta, Fulton, Habersham, Hall, Haralson,

Murray, Polk, and Rabun counties (FSCA, MEM,NCSM, MNMH, ZMUC).

Southernmost record: Coweta Co., locality unknown, 2 spmns., date

and collector unknown (ZMUC).

ALABAMA: Expected in the eastern part of state south to the

Fall Zone, with one segregated record from western Alabama near

the western terminus of the Cumberland Plateau. DeKalb Co., DeSoto

St. Pk., 1 spmn., 5 September 1966, F. A. Coyle (NCSM). Tallapoosa

Co., Dadeville, 1 spmn., 13 July 1900, W. R. Maxon (NMNH). Southernmost

record: Lee Co., Auburn, 1 spmn., April 1896, collector unknown (NMNH)

and 1 spmn., 14 June 1959, collector unknown (FSCA). Westernmost

record: Marion Co., Hamilton, bank of Buttahatchee R., 1 spmn, 18

June 1958, collector unknown (FSCA).

The following literature records are considered valid and are incorporated

into Fig. 31.

ILLINOIS: McLean, Champaign, Randolph, Williamson, Gallatin,

and Pulaski cos., (Summers et al. 1980, 1981).

MISSOURI: St. Francois Co., Libertyville (Chamberlin 19446).

Franklin Co., Sullivan; and Jefferson Co., High Ridge and Vaugirard

(Crabill 1955c).

ARKANSAS: Sevier Co., Ben Lomond; and Logan Co., Mt. Magazine

(Chamberlin 19446).

NORTH CAROLINA: Haywood/Buncombe cos., Mt. Pisgah (Wray

1950, 1967).

Remarks. Theatops spinicaudus does not occur near Pennsylvania,

so past records from this state in general, none from specific sites

or counties (Wood 1862, 1865; Saussure and Humbert 1872, Underwood

1887, Bollman 1893, Kevan 1983), represent misidentifications of T.

posticus.

Theatops phanus Chamberlin

Figs. 33-39

Centipede Subfamily Plutoniuminae

95

Figs. 33-38. Variation of the caudal legs and segment of T. phanus from

selected localities. 33-34, dorsal views. 33, Atascosa County, Texas. 34, Menard

County, Texas. 35-38, ventral views. 35, Menard County, Texas. 36, Atascosa

County, Texas. 37, Williamson County, Texas. 38, Jim Wells County, Texas.

Scale lines = 1.00 mm for each figure.

96 Rowland M. Shelley

Theatops phanus Chamberlin, 19516:101. Reddell, 1965:166.

Theatops spinicauda (nee Wood): Reddell, 1965:166.

Type specimen. Holotype (NMNH) taken by G. G. Stevenson,

16 April 1926, from an unnamed cave on his ranch near Sonora,

Sutton County, Texas.

Diagnosis. Ultimate tergite with complete median suture; ultimate

legs dorsally with a distomedial spur on each prefemur (Figs. 33-

34).

Variation. The most striking variation in T. phanus involves its

adaptability to subterranean environments and the differences between

individuals from cave and epigean environments. Most of the cave

specimens that I examined were quite large, much longer and broader

than the surface specimens from Atascosa and Jim Wells counties,

which were small and similar in size to individuals of T. posticus

from southwestern deserts. Cave individuals also display troglobitic

modifications like pallid color and longer, narrower appendages. Their

antennae extend backwards to tergites 6-7; the antennomeres are three

to five times longer than wide; and the podomeres on the penultimate

legs are four to five times longer than wide. By contrast in epigean

specimens, the antennae reach back only to tergites 3-4; the antennomeres

are approximately twice as long as wide; and the podomeres on the

penultimate legs are only two to three times longer than wide. In

both cave and surface specimens, the dorsal and ventral edges (ridges)

of the flattened inner (medial) surfaces of the ultimate legs vary as

in T. spinicaudus. The dorsal edge varies from wavy and lightly scalloped

to highly irregular with variably minute teeth (Figs. 33-34), and on

the left prefemur of the individual from Menard County, the distalmost

tooth is almost as long as the adjacent spur (Fig. 34). Ventrally, all

specimens show at least one tooth on each prefemur (examples of

variation depicted in Figs. 35-38) and epigean specimens from Atascosa

and Jim Wells counties also have teeth on the femora (Figs. 36, 38).

As with T. spinicaudus, the medial borders of the coxopleurae exhibit

subapical pigmented spots, which are developed into teeth on the indivi-

dual from Williamson County (Fig. 37).

Ecology. Chamberlin (19516) stated that the holotype was found

beneath a stone on the bottom of the first drop in the cave, and Reddell's

specimens (1965) were discovered along the banks of the stream in

Powell's Cave, Menard County. Because previous cave specimens display

elongated legs and antennae, classical adaptations to subterranean life,

T. phanus was thought to be exclusively troglobitic, but the epigean

specimens in Atascosa and Jim Wells counties lack these modifications.

The Terrell County cave specimen was found on silt 60 m (200 ft.)

Centipede Subfamily Plutoniuminae

9 7

39

Fig. 39. Distributions of F. phanus (dots) and J. posticus (stars) in Texas.

from the entrance; other vial labels lack habitat data.

Distribution. No distributional information has been published

for T. phanus. Crabill (1960) merely listed Texas without specification,

and the only published record in addition to the type locality is Powell's

Cave, Menard County (Reddell 1965, Shelley 1990a). The species occurs

in south Texas from Menard and Williamson to Jim Wells counties,

suggesting general occurrence in this region of the state; it ranges

westward onto the Edward's Plateau and may extend southward to

near the Rio Grande (Fig. 39). Specimens were examined as follows;

the exact locations of 0-9 well in Crockett County and caves in Bexar,

Burnet, Menard, Travis, and Williamson counties are unknown.

TEXAS: Terrell Co., Longley Cv., 4.8 km (3 mi) W Val Verde

co. line, 1 spmn., date unknown, J. Reddell, W. Russell (NMNH).

Crockett Co., 0-9 well, 1 spmn., 31 July 1988, G. Veni, A. Cobb,

J. Ivy (TMM) and 1 spmn; 15 August 1992, C. Savvas (TMM). Sutton

Rowland M. Shelley

Co., nr. Sonora, cave on Stevenson's Ranch, 1 spmn., 16 April 1926,

G. G. Stephenson (NMNH) TYPE LOCALITY. Menard Co., Powell's

Cv., 1 spmn., date and collector unknown (NMNH), 2 spmns., 7 September

1964, J. Reddell, D. McKenzie, B. Russell (NMNH), 1 spmn., 25 October

1980, J. Reddell, D. McKenzie (TMM), and 1 spmn., 28 January 1989,

W. Steele (TMM); Silver Mine Cv., 1 spmn., 23 January 1982, M.

Minton (TMM); 8 km (5 mi) NW Menard, 1 spmn., 5 May 1957,

S. Fowler (AAW). Burnet Co., Simons Water Cv., Lost Falls Passage,

1 spmn., 3 August 1991, M. Warton (TMM). Travis Co., Ceiling Slot

Cv., 1 spmn., 31 March 1991, J. Reddell, M. Reyes (TMM). Williamson

Co. Inner Space Caverns, 1 spmn., October 1966, B. Russell (TMM)

and 1 spmn., 22 December 1968, W. Elliott (TMM); Formation Forest

Cv., 1 spmn., 31 March 1993, J. Reddell, M. Reyes (TMM); and Water

Tower Cv., 1 spmn., 15 May 1993, J. Reddell, M. Reyes (TMM).

Bexar Co., Robber Baron Cv., 1 spmn., 19 June 1993, J. Loftin (TMM).

Atascosa Co., Jourdanton, 1 spmn., 27 November 1935, S. Rutherford

(NMNH). Jim Wells Co., Alice airport, 1 spmn., 3 February 1962,

R. O. Albert (NMNH).

Theatops erythrocephalus (C. L. Koch)

Figs. 40-41

Cryptops erythrocephalus C. L. Koch, 1847:173-174; 1863:99-100, Figs.

221a,b. Kohlrausch, 1881:130.

Opisthemega erythrocephalum: Latzel, 1880:147-149. Kohlrausch. 1881:131.

Latzel, 1880:147-148. Daday, 1889:92.

Opisthemega lusitanum Verhoeff, 1896:78-79.

Theatops erythrocephalus: Kraepelin, 1903:66-67, Fig. 26. Attems, 1929:299.

Verhoeff, 1941:figs. 89-90.

Theatops erythrocephala: Attems, 1930:251-252, Figs. 4, 27, 32, 331-

335; 1959:319. Foddai et ah, 1995:8.

Theathops erythrocephalus breuili Matic 1960:446-447, Figs. 7-8.

Theatops erythrocephala erythrocephala: Matic, 1960:447.

Type specimen. Most of Koch's centipede types are deposited

in the NHM, but that of C. erythrocephalus is not labeled as such

(Minelli, in litt.) and could not be located by the curator. According

to Koch (1847) the type was collected by Prof. Dr. V. Siebold at Pula

on the Istrian peninsula, Croatia.

Diagnosis. Ultimate tergite usually with incomplete median suture,

running from anterior margin to just short of caudal edge; ultimate

legs without dorsal distomedial prefemoral spurs; ultimate prefemora

and femora usually with four strong, distinct ventral spurs, one on

Centipede Subfamily Plutoniuminae

99

Figs. 40-43. Variation of the ventral surfaces of the caudal legs and segment

of T. erythrocephalus. 40, specimen from Sipan Island, Croatia. 41, specimen

from Portugal, locality unknown. 42-43, caudal segment and legs of a speci-

men of P. zwierleini from Sardinia. 42, dorsal view. 43, ventral view. Scale

lines = 1.00 mm for each figure.

100 Rowland M. Shelley

each podomere, rarely with fewer spurs; caudal coxopleurae with medial

borders strongly elevated and extended caudad, apically acuminate with

blackened subapical spurs (Figs. 40-41).

Variation. The NCSM specimen from Sipan Island, Croatia, lacks

the ventral spur on the left prefemur (Fig. 40), which contrasts with

the normal condition as in the specimen from Portugal (Fig. 41). One

caudal leg on a ZMH specimen from Rijeka (= Fiume), Croatia, is

much smaller, appears to be regenerating, and lacks both spurs.

Ecology. To my knowledge, no habitat information has been published

on T. erythrocephalus. However, Kos (1992) records it from mediterranean

and submediterranean districts in Croatia, Bosnia-Hercegovina, and

Montenegro.

Distribution. European, occurring in two areas segregated by some

992 km (620 mi), one in the Balkan Peninsula along the Adriatic

Sea, extending from the Istrian peninsula of Croatia to the southern

coastal extremity of Montenegro below Lake Scutari and probably also

into Albania, including offshore islands along the coast of Croatia,

and the other in Spain and Portugal, probably along the Mediterranean

Coast south of Barcelona and the Atlantic Ocean west of the Strait

of Gibralter (Figs. 44, 45). The following literature records cannot

be placed today because they refer to general areas instead of specific

sites and because of political changes in the Balkan peninsula during

the First and Second World Wars and the currently chaotic situation

in this war-torn region: Kraepelin (1903) — Hungary, Dalmatia (southern

coastal Croatia), Portugal, and Italy, the last erroneously referring to

P. zwierleini; and Attems (1930) — Montenegro, Dalmatia, "Kroatisches

Litorale" (roughly equivalent to Dalmatia), Istria, South Hungary, and

Portugal. However, enough specimens and specific literature records

exist that the distribution in the Balkans can be defined as the Adriatic

coastal region from the Istrian peninsula to the southern extremity

of Montenegro, extending inland some 48 km (30 mi) to Mostar, Bosnia-

Hercegovina. The last record may represent dispersal up the Neretva

River Valley, which flows through Mostar to the Adriatic Sea. Fewer

specimens and specific literature records are available from the Iberian

Peninsula, but they suggest occurrence in a narrow band along the

Mediterranean Coast from Barcelona to Gibralter, and continuing along

the Atlantic Ocean into Algarve Province, Portugal. The available evidence

thus indicates a primarily coastal distribution for T. erythrocephalus

in both the Balkan and Iberian peninsulas, and the question mark in

Figure 44 is placed in southern Portugal because of the known coastal

records in Spain. The specimens examined, and literature and other records,

are as follows; where the name of a city has changed from that in the

Centipede Subfamily Plutoniuminae 101

literature or on the vial label, the modern name is cited first with the

older equivalent in parentheses:

CROATIA: Istrian Peninsula, Rijeka (=Fiume), 2 spmns., 1897,

collector unknown (ZMH). Sipan Island, Luka, 1 spmn., date and collector

unknown (NCSM). Dalmatia, locality unspecified, 1 spmn., 1 January

1899, collector unknown (ZMH).

BOSNIA-HERCEGOVINA: Exact locality unknown, 3 spmns.,

1903, collector unknown (ZMH).

MONTENEGRO: Bar (=Antivari), 2 spmns., date and collector

unknown (MCZ).

SPAIN: Barcelona Prov., Barcelona, 3 spmns., October 1927,

collector unknown (ZMH).

PORTUGAL: Province and locality unspecified, 2 spmns., 30 January

1900, collector unknown (ZMH) and 2 spmns., date unknown, K. W.

Verhoeff (MCZ).

The following literature records are incorporated into Figures

44 and 45.

CROATIA: Velebit Mtns., Senj (=Zengg) (Attems 1929). Istrian

Peninsula: Pula (Attems 1929) TYPE LOCALITY. Dalmatia: Zadar

(=Zara) (Attems 1929); Kali (a community on Pasman Island directly

west of Zadar) (=Kali Pecina) (Attems 1959); Dugi Island (=Isola Grossa,

the outermost island in the Adriatic Sea due west of Zadar), Dubrovnik

(=Ragusa), Lapad (a town in the metropolitan area of Dubrovnik, at

the tip of its peninsula), and Pridworje, ca. 24 km (15 mi) SE Dubrovnik,

below Zupski Bay (Attems 1929).

BOSNIA-HERCEGOVINA: Trebinje (a town below Mostar and

ENE of Dubrovnik) (Attems 1929); along the Trebisinca River (a river

just inside Bosnia-Hercegovina border that flows through Trebinje and

parallel to the border) (=Vodena Dolina am Popovo Polje) (Attems

1959); Diklici (a community near Trebinje on the Trebesinca River)

(=Pecina bei Diklici) (Attems 1959); Mostar (Attems 1929); Konjsko

(a small town just SE of Trebinje) (Attems 1929); Prenj (a community

just inside the Bosnia-Hercegovina border south of Mostar) (Attems

1929); Plasa, exact location unknown (Attems 1929); and Mljet (=Meleda)

Island (Attems 1929, 1959).

MONTENEGRO: Hercegnovi (=Castelnuovo) (Attems 1959); Kotor

(on south end of Kotorski Bay) (=Cattaro), Njegus (a small community

slightly north of Kotor), Virpazar (on the western shore of the northern

end of Lake Scutari), Cetinje, Titograd (=Podgorica), and Ulcinj (=Dulcigno)

(Attems 1929).

SPAIN: Alicante Prov., Denia, ca. 72 km (45 mi) NE Alicante,

Cueva de la Punta de Benimaquia (Ribaut 1915), Valencia Prov., Gandia,

102 Rowland M. Shelley

ca. 60.8 km (38 mi) SSE Valencia, Cueva Negra de Palma (Ribaut 1915).

Malaga Prov., Cueva del Cerro de la Pileta, nr. Ronda, nearest town

Benaojan, ca 100 km (62.5 mi) NNE Gibraltar (Matic 1960, Wiirmli

1975).

The following unpublished records were communicated by A. Minelli

and are incorporated into Figures 44 and 45.

CROATIA: Krk Island.

SPAIN: Granada Prov., Sierra Nevada and Capileira de Poqueira

(possibly referring to a small town on the southern slope of the Sierra

Nevada).

Deleted records. The following literature records are deleted. Minelli

(1991) does not include T. erythrocephalus in his list of centipedes

in northeastern Italy and does not anticipate its discovery at Trieste;

a voucher specimen has never been located, and an old Trieste label

could refer to a site in Istria (Minelli, in lift.). Foddai et al. (1995)

list T. erythrocephalus as a questionable inhabitant of northern Italy

and state that its presence should be confirmed. Kos (1992) does not

record T. erythrocephalus from Slovenia, and this location, some 128

km (80 mi) inland, is implausible for the species.

ITALY: Trieste Prov., Trieste (Attems 1929).

SLOVENIA: near Brestanica (=Bucerca-Hohle bei Reichenburg,

Sudsteiermark (Attems, 1959)).

Genus Plutonium Cavanna

Plutonium Cavanna, 1881:169. Kraepelin, 1903:67. Verhoeff, 1907:253.

Attems, 1930:253. Foddai et al., 1995:8.

Type species. P. zwierleini Cavanna, 1881, by monotypy.

Diagnosis. With 19 pairs of spiracles, on segments 2-20.

Distribution. Known from Granada Province, Spain, and the following

regions of Italy: Sicily, southeastern Sardinia, and coastal Campania,

particularly the Sorrento Peninsula (Fig. 44) (Wiirmli 1975, Foddai

et al. 1995).

Species. One.

Remarks. A check of Attems (1930) and subsequent publications

reveals that Plutonium and Tonkinodentus are the only known cryptopid

genera that are absent from the Western Hemisphere and that the family

Cryptopidae is primarily an "American," New World, taxon.

Plutonium zwierleini Cavanna

Figs. 42-43

Plutonium zwierleini Cavanna, 1881:169-170, Figs. 1-7. Kraepelin, 1903:67-

68, Fig. 27.

Centipede Subfamily Plutoniuminae

103

Fig. 44. Distributions of T. erythrocephalus (stars) and P. zwierleini (dots).

Some symbols of T. erythrocephalus in Croatia and Montenegro denote more

than one locality; the site in Portugal is unknown and indicated by the question

mark.

Plutonium zwierleinii: Verhoeff, 1906:387. Attems, 1926:Fig. 433; 1930:253-

254, Figs. 336-338. Foddai et al., 1995:8.

Type specimen. Neotype (ZMH) collected by G. A. Markens, 16

May 1898, at Palermo, Sicily, Italy. The holotype is not known to

exist; it is not housed at either museum in Florence, Italy, the Museo

Zoologico de La Specola (Zoological Museum of Florence University)

or the Instituto Sperimentale per la Zoologia Agraria (Minelli, in litt.).

According to Cavanna (1881), the holotype was collected in 1878 at

an unspecified location, probably in Sicily, by Dom. Eq. Zwierlein.

Diagnosis. With the character of the genus (Figs. 42-43).

Variation. The few specimens that I examined agree with the description

by Cavanna (1881). Wiirmli (1975) also does not indicate significant

variation.

Ecology. According to Wiirmli (1975) and Minelli and Iovane

(1987), P. zwierleini occurs from 0-900 m (0-2,952 ft.) in seashore

environments and woodlands; it is usually encountered under large rocks

or small stones, where soil moisture is preserved.

104

Rowland M. Shelley

Fig. 45. Distribution of T. erythrocephalus in the Balkan Peninsula.

Distribution. Same as that of the genus (Fig. 44). Wiirmli (1975)

discusses the authentic localities and erroneous ones reported by previous

authors. In addition to the neotype, specimens were examined as follows:

ITALY: Salerno Prov., Salerno, 1 spmn., 1928, collector unknown

(ZMUC). Sardinia, Assuni, 1 spmn., 30 June 1911, N. L. H. Krausse

(NMNH); and Lanusei, 1 spmn., 23 October 1899, collector unknown

(ZMH).

The following localities were communicated by the indicated colleague

and are incorporated into Figure 44.

ITALY: Napoli Prov., Sorrento. Sicily, San Cataldo, ca. 6.4 km

(4 mi) W Caltanissetta (H. Enghoff).

SPAIN: Granada Prov., Orgiva (A. Minelli).

Remarks. The ultimate legs and segment of the examined specimens

Centipede Subfamily Plutoniuminae

105

of P. zwierleini are identical to those in the eastern population of T.

posticus (compare Figs. 6 and 42 and Fig. 43 with Fig. 6 in Shelley

(1990a)). The legs lack dorsal or ventral spurs, and the segment possesses

a complete median dorsal suture and has rounded, nonextended coxopleurae.

The only detectable distinction between P. zwierleini and eastern forms

of T. posticus is the different number of spiracles.

RELATIONSHIPS

In assessing relationships among the plutoniuminine taxa, P.

zwierleini is obviously the sister-group to the five species of Theatops.

Within the latter, T. spinicaudus, which is unique in lacking a median

suture on the ultimate tergite, is the sister-group to the other four

species. Theatops phanus, with the dorsal prefemoral spur, is then

sister to the three species lacking this structure, and because T. posticus

and T. californiensis were once geographic races of a single species,

as shown by the residual intergrade specimens in the southwestern deserts

(Shelley, 1990a), T. erythrocephalus is sister to its American counterparts

(Fig. 44). Most of these proposed lineages cannot now be defined by

autapomorphies, and comparative biochemical investigations may be necessary

to elucidate such characters because of the high degree of phenotypic

similarity among the members of this subfamily.

Their Holarctic distributions indicate a Laurasian origin for both

the subfamily and the genus Theatops. Aside from Plutonium and Theatops,

Theatops

R zwierleini

spinicaunus

phanus erythrocephalus posticus

califomiensis

Fig. 46. Relationships in the Plutoniuminae.

106 Rowland M. Shelley

the only European cryptopid genus is Cryptops Leach (Cryptopinae),

which also has 21 leg pairs and pedal segments, with slightly enlarged

caudal legs. The Cryptopinae, a global taxon, also is indigenous to

the Nearctic, whereas the other cryptopid subfamily, the Scolopocryptopinae,

with narrow caudal legs and 23 leg pairs and pedal segments, is primarily

a New World group with minor representation along the western Pacific

Rim from Japan to New Guinea (Attems 1930). The Plutoniuminae

and Cryptopinae therefore logically share ancestry and may antedate

the Scolopocryptopinae, whose concentration in the Americas suggests

a post-Laurasian origin. Its diversity and abundance in North and South

America probably reflect considerable northward and southward dispersal

after closure of the Panamanian portal, and the occurrence of Scolopo-

cryptops Newport in Japan, Korea, and China surely represents a Pleistocene

invasion of Asia via the Bering Land Bridge. However, this genus

and the subfamily also occur in the Philippines, Viet Nam, New Guinea,

Sulawesi, and the Sunda and Fiji Islands (Attems 1930, Schileyko

1995), and their existences in these areas, if native and not the result

of introductions, hardly represent trans-Beringian dispersal. With this

circum-Pacific distribution, the Scolopocryptopinae may be a chilopod

analog to the diplopod family Cambalidae (order Spirostreptida), whose

biogeography was attributed to the lost continent "Pacifica" by Jeekel

(1985). Not nearly enough is known about scolopocryptopinine bio-

geography to further explore this possibility, but it raises questions

about the composition of the Cryptopidae, because an independent bio-

geography for the Scolopocryptopinae implies a different origin and

phylogeny. This in turn implies that concordance with the Cryptopinae

and Plutoniuminae in the absence of ocelli represents convergence rather

than shared ancestry; consequently, the Scolopocryptopinae may merit

separate family status.

The prevailing concept of the Scolopendromorpha recognizes two

families, Scolopendridae and Cryptopidae, based primarily on the presence

and absence, respectively, of four ocelli on each side of the cephalic

plate. Schileyko (1992) proposed a new arrangement derived in part

from that of Haase (1887), but this system is incomplete, omitting

at least three cryptopid genera, Dinocryptops Crabill (1953), Thalkethops

Crabill (1960), and Ectonocryptops Crabill (1977). Furthermore, it is

not based on a rigorous assessment of shared, derived features, so there

is no assurance that the groupings are monophyletic lineages representing

true lines of affinity. Many more alpha- and beta-level generic studies

must be conducted in the Scolopendromorpha before the families can

be reappraised and subjected to an intensive cladistic analysis, but no

longer should the present division, based primarily on the presence or

Centipede Subfamily Plutoniuminae 107

absence of eyes, be uncritically accepted.

ACKNOWLEDGMENTS— Vox loan of pertinent type specimens I

thank P. Hillyard (NHM) (C. postica), H. W. Levi (MCZ) (O. insulare),

and J. A. Coddington (NMNH) (T. californiensis, O. spinicauda, T.

phanus). The following curators and collection managers loaned material

from the indicated collections: N. I. Platnick (AMNH), D. Azuma and

the late S. S. Roback (ANSP), W. E. Clark (AU), A. R. Hardy (CDFA),

W. J. Pulawski (CAS), A. H. Barnum (DC), R. E. Funk (EIU), D.

Summers (FMNH), G. B. Edwards (FSCA), K. C. McGiffen (INHS),

the late C. L. Hogue (LACMNH), the late D. K. McE. Kevan (LEM),

H. W. Levi (MCZ), R. W. Brown (MEM), J. P. Jass (MPM), J. A.

Coddington (NMNH), R. Irving (OPCNM), R. W. Brooks (SEM), J.

R. Reddell (TMM), J. B. Whitfield (UAR), C. A. Olsen (UAZ), J.

A. Chemsak and C. B. Barr (UCB), the late R. O. Schuster (UCD),

J. E. O'Donnell (UCT), S. I. Frommer (UCR), C. L. Smith (UGA),

C. V. Covell (UL), M. W. O'Brien (UMMZ), R. W. Blinn (UMO),

R. L. Hoffman (VMNH), M. C. Thomas (WVDA), H, Enghoff (ZMUC),

and G. Rack (ZMH). W. A. Shear loaned his private collection, and

A. A. Weaver donated his chilopod holdings, containing numerous

specimens of Theatops, to the NCSM. Collecting in Yosemite and

Sequoia/Kings Canyon National Parks in 1990 and Yosemite in 1991

was conducted under permits issued by the park offices. Special thanks

are extended to G. C. Steyskal, for advice on the "-ops" suffix; to

J. G. E. Lewis and R. L. Hoffman, for valuable commentaries on preliminary

drafts of the manuscript; and to A. Minelli, for advice on T. erythrocephalus,

P. zwierleini, and Balkan and Italian localities, for permission to cite

his Spanish record of P. zwierleini, and for an insightful prepublication

review. Balkan localities of T. erythrocephalus, many from early publications

with pre-World War I geography and archaic place names, were located

with the invaluable assistance of Celia Pratt, librarian at the University

of North Carolina Map Library. Figures 2-9, 13-17, 19-30, 33-38,

and 40-43 were prepared by R. G. Kuhler, NCSM scientific illustrator;

Cathy Wood performed repeated word processing chores.

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Received 15 April 1996

Accepted 1 May 1996

114

DATE OF MAILING

Brimleyana 23 was mailed on 16 July 1996.

ERRATA

The following correction should be made in "Mensural

Discrimination of Four Species of Peromyscus (Rodentia: Muridae)

in the Southeastern United States" by Joshua Laerm and James

L. Boone {Brimleyana 21:107-123, December 1994). In Table

2 (p. 114), the constant given for Peromyscus leucopus is -32.229.

The correct constant is -38.229.

115

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BRIMLEYANA NO. 24, APRIL 1997

CONTENTS

The Rock Shrew, Sorex dispar (Insectivora: Soricidae), in Georgia with Comments on its

Conservation Status in the Southern Appalachians.

Joshua Laerm, Charles H. Wharton, and William Mark Ford 1

Dog Burials from the Eighteenth Century Cherokee Town of Chattooga, South Carolina.

Gerald F. Schroedl and Paul W. Parmalee 7

Mensural Discrimination of Sorex longirostris and Sorex cinereus (Insectivora: Soricidae)

in the Southeastern United States. Joshua Laerm, Michael A. Menzel, and

James L. Boone 15

Clam Siphon Tip Nipping by Fishes in the Estuarine Cape Fear River, North Carolina.

Frank J. Schwartz 33

Condylura cristata (Insectivora: Talpidae) in the Blue Ridge Province of Western

South Carolina. Joshua Laerm, Gayle Livingston, Christine Spencer, and

Bryan Stuart 46

The Holarctic Centipede Subfamily Plutoniuminae (Chilopoda: Scolopendromorpha:

Cryptopidae) (Nomen Correctum Ex Subfamily Plutoniinae Bollman, 1893).

Rowland M. Shelley 51

Miscellany 114