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JOHN E. COOPERE,d itor
ALEXA C. WILLIAMS, Managing Editor
JOHN B. FUNDERBURGE, ditor-in-Chief
Board
1 ALVINL . BRASWELLC,u rator of DAVIDS. LEE, ChzefCurator
Lower VerfebraterN, .C. ojBlrds and Mammals, N C.
State Museum State Museum
JOHN C. CLAMPA,s so~tateC uralor WILLIAMM. PALMERC,h lcfCuralot.
(Invertebrates), N.C. of Lower Vertebrates, N.C.
State Museum State Museum
MARTHRA. COOPERA, ssoc~ate THOMLA. SQ UAY,D epartmmt
Curalor (Crustaceans),N .C. of ~ o o l oN~.C, . State
State Museum Untverstty
JAMES W. HARDIND, epartment ROWLANMD SHELLEYCh, tef
of Botany, N C. State Curator of Inverlebratcs, N C.
Unauers~ty State Museum
Brirnleyana, the Journal of the North Carolina State Museum of Natural His-tory,
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CODN BRIMD 7
ISSN 0193-4406
The Milliped Fauna of the Kings Mountain Region
of North Carolina (Arthropoda: Diplopoda)
MARIANNEE. FILKA
Department of Zoology,
North Carolina State University,
Raleigh, North Carolina 27607
AND
ROWLANDM . SHELLEY
North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh, North Carolina 276111
ABSTRACT.-The millipeds of the Kings Mountain region of Cleve-land
and Gaston counties, one of five inselberg areas in the Piedmont
Plateau of North Carolina, were sampled to determine seasonal varia-tion
in faunal composition. Comparative collections also were made at
Spencer Mountain, an inselberg located northeast of Gastonia. Of 24
species taken, only Narceu americanus (Beauvois) and Aururus
eryrhropygos (Brandt) were present as adults and juveniles in all three
sampling seasons. The most diverse assemblage was encountered in Oc-tober.
Five species were more common in April and October than in
Julv. four were more common in Julv than in either of the cooler
mdnths, and five others were collected~in but a single season (three in
Julv. two in October). A more diverse fauna was encountered in the con-tigkous
~ings-CroGders ridge than at the isolated Spencer Mountain
outcrop, from which three xystodesmids were conspicuously absent. A
notabli dilfeercnce between millipeds of the trr o loca-lilies ~nvolved color
pattern of the intergrades oFSlgmoria lal~u(rB r3lsmann). P~yoiulurw as
ihe sole genus represented by~moreth an one species, and the overall
species/genus ratio is indicative of a lowland rather than a montane
fauna.
The Kings Mountain region shares eight species with the eastern
Piedmont and five with the Appalachian Mountains. Seven widespread
species occur in all three regions, but three species are unique to the
Kings Mountain region. This area is the northeastern range limit of the
genus Pachydesmu; and the easternmost populations of four montane
diplopods, the westernmost population of Ptyoiulus ectenes (Bollman),
and the southernmost known locality of Cleidogona medialis Shelley, oc-cur
there. The conservation status of three species of concern to North
Carolina is discussed. and the Kines Mountain region is considered a
"cluster area" due to'its unique didopod fauna. d he ranges of Bornria
stricta (Brolemann) and Deltotaria lea Hoffman are extended into South
~arolin'aA. key tdgenera and species is provided along with pertinent
diagnostic illustrations.
'Adjunct Asststant l'rofcssor ofZoology, North Carolina State University
Brimlmna No. 4: 142. Dmmber 1980. 1
2 Marianne E. Filka and Rowland M. Shelley
INTRODUCTION
The importance of the Appalachian Mountains to the arthropod
class Diplopoda has been evident since 1969, when Hoffman identified
the mountains as a global center of milliped evolution. This opinion was
based on the diversity and abundance of known indigenous taxa. Four
other areas also were cited as important global centers of evolution and
dispersal, and since all are mountainous to some extent, Hoffman sur-mised
that vertical relief probably allows for a greater variety of
ecological niches than occur in lowland or flat areas.
The Appalachian Highlands, one of eight physiographic divisions of
the United States, is comprised of seven physiographic provinces (Hunt
1967). The most important in terms of known diplopod faunas are the
Ridge and Valley and Blue Ridge Provinces, especially the southern sec-tion
of the latter (the region south of the Roanoke River). The
Xystodesmidae, the dominant Nearctic polydesmoid family, attains its
greatest known diversity in the part of the southern Blue Ridge Province
south of the Kanawha River System (Hoffman 1969). The bulk of the
southern Blue Ridge Province is in western North Carolina, where it is
demarcated from the Piedmont Plateau by a prominent escarpment, the
Blue Ridge Front. Thus, for all practical purposes one of the five regions
of greatest milliped diversity in the world lies in the western part of this
state.
Although most of the mountains of North Carolina are west of the
Blue Ridge Front, a number of prominent hills and ridges also occur to
the east in the Piedmont Plateau. Some of these are quite properly called
mountains and extend to altitudes of well over 300 meters. These isolated
mountains protruding from a surrounding flat plain are known as in-selbergs
and are erosional remnants of previously more extensive moun-tain
masses (Kesel 1974). Five main groups of inselbergs occur in Pied-mont
North Carolina (Fig. 1): the Sauratown Mountains of Stokes
County (including Pilot Mountain, Surry County); the Brushy Moun-tains
of Wilkes, Caldwell, Alexander and Iredell counties; the South
Mountains of Burke, Rutherford, McDowell and Cleveland counties; the
Kings Mountain region of Cleveland and Gaston counties; and the
Uwharrie Mountains of Davidson, Randolph, Montgomery, and Stanly
counties. The faunas of these inselberg regions are of particular
biogeographic interest and raise a number of questions. Do they, for ex-ample,
reflect previous direct connection with the Blue Ridge chain? If
so, their later isolation may have separated previously continuous
diplopod populations and led to speciation by geographical isolation.
Accordingly, knowledge of the inselberg diplopod faunas may provide
insights into processes affecting milliped evolution, and an investigation
of one such area was conducted in this study.
The Kings Mountain region straodles the South Carollna border
about 13 km southwest of Gastonia. Preliminary studies there had dis-closed
a milliped fauna with southern elements found nowhere else in
Kings Mountain Milliped Fauna 3
,. "2".
z, ..".h. ",". S.",h St". * "I. "1" 1.s1.1
""I..."I,".. - "
Fig. I. Major inselberg regions of North Carolma.
North Carolina. The other inselberg regions have no faunas of such
singular importance to the state. Information on unique areas in North
Carolina is timely in regard to recent concerns about environmental
management and planning, as reflected by the North Carolina Environ-mental
Policy Act of 1971; the State Land Policy and Coastal Area
Management Acts, both enacted in 1974; and the Symposium on En-dangered
and Threatened Biota of North Carolina (see Cooper et al.
1977). Moreover, the North Carolina Natural Heritage Program, ad-ministered
by the Department of Natural Resources and Community
Development through a contract with The Nature Conservancy, is
presently conducting an inventory of the state's most significant natural
areas. In order to realize the goals of these programs and to effectively
manage the resources of the state, knowledge of its indigenous flora and
faunamuct be substantiated. Another ohjectlve of this project, therefore,
was to furnish such knowledge for the Diplopoda of the K~ngsh lountain
area, and categories of concern are suggested in some of the species ac-counts.
This report includes a key to genera and species, and gonopod il-lustrations
to assist in determinations. Accounts are presented for each
milliped species collected, along with synonymies for the two species
previously reported from the region or vicinity. Numeric ratios of or-ders/
families/genera/species (o/f/g/s) and species/genera (s/g) are used
to show diversity and seasonal variation within the Kings Mountain
fauna and to compare it with the faunas of theeastern Piedmont and the
Great Smoky Mountains (Tables 10-12). Comments on seasonal oc-currence
of adults and juveniles are provided in the species accounts and
summarized in Table 10. Noteworthy behavior and gonopodal variation
are discussed for each species where appropriate. Localities are listed for
species collected from fewer than six sites and for three diplopods con-sidered
of Special Concern in North Carolina, as defined in Cooper et al.
(1977). Due to present nomenclatorial confusion and in deference to
current work by other specialists, as explained in appropriate accounts,
4 Marianne E. Filka and Rowland M. Shelley
specific names cannot he assigned for two millipeds and provisional
names are used for two others. The major concern of this study was the
fauna of natural habitats, and synanthropic diplopods were thus incom-pletely
sampled. Additional species that might be discovered in future in-vestigations
are discussed in the final section.
Fig. 2. The Kings Mountain region of North Carolina.
THE KINGSM OUNTAINR EGION
The Kings Mountain range extends northeastward as a linear ridge
some 26.5 km from the southern tip of Cherokee and York counties,
South Carolina, to the southeastern corner of Cleveland and south-western
part of Gaston counties, North Carolina (Fig. 2). It is hounded
on the east and west by Crowders and Kings creeks, respectively, and sur-rounded
by Piedmont Plateau. Isolated outlying peaks, inselbergs of the
Kings Mountain ridge, continue northeastward approximately 64 km to
Anderson's Mountain in Catawha County. The hulk of the region is
located about 136 km east of the Blue Ridge Front in Cleveland and Gas-ton
counties, where it covers an area of approximately 3108 hectares. It
consists of four main groups of lowlying peaks separated by gaps. Mean
elevation is 361 m above sea level with maxima of 570 m (the Pinnacle)
and 474 m (Crowders Mountain). Spencer Mountain, a 378 m high in-selberg
of the Kings Mountain ridge, is located about 14.5 km northeast
of Crowders Mountain on the opposite side of Gastonia.
The Kings Mountain geologic belt, composing the range, is a narrow
zone of metamorphosed sedimentary rocks (schist, marble, and
quartzite) of Paleozoic age (Stuckey 1965). The porous nature of this
rock produces a bountiful supply of ground water, and natural springs
and seeps are characteristic of the area (Keith 1931). Soil composition
Kings Mountain Milliped Fauna 5
varies from thick black peaty humus in forested areas, to exposed red
clays and yellow silts on eroded downslopes, to glittering micaquartzite
sand along stream banks and on summits. Xeric scrub forests similiar to
those found in the Blue Ridge characterize these summits, and hardwood
forests, remnants of the previous oak-hickory and beech-maple climaxes,
distinguish relatively undisturbed regions on surrounding lower slopes.
In clear-cut or burned areas, dense stands of Virginia pine, Pinus
virginiana, and shortleaf pine, P. echinata, dominate to the exclusion of
other species. Various pine-hardwood mixtures occur in disturbed areas
throughout the Kings Mountain region (Burney 1974).
MATERIALS AND METHODS
Millipeds were sampled in July and October 1976, and April 1977, to
investigate seasonal variation in faunal composition. Collecting was done
primarily in the contiguous ridge area around Kings and Crowders out-crops,
but four sites around Spencer Mountain also were sampled for
comparison. The South Carolina state line was selected as the southern
boundary, and collecting limits were set in other directions using
topographical and county road maps. Collecting sites, chosen to provide
a maximum diversity of habitats, included pine, mixed pine-hardwood,
and deciduous forests; banks of streams and ponds; seepage areas; bor-ders
of flat meadows; gradual slopes and steep hill terrains; bases, slopes,
and summits of outcrops; and,trash dumps. Climatic conditions varied
from hot and dry in July to cool and damp in October and April.
Twenty-five sites were examined during the first trip (July). Five of
these yielded few millipeds because of scant leaf litter, so only the twenty
remaining sites were routinely sampled on all trips. Several new prospec-tive
sites were visited during each ensuing trip. Specimens were collected
from beneath leaf litter, bark of decaying logs, and large rocks, and
preserved in 70% isopropanol. Pine, hardwood, and mixed pine-hard-wood
litter samples were collected for extraction with Berlese funnels.
Notes on color, behavior, and habitat were recorded at each site.
Measurements of length and width in mm were taken with vernie~
calipers. Drawings of most gonopods and all other structures were
prepared with the aid of a grid reticle with 0.5 mm squares, but a camera
lucida was used for the smallest gonopods, which were mounted tem-porarily
in glycerine jelly and examined with a compound microscope.
All other structures were examined using a stereomicroscope, with the
specimens immersed in 70% isopropanol and stabilized by cotton. More
than 1000 specimens were examined, some of which were collected in
preliminary studies. All specimens are deposited in the invertebrate
collection of the North Carolina State Museum of Natural History
(NCSM), the invertebrate catalogue numbers of which are indicated in
parentheses with appropriate citations. A single pertinent specimen was
found in the collection of the American Museum of Natural History
(AMNH).
In the species accounts and legends, CR means country road.
6 Marianne E. Filka and Rowland M. Shelley
KEY TO GENERA AND SPECIES OF
KINGS MOUNTAIN DIPLOPODS
1. Body soft, exoskeleton noncalcified; clusters of modified setae
adorning head and tereites: terminal setal tufts aresent aosterior- - ,
ly; male; without gonopods; adults less than 3'mm long (Penicil-ata,
Polyxenida, Polyxenidae ...... Polyxenus fasciculatur Say
Exoskeleton hard, calcified; setae normal, scattered, males with
gonopods on 7th or 7th and 8th segments; adults varying in size
but always longer than 3 mm (Helminthomorpha) ........... 2
2. Head reduced; males with 8 pairs of legs preceding gonopods . . 3
Head normal; males with 7 pairs of legs preceding gonopods ... 4
3. Three pairs of ocelli present; paranota absent; segments arched, con-vex
dorsally (Polyzoniida, Polyzoniidae) . . . . . . . . . . . . . . . . . . . . .
.............................. Polyzonium strictum Shelley
Ocelli absent; paranota present, bilobed on segment five (Fig. 4);
segments narrow, flattened (Platydesmida, Andrognathidae)
............................ Andrognathus corticarius Cope
4. Ocelli present; paranota absent; adults with more than 20 seg-ments
................................................5
Ocelli absent; paranota present; adults with 19 or 20 segments
(Polydesmida) ..................... ... .............. 16
5. Segments with dorsal longitudinal crests .................... 6
Without this character ................................... 9
6. Collum enlarged, hoodlike, covering epicranial region of head . 7
Collum reduced, head completely. ex.p osed (Callipodida, Casp.i op.e -
talidae) .............................................. 8
7. Epiproct trilobed (Fig. 11); adults with 30 segments; adult length
not exceedin-g 26 mm (Cho.r.d.e umida. Stridriidae) ............. ..................... . . . . . . . . . . . . . . . . . . . . Striaria sp.
Epiproct entire; adults with more than 30 segments; adults 40-50 mm
long (Spirostreptida, Cambalidae) .... Cambala annulata (Say)
8. Coxal process of gonopod thin, translucent, ensheathing stalk of
telopodite; flagellum absent (Figs. 25-26) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Delophon aeoraianum Chamberlin
Coxal process of gonopod thick, opaque, bsnt Gterad apically, not
ensheathing stalk of telopodire: flagellum present (Fig. 20) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A bacion magnum (Loomis)
9. Adults with 28-30 segments (Chordeumida) . . . . . . . . . . . . . . . . 10
Adults with more than 30 segments ....................... 11
10. Ocelli arranged curvilinearly, 6 per row; adults not exceeding 5 mm
long (Trichopetalidae) ...... Trichopetalum dun (Chamberlin)
Ocelli in triangular patch, 26 per patch; adults longer than 5 mm
(Cleidogonidae) . . . . . . . . . . . . . . . .C leidogona medial& Shelley
11.. Coxae of legs 3-7 of males with lobed extensions (Fig. 16); ocelli
in ovoid patch; gonopods concealed within body; adults large,
robust, 80-100 mm long (Spirobolida, Spirobolidae) ..........
. . . . . . . . . . . . . . . . . . . . . . . . . . . .N arceus americanus (Beauvois)
Kings Mountain Milliped Fauna
Coxae of pregonopodal legs of males without lobed extensions;
ocelli variable in arrangement; gonopods completely visible or
concealed within body; adults slender, never exceeding 5 mm
long (Julida) .................... ... ............... 12
Ocelli arranged linearly (Blaniulidae) Nopoiulus minutus (Brandt)
Ocelli in triangular patch ................................ 13
Gonopods completely concealed within body; first pair of legs of
male reduced, hooklike, dorsum with 2 yellow longitudinal
strips containing narrow median black line (Julidae) .........
............................ Brachyiulus lusitanus Verboeff
Gonopods visible externally; first pair of legs of male enlarged;
body uniformly gray in color, without stripes (Parajulidae) . 14
Epiproct decurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T eniulus sp.
Epiproct extending into straight spine . . . . . . . . . . . . . . . . . . . . . 15
Peltocoxites of anterior gonopods with flared, serrate calyx (Fig. 6)
.................................P tyoiulus impressus (Say)
Calyx of peltocoxites cupped, smooth (Fig. 5) .................
.............................. Ptyoiulus ectenes (Bollman)
Midbody metatergites with transverse groove; rim of paraprocts
without setae (Paradoxosomatidae) . . . . . Oxidus gracilis (Koch)
Midbody metatergites without transverse groove; rim of para-procts
with one pair of setae . . . . . . . . . . .: . . . . . . . . . . . . . . . 17
Prefemora of legs with ventrodistal spines; gonopod usually bearing
prefemoral process; adults large, robust, color bright yellow-black
or yellow-brown (Xystodesmidae) . . . . . . . . . . . . . . . . . 18
Prefemora of legs without ventrodistal spines; gonopod without
preformal process; adults slender, coloration otherwise . . . . 22
Gonopods with coxal apophysis (Figs. 55-57) ............... 19
Without this character . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Cranial setae present on frons and epicranium in both sexes; gono-pods
with one prefemoral and two tibiotarsal processes (Fig.57);
podosterna present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
................ Pachydesmus crassicutis innrrsus Chamberlin
Cranial setae absent from frons and epicranium in both sexes;
gonopod with or without small prefemoral process; telopodite
broadly curved, falcate in shape (Figs. 55-56); sterna unmodified
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deltotaria lea Hoffman
Telopodite of gonopod with irregularly notched expansion along
proximomedial edge; prefemoral process large, extending beyond
tip of telopodite (Fig. 54); membrane of cyphopod enlarged and
folded, protruding through medial portion of aperture. ........
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C.r oatania catawba Shelley
Without these characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2.1
Prefemoral process short, blunt, never two-thirds length of telopo-dite;
telopodite curved, with medial flange at midlength (Fig. 60);
adults 40-45 mm long . . . . . . . . . . . Sigmoria latior (Briilemann)
8 Marianne E. Filka and Rowland M. Shelley
Prefemoral process of gonopod acicular, approximately two-thirds
length of telepodite; telopodite nearly straight, bent slightly
mediodorsad distally, without flange (Fig. 47); adults 28-32 mm
long . . . . . . . . . . . . . . . . . . . . . . . . .B oraria stricta (BrGlemann)
22. Epiproct broad, truncate; adults gray with orange paranota and
middorsal spots (Platyrhacidae) . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . .A uturus erythropygos (Brandt)
Epiproct subtriangular; adults with essentially uniform coloration
(Polydesmidae) . . . . . . . . . . . . . . . . . . . . . . . . . .... ........ 23
23. Adults with 19 segments; metatergites with four rows of small seti-ferous
tubercles . . . . . . . . . . . . . . . . . Scytonotus granulatuF(Say)
Adults with 20 segments; dorsum without setae and distinct rows of
tubercles ............. Pseudopolydesmus branneri (Bollman)
CLASSIFICATION OF KINGS MOUNTAIN DIPLOPODS
CLASS DIPLOPODA
SUBCLASS PENICILLATA
ORDER POLYXENIDA
Family Polyxenidae
Polyxenus fasciculatus Say
SUBCLASS HELMINTHOMORPHA
ORDER POLYZONIIDA
Family Polyzoniidae
~ol&onium stricturn Shelley
ORDER PLATYDESMIDA
Familv Androenathidae
~nBro~nath&co rticarius Cope
ORDER JULIDA
Family Blaniulidae
Nopoiulus minutus (Brandt)
Family Julidae
Brachyiulus lusitanus Verhoeff
Family Parajulidae
Ptyoiulus ectenes (Bollman)
Ptyoiulus impressus (Say)
Teniulus sp.
ORDER CHORDEUMIDA
Family Cleidogonidae
Cleidogona medialis Shelley
Family Trichopetalidae
Trichopetalum d m (Chamberlin)
Family Striariidae
Striaria sp.
Kings Mountain Milliped Fauna
ORDER SPIROBOLIDA
Family Spirobolidae
Narceus americanus (Beauvois)
ORDER CALLIPODIDA
Family Caspiopetalidae
Abacion magnum (Loomis)
Delophon georgianum Chamberlin
ORDER SPIROSTREPTIDA
Family Cambalidae
~a&b a l aa nnulata (Say)
ORDER POLYDESMIDA
Familv Paradoxosomatidae
0xi;ius graciiis ( ~ o c h )
Family Polydesmidae
Pseudopolydesmus branneri (Bollman)
Scytonotus granulatus (Say)
Family Platyrhacidae
Auturus erythropygos (Brandt)
Family Xystodesmidae
Boraria stricta (Brolemann)
Croatania catawba Shelley
Deltotaria lea Hoffman
Pachydesmus crassicutis incursus Chamberlin
Sigmoria latior (Brolemann)
SPECIES ACCOUNTS
Polyxenidae
Polyxenus fasciculatus Say, 1821
Polyxenus fasciculatus, a small, pale milliped, was recovered from
pine and mixed pine-hardwood litter using Berlese funnels but was absent
from hardwood litter. More adults and juveniles were taken in July than
in October or April. Previous North Carolina records are from Duke
Forest (Brimley 1938; Wray 1967) and the eastern Piedmont in general
(Shelley 1978). The species is known to range from Long Island through
the southeastern and midwestern United States to Texas (Chamberlin
and Hoffman 1958). The presence of P. fasciculatus in the Appalachian
Mountains is questionable, since Chamberlin and Hoffman (1958) re-ported
it absent or very scarce there.
10 Marianne E. Filka and Rowland M. Shelley
Figs. 3-8. 3, Polyzonium striclum, anterior gonopods, cephalic view. 4,
Andrognathus corticarius, head and first six segments, dorsal view. 5, Ptyoiulusec-renes,
anterior gonopods, caudal view, calyx (c) and peltocoxites (p). 6, Ptyoiulus
impressus, anterior gonopods, caudal view, calyx (c) and peltocoxites (p). 7-8
Teniulus sp. 7, anterior gonopods, caudal view. 8, left posterior gonopod, lateral
view. Scale line = 0.1 mm.
Kings Mountain Milliped Fauna
Polyzoniidae
Polyzonium strictum Shelley, 1976
Fig. 3
Yellow adults of P. strictum were taken from beneath the bark of
decaying logs in July and October. A large number of juveniles was ex-tracted
from mixed pine-hardwood Berlese samples in October. In North
Carolina P. strictum ranges from the mountains to the inner Coastal
Plain, and it also occurs in the mountains of Virginia (Shelley 1976a).
Andrognathidae
Andrognathus corticarius Cope, 1968
Fig. 4
This slender, cream-colored diplopod typically occurs beneath the
bark of decaying pine logs (Shelley 1978), a habitat that was examined ex-tensively
in the study area. Only two specimens were encountered, how-ever,
both in July from a single log at a Spencer Mountain site. Cham-berlin
and Hoffman (1958) reported A. corticarius from western Virginia,
southeastern Kentucky, Tennessee, Georgia, and northern Florida.
Gardner (1975) examined material from Graham and Madison counties
in the Appalachian Mountains of North Carolina, and Shelley (1978)
noted that the species was more prevalent in the southern subregion of
the eastern Piedmont than north of the Deep-Cape Fear Rivers.
Locality. Gaston Co.-7.2 km NE Gastonia, along CR 2200,2.2 km
SW jcr. NC Hwy. 7, base of Spencer Mountain, 2 9, 7 July 1976, M..
Filka and W.W. Thomson (NCSM A1032).
Blaniulidae
Nopoiulus minutus (Brandt, 1841)
This narrow brown milliped occurs in habitats similar to those of A.
corticarius. Four immature specimens were encountered, in July and Oc-tober,
but no adults were found. The dearth of specimens in the Kings
Mountain region contrasts markedly with the abundance of the species
farther east in the fall zone region of North Carolina, where it also occurs
in summer and autumn (Shelley 1978). Nopoiulus minutus is widespread
east of the Great Plains, ranging from Illinois, Indiana, and Ohio south
to Georgia (Enghoff and Shelley 1979).
Localities. Cleveland Co.-9.1 km SE Kings Mountain (town), along
CR 2286, 1.6 km S jct. CR 2283,8 July 1976, 1 juv., M. Filka and W.W.
Thomson (NCSM A1973); and 1.9 km SW Kings Mountain (town),
12 Marianne E. Filka and Rowland M. Shelley
along 1-85, jct. NC Hwy. 161, 3 juvs., 18 October 1976, M. Filka and G.
Wicker (NCSM A2197).
Julidae
Brachyiulus lusitanus Verhoeff, 1898
Brachyiulus Iusitanus is easily identified by its characteristic dorsal
color pattern -two pale longitudinal stripes surrounding a narrow black
mid-dorsal line. Introduced from Europe, B. lusitanus has been reported
from developed areas of North America as far south as the "Triangle"
(Raleigh-Durham-Chapel Hill) region of North Carolina, where it was
erroneously reported as B. pusillus (Leach) by Shelley (1978). However,
three females were encountered during this study, all in April, under
debris at a public campsite. The Kings Mountain region thus becomes
the southernmost known locality for B. lusifanus in the New World.
Locality. ~levelknd Co.-1.9 km SE Kings Mountain (town), jct.
1-85 and NC Hwy. 161, 3 0 , 10 April 1977. M. Filka.
Parajulidae
Ptyoiulus ectenes (Bollman, 1888)
Fig. 5
Juvenile and female Ptyoiulus are unidentifiable to species. Those
found associated with males of a single species (all only in October) were
adjudged to be that species and are so shown in Table 3. Those collected
without associated males (all only in July) are tabulated by genus in
Table 3. No juveniles or females were found with males of both species at
a single collecting site. These identification problems may have influ-enced
the apparent seasonal distribution patterns of both species of
Ptyoiulus in the region, although examination of Table 3 reveals similar
July and October patterns for the two. Adults of P. ectenes were most
numerous in October but also occurred in April; immatures were found
only in July and October. These data suggest that P. ectenes reproduces
during fall and spring, and juveniles mature the following summer and
fall. All specimens were collected from deciduous litter. Gonopods of the
31 males were examined but no variation was apparent.
This species was reported from the fall zone region as Ptyoiulus sp.
by Shelley (1978) who declined to assign a specific name in deference to
studies being conducted by the late Dr. Nell B. Causey. The oldest
available specific name is in doubt, but that of Bollman is used tentatively
here since it is one of the earliest names and the first applied to specimens
from North Carolina. However, there is some question as to whether
ectenes is referable to Ptyoiulus, since Bollman (1887) remarked that the
Kings Mountain Milliped Fauna 13
species differed in its "slender body and peculiar form of the male geni-talia,"
which he neither illustrated nor described verbally in his descrip
tion. Unfortunately, the male from the type series is lost, but collections
made by Shelley in and around the type locality - Chapel Hill, Orange
County, North Carolina - have produced male parajulids whose
gonopods are virtually identical to that illustrated in Figure 5. This
suggests that ectenes may be the species under consideration here, but a
final judgment can only result from a comprehensive revision of
Ptyoiulus in which female cyphopods are studied. Wray (1967) may have
been correct in transferring ectenes to Aniulus, and this combination may
be a senior subjective synonym of A. orientalis Causey, the only other
parajulid known from the "Triangle" region of the state.
Ptyoiulus impressus (Say, 182 1)
Fig. 6
Adults of P. impressus were abundant in October and absent in April
and July; juveniles were taken only in July and October. Thus, P. im-pressus
appears to have a slightly different life history from that of P. ec-tenes,
with summer growth and maturation preceding fall reproduction.
Both species are uniformly gray and both were found in deciduous forest
litter. Adult P. ectenes are slightly smaller and less robust than adult P.
impressus, although this difference can be misleading and should not be
the sole criterion for identification. The most reliable character is the
configuration of the calyx of the peltocoxites of the anterior gonopod
(Figs. 5-6, c, p,), which is flared and serrate distally in P. impressus and
cupped and smooth in P. ectenes. As with its congener, the gonopods of
P. impressus were essentially uniform.
Ptyoiulm impressus ranges from the northeastern United States west
to Indiana and south to western North Carolina and Kentucky (Cham-berlin
and Hoffman 1958). Shelley (1978) deleted this species from the
eastern Piedmont fauna, stating that it was known definitely only from
themountains and western Piedmont. The Kings Mountain region is the
easternmost authentic locality in North Carolina.
Teniulus sp.
Figs. 7-8
This uniformly gray species is similar in appearance to both species
of Ptyoiulus, but is distinguished by the decurved epiproct. Adults were
collected in October from moist deciduous leaf litter in association with
both species of Ptyoiulus. No juveniles were found.
The genus currently contains only two species, T. parvior and T.
setosior, both described by Chamberlin (1951) from Gatlinburg, Seyier
County, Tennessee. Gatlinburg is about 200 km west-northwest of the
14 Marianne E. Filka and Rowland M. Shelley
Kings Mountain region, and such wide geographic separation suggests
that the forms in the two areas are not conspecific. If true, the Kings
Mountain species is undescribed. However, drawings by Chamberlin
(1951) accompanying the descriptions of the Tennessee species are un-clear
and were prepared from different views, which prevents close com-parisons.
Examination of the type specimens of both species by Shelley
revealed that males and/or gonopods were absent. These two species may
be synonymous, but adult males are needed before their identities can be
determined and an accurate statement can be made on the status of the
Kings Mountain form.
Locality. Gaston Co.-9.9 km S Bessemer City, along CR 1112, 0.3
km E ict. CR 1125. 6. SF. 17 October 1976. M. Filka and G. Wicker
Cleidogonidae
Cleidogona medialis Shelley, 1976
Figs. 9-10
The Kings Mountain region is the second known locality for this
light brown chordeumid, whose range is now extended some 117 km
from Blowing Rock, Watauga County, North Carolina (Shelley 1976b).
The single male and female taken during this study and all those from the
type locality were collected in October, suggesting autumnal maturation.
Juveniles would therefore be expected in the summer; although none
were encountered in July, they may be present in late August or early
September.
The medial processes of the gonopods of the Kings Mountain speci-men
were more jagged than those of the holotype (Shelley 1976b, Fig.
lo), which conforms to known variation in the species. A single oversized
telopodite variant was reported in one male paratype, but those of the
Kings Mountain male were as illustrated for the holotype. No other
gonopodal variations were observed. Two additional records are cited
below from material that has recently become available.
Localities. Gaston Co.-9.9 km SE Bessemer City, along CR 1126,
0.8 km SW jct. CR 1113, P, 16 October 1976, M. Filka and G. Wicker
(NCSM A2770); and 9.9 km SW Bessemer City, along CR 1104, 1.3 km
W jct. CR 1115, r?, 17 October 1976, M. Filka and G. Wicker (NCSM
A2771). Davidson Co.-Boone's Cave State Park, 8, 6 November 1976,
R.M. Shelley (NCSM A1434). Watauga Co.-16 km NE Deep Gap, d,
9 , 17 October 1965, 1. & W. Ivie (AMNH).
Kings Mountain Milliped Fauna
Figs. 9-19. 9-1 1, Chordeumida. 9-10, Cteidogona medialis. 9, anterior gonopods,
cephalic view, sternum broken in dissection. 10, left anterior gonopod, lateral
view. 11, Striaria sp., epiproct, dorsal view. 12-19 Narcnrs americanus. 12, an-terior
gonopods, cephalic view. 13, left posterior gonopod, cephalic view. 14.15,
distal portions of left posterior gonopods, cephalic views, showing variation in
prefemoral endite (pe). 16, coxae and lobes of legs 3-7 of male, ventral view. 17-
19, coxae and prefemora of left third legs of females, cephalic views, showing
variation in coxal lobes. Scale line = 0.1 mm.
Marianne E. Filka and Rowland M. Shelley
Trichopetalidae
Trichopetalum dux (Chamberlin, 1940)
The specific identification of this milliped is tentative and based
solely on previous North Carolina records. Only one female was collec-ted,
discovered in berlesate from a deciduous litter sample taken in Gas-ton
County in April. This is the only species of Trichopetalum known
from North Carolina, where it was previously reported from Duke
Forest (Chamberlin 1940a; Wray 1967) and Chatham County (Shelley
1978). Positive identification of the Kings Mountain species awaits the
collection of males.
Locality. Caston Co.-8 km NE Gastonia, base of Spencer Moun-tain,
along CR 2200, 1.7 km SWjct. CR 2003, 9 , 10 April 1977, M. Filka
(NCSM A2185).
Striariidae
Striaria sp.
Fig. 11
Striaria is readily distinguished from the other chordeumids by its
enlarged collum, crested segments, and trilobed epiproct. One adult
female and two juveniles were taken in Gaston County in October and
April, respectively. The adult female and one juvenile were brown with a
pale white collum, while the other juvenile was uniformly brown. Three
species of Striaria are known from North Carolina: two with a white
collum - S. zygoleuca Hoffman, from Highlands, Macon County (Hoff-man
1950; Wray 1967), and an undescribed form from High Falls, Moore
County (Shelley 1978) - and one with a brown collum, S. causeyae
Chamberlin, from eight counties in the eastern Piedmont (Shelley 1978).
The presence of differently colored collums on the Gaston County
material suggests the presence of at least two species, but again the ab-sence
of males precludes final determinations. For the purposes of this
report, therefore, only one species is considered.
Localities. Caston Co.--4.0 km S Bessemer City, along CR 1125,0.2
km S jct. U.S. Hwy. 74.29, 0, 17 October 1976, M. Filka and G. Wicker
(NCSM A2215); juv., 9 April 1977, M. Filka (NCSM A2219); and 6.4 km
SE Bessemer City, along CR 1103, jct. CR 11 12, juv., 9 April 1977, M.
Filka (NCSM A2202).
Spirobolidae
Narceus americanus (Beauvois, 1805)
Figs. 12-19, Tables 1-2
Body coloration of N. americanus is dark brown with the head, legs,
and body segments bordered in red. As indicated in Table 3, this species
is common in the Kings Mountain region, and adults and juveniles were
Table I. Comparison ofthe number ofsegments, clypcal and labral sctae, ocelli, and formula value for the Kings Mountain spirobolid (KM),
Norem onnuloris (Nan), and N. omerieonus (Nam). Data for Narcem f ~ o mK a ton (1960). d = difference from the Kings Mouniain
spirobolid.
K M 1 4 6 9 1 22 49.4
Nan 1 5 22 66 77 60 37 5 4 277 55.1 5.7
Nam 3 7 15 16 28 19 18 8 2 116 51.1 1.7
Number afclypml reroe -F.
-4 -5 -6 -7 -8 -9 -10 -11 -12 -13 -14 -I5 -16 -N -F -d 2J
KM 1 7 5 8 - 1 22 11.1
Nan 4 1 21 56 203 84 26 5 2 402 8.1 3.0 z 0
Nam 1 32 35 38 21 5 3 I I 1 138 9.7 1.4 G 3
Number oflabroiselae - 5.
KM
-.
1 4 9 4 2 2 22 16.4 -
Nan 2 1 - 13 19 136 217 125 27 5 2 2 549 13.0 3.4
Nam 2 7 18 22 24 19 15 6 9 1 - I 1 125 16.3 . 0.1 3
Number o f o d i 2
KM I - 3 2 2 3 4 1 2 I 22 39.6
Nan I - 2 3 5 10 21 38 34 43 43 36 35 24 28 I5 15 10 4 3 1 2 - - - I 374 42.5
Nam
2.9
I 2 9 7 1 7 1 1 1 2 1 4 1 0 9 7 1 7 1 1 8 1 0 3 5 - 1 2 2 - I 1 160 . 43.7 4.1
Fornulo wlueB
18 Marianne E. Filka and Rowland M. Shelley
collected from a diverse array of habitats on each trip. Continuous
breeding and maturation throughout the year is suggested by these data.
Two species of Narcm occur in North Carolina; N. americanus,
known to range throughout the southeastern United States, and N.
annularis (Rafinesque), known from the northeastern and midwestern
United States (Chamberlin and Hoffman 1958). Both are reported from
the mountains (Keeton 1960) and eastern Piedmont (Shelley 1978) of
North Carolina.
In his monograph on the Spirobolidae, Keeton (1960) distinguished
between N. annularis and N. americanus by a formula computing values
based on somatic features (set footnote Table 1 for explanation) and by
comparison of gonopodal characters. He found that differences were dif-ficult
to define due to overlap of characters. Consequently, identifica-tions
are difficult and distinctions between the species are vague, facts
corroborated by the Kings Mountain material.
Thirty-four adults were collected but only 22 of these, 16 males and 6
females, were in sufficiently good condition for detailed study. As shown
in Tables 1-2, the number of segments, clypeal setae, and labral setae of
the Kings Mountain spirobolid are closer to the values for N. americanus;
but the mean number of ocelli is closer to that for N. annularis. Mean
length and width, the lengthlwidth ratio, and the formula value, also are
nearer the figures for N. americanus.
The distal prefemoral endite of the posterior gonopod, normally
rounded in N. annularis and acute in N. americanus, is rounded in 72% of
the specimens (Figs. 13-14, pe) and acute in the rest (Fig. 15, pe).
Likewise, the cephalic groove on the coxal lobes of the third pair of legs
of males, a characteristic of N. americanus, is absent from all Kings
Mountain males (Fig. 16). Furthermore, the third coxal lobe of adult
females, enlarged in N. americanus but only slightly produced ventrad in
N. mmularis, conforms to the latter condition in all but one specimen
(Figs. 17-19). The Kings Mountain spirobolid therefore could be iden-tified
as either species of Narcm depending upon the character used, and
the question becomes one of the relative importance of the characters.
Although gonopods are the most important taxonomic character in
the Diplopoda, many genera show few specific gonopodal differences
whereas there is wide variation in body forms. This appears to be the
situation in Narceus, and the few gonopodal similarities between the
Kings Mountain spirobolid and N. annularis do not seem to outweigh the
close agreement of nearly all the somatic features with those of N.
americanus. Thus, the Kings Mountain spirobolid is identified as N.
americanus. The great variability of the somatic features of Narceus, as
demonstrated by Keeton (1960) and our tables, indicates a need for
reassessment of the status of the two nominal eastern species. Such a
study might show them to be conspecific.
Table 2. Comparison of length, width, and length/width ratio for the Kings Mountain spirobolid (KM), Narceur annulark (Nan) and N.
americanus (Nam). Data for Narceus from Keeton (1960). d = difference from Kings Mountain spirobolid; C x = combined mean
for both sexes; Cd = combined difference for both sexes.
T.enclh (rml
Nan c+ I 2 13 22 26 22 13 13 4 3 1 I
0 1 4 16 19 21 23 18 13 9 5 2 I 1 5:
Nam or 2 2 4 6 4 6 1 3 2 4 I I 37 78.5
F 2 4 . 4 5 . 5 - 5 8.--9 9 6 6 - 2 3.4 2.5 %
65 78.5 78S 0.2
Width (mm) ' 3
0
Nan ol 2 15 42 40 36 9 3 1
0 2 7 19 37 29 24 18 6 2 2
e;'
Nam d 2 5 5 3 8 6 1 4 1 7 I 1
6 2 8.1
c
0 I 1 3 4 9 8 9 9 1 0 6 0.0 m
Ct
Lengthlwidth ratio ('3)
7.0 7.5 8.0 - - _ -8.5 -9.0 9-.5 1-0.0 -10.5 1_1.0 _11.5 _12.0 _12.5 _13.0 -13.5 _14.0 _14.5 _15.0 -N -X C-X -d C-d
Nan ol
0
Nam w 1 5 9 1 0 1 0 5 2 1 43 10.1 0.7
P 1 5 1 3 I 2 1 6 9 8 2 66 9.8 'O" 0.3 O3 w
\O
20 Marianne E. Filka and Rowland M. Shelley
Abacion magnum (Loomis, 1943)
Figs. 20-24
Abacion magnum is a crested diplopod, brown with a light middorsal
stripe. It was one of the few species found in drier parts of deciduous,
pine, and mixed leaf titter. Juveniles of the two callipodids, A. magnum
and D. georgianum, could not be identified to species, but all were found
with adult males of a single species and, as with Ptyoiulus, were identified
as such. Adults of A. magnum were taken during all three months, while
juveniles were collected only in July and October. This implies that
reproduction and maturation occur throughout the year. The coxal
processes of the gonopods of the eastern Piedmont specimens varied in
degree of apical serration and configuration of the midlength angulation
(Shelley 1978). These structures were found to vary similarly in the Kings
Mountain specimens (Figs. 21-24). No other gonopodal variations were
detected. This species has been collected in Macon and Transylvania
counties in the North Carolina mountains (Hoffman 1950) and in eight
counties of the eastern Piedmont (Shelley 1978).
,
Delophon georgianum Chamberlin, 1943
Figs. 25-26
This callipodid is similar in coloration to A. magnum but is smaller
and differs in the structure of the gonopod. Abacion magnum has a
flagellum and a serrate coxal process lateral to the telopodite (Fig. 20, fl.
cp). Delophon georgianum lacks the flagellum, and its coxal process
ensheaths the stalk of the telopodite (Figs. 25-26, cp) (Shelley 1979a). Lit-tle
gonopodal variation was found in this study. Like A. magnum, adults
of D. georgianum were taken on all three trips, but only one juvenile was
encountered, in April. Delophon georgianum was typically found in
moister habitats than Abacion.
This species has been previously reported from the mountains of
North Carolina as D. carolinum Hoffman (Hoffman 1950; Chamberlin
and Hoffman 1958; Wray 1967). However, Shelley (1979a) concluded
that this binomial was a synonym of D. georgianum. The Kings Moun-tain
population is disjunct from that occurring in the Appalachians, and
no specimens have ever been taken in the intervening lowlands. Shelley
speculated that the Kings Mountain population might he a Pleistocene
relict that has survived due to a slightly cooler microclimate afforded by
the peaks and coves of the area. Hardin and Cooper (1967) concluded
that this was the explanation for the occurrence of several disjunct pop-ulations
of montane plants, most notably Tsuga canadensis L. and Pinus
strobus L., in the Piedmont.
Kings Mountain Milliped Fauna 21
Figs. 20-27. 20-26, Callipodida. 20-24, Abacion magnum. 20, left gonopod, caudal
view, coxal process (cp), flagellum (fl). 21-24, coxal processes, lateral views, of
four specimens from the Kings Mountain Region. 25-26, Delophon georgianum.
25, left gonopod, lateral view, coxal process (sheath) (cp). 26, left gonopod,
caudal view, coxal process indicated. 27, Cambala annulafa, left posterior
gonopod, lateral view. Scale line = 0.1 mm.
22 Marianne E. Filka and Rowland M. Shelley
Cambalidae
Cambala annulata (Say, 1821)
Fig. 27
Shelley (1978) reported that C. annulata seemed to prefer cooler tem-peratures,
and this was apparent in the Kings Mountain region where the
dark purple adults were abundant in both April and October. Only one
adult, a female, was collected in July. Juveniles were taken in July and
October but not in April. Hoffman (1958) indicated that individuals of
this species were usually found grouped together, but in the Kings Moun-tain
region this was true only of females; adult males were always found
alone. All stages were collected from moist humus. Hoffman found no
gonopodal variation in material from high elevations, and Shelley
(1979b) noted homogeneity in the gonopods of C. annulata throughout
its range. This was evident in the Kings Mountain material, as the struc-tures
were virtually uniform. Cambala annulata has been reported from
the northeastern and central subregions of eastern Piedmont North
Carolina (Shelley 1978), and its distribution in the Appalachian Moun-tains
was illustrated by Hoffman (1958).
Paradoxosomatidae
Oxidus gracilis (Koch, 1847)
Causey (1943) reported nearly year around oviposition by 0. gracilis
under favorable conditions in a Durham County greenhouse, and Shelley
(1978) collected fifth instarjuveniles (adults rre the seventh instar) in Oc-tober
from William B. Umstead State Park in the eastern Piedmont. The
preponderance of juveniles in October and April and of adults. in July
and October in the Kings Mountain region suggests that maturation oc-curs
in the fall and spring and breeding in the late summer and early fall.
White juveniles often populated several square meters of deciduous leaf
litter, and shiny black adults also were common. Oxidus gracilis is nearly
worldwide in distribution, and was introduced into the United States
from the East Indies via imported soil in greenhouses (Causey 1943).
Polydesmidae
Pseudopolydesmus branneri (Bollman, 1887)
Figs. 28-40
Pseudopolydesmus branneri is the sole representative of its genus in
the Kings Mountain region; neither P. collinus Hoffman nor P. serratus
Kings Mountain Milliped Fauna 23
(Say), both of which occur in the eastern Piedmont (Shelley 1978), were
encountered during this study. This species has been previously reported
from Rutherford, Wilkes and Alexander counties in the western Pied-mont,
as well as the mountains and eastern Piedmont (Shelley 1978), so
its occurrence in the Kings Mountain region was expected. Juveniles and
adults were collected in April and October, but none were found in July.
Adults were dull reddish brown with light brown paranota, similar to
Richmond County specimens (Shelley 1978). Material from the two areas
was also similar in length.
Past discussion of gonopodal variation in the Polydesmidae has been
hampered by the absence of a standardized nomenclature for the spines,
branches, and other projections of the posterior faces of the telopodites.
Hoffman (1974) devised a labeling system based upon orientation of
these processes, with letters "m" and "e" designating mesial and ectal
position, respectively, and numbers indicating position relative to the
base of the telopodite, the most proximal designated by 1. Thus, in P.
branneri the four mesial processes are labeled ml, m2, m3, and m4; the
four ectal processes are el, e2 + 3 (indicating that they share a common
pedicel), and e4 (Figs. 28-29). Examination of the left gonopods of 5 1
specimens collected from the Kings Mountain region revealed consid-erable
gonopodal variation.
The only evident variation in the mesial processes involved suppres-sion
and division of m4. Nearly three-fourths of the individuals examined
had a normal m4 lobe with a projecting setaceous shoulder (Figs. 28,30,
34); the remaining individuals had a reduced or vestigial m4, with a
shoulder lacking setae (Figs. 31-33). In 75% of the specimens, m4 con-sisted
of a large lobe contiguous with a smaller shoulder (Figs. 28,30-33).
This lobe was divided into two separate processes in the remaining
specimens (Fig. 34). The other mesial processes, ml-m3, were virtually
uniform, although one individual lacked m2 (Fig. 35).
The most variable ectal process, el, displayed four configurations;
spiniform (Figs. 28, 36) in 31% of the males; reduced (Fig. 37) in 29%;
vestigial (Fig. 38) in 18%; and absent (Fig. 39) in 18%. Two individuals
(4% of the males) carried a vestigial secondary spine distal to a reduced el
spine (Fig. 40). The other ectal processes, e2+3 and e4, were uniform.
Terminal macrosetae of the telopodite varied in abundance and dis-tribution.
Thirty-one percent of the males had numerous macrosetae oc-curing
from just distal to m4 to the telopodite tip (Figs. 28-30, 33-34), and
fifty-one percent carried fewer macrosetae distributed from e4 to the tip
(Fig. 31). Most remaining individuals (14%) possessed very few mac-rosetae,
occurring only apically on the telopodite (Fig. 35). Two in-dividuals
(4%) lacked terminal macrosetae (Fig. 32).
Hoffman (1974) reported that m2, m3, e2, and e3 were the most
variable processes in P. branneri, but these were found to be the most
stable in the Kings Mountain population, where most of the variation in-volved
m4, el, and the terminal macrosetae. Many combinations of these
24 Marianne E. Filka and Rowland M. Shelley
Figs. 28-41. Polydesmidae. 28-40, Pseudopo/ydesmus branneri. 28, left gonopod,
medial view. 29, the same, lateral view, medial processes (ml-m4), ectal processes
(el-e4), endomerite (end), terminal macrosetae (tm). 30-34, distal ends of
telopodites, medial views, showing variation of m4 and terminal macrosetae. 35,
distal half of telopodite, medial view, showing absence of process m2. 36-40, en-domerite
regions of telopodites, medial views, showing variation of process el.
41, Scytonorur granulalur, left gonopod, medial view. Scale line = 0.1 mm.
Kings Mountain Milliped Fauna 25
gonopodal variants occurred, and there was no correlation between
them. The length of the patch of terminal macrosetae and the configura-tion
of m4 and el varied independently, and seem to be controlled by dif-ferent
genes.
These findings greatly expand current knowledge of variation for the
species and illustrate the degree of variability that may occur within a
local population. To Hoffman's (1974) characterization of P. branneri
may now be added the occasional appearance of a new process, the
secondary el spine, the division of a single process into separate compo-nents,
m4 lobe and shoulder, and the occasional loss of all terminal
macrosetae.
Hoffman (1974) described P. collinus as differing from P. branneri in
the absence of m3 and either the absence or vestigial condition of el. In
the Kings Mountain specimens, el and/or m3 were present on all
specimens, although el varied considerably in size. Consequently, only
one species, P. branneri, is represented by this material.
Scytonotus granulatus (Say, 1821)
Fig. 41
This species was rare in the Kings Mountain region. Isolated brown
adults were found in April and October, and white juveniles were taken
in July. Both were found in moist humus. The widespread occurrence of
the species in western North Carolina and several other states was noted
by Hoffman (1962), and Shelley (1978) reported additional localities in
eastern Piedmont North Carolina.
Platyrhacidae
Autuw erythropygos (Brandt, 1841)
Figs. 42-46
Adults of A. erythropygos exhibit striking body coloration, with each
blue-gray metatergite bearing a bright orange middorsal spot and orange
paranota. Juveniles, though lighter, have a similar pattern. All stages
kcre collected from under bark of decaying d e c ~ d ~ o u ~ lino h~usm,u s un-der
logs, or in associaled bark litter. Both adults and juveniles were most
abundant in October. but the svecies was auite common in Aoril and
July.
Flattened, round molting chambers, built under the bark of logs in-habited
by A. erythropygos, were observed on each collecting trip (Figs.
45-46). They are constructed of cemented wood particles and provide
protection from desiccation and predation during intermolts. The dimen-sions
were proportional to the inhabitant's size, the largest being 20-22
mm diameter. An adult or juvenile accompanied by cast exuvium was
26 Marianne E. Filka and Rowland M. Shelley
seen in each chamber. The exoskeleton of newly molted individuals was
whitish and incompletely sclerotized.
This species was unknown from North Carolina until reported from
Northampton County by Shelley (1978), who listed it as A. georgianus
Chamberlin, now considered a junior synonym.
Figs. 42-54. 42-46, Aurum eryrhropygos. 42, left gonopod, medial view. 43, the
same, lateral view. 44, distal end of telopodite, cephalic view. 45-46, molting
chamber. 45, side view in situ on log, bark lifted. 46, top view. 47-53, Boraria
stricra. 47, left gonopod, medial view. 48-51, distal halves of prefemoral processes
of left gonopods, medial views. 52-53, molting chamber. 52, side view in situ.
attached to plant roots. 53, top view. 54, Croarania camwba, telopodite of left
gonopod, medial view. Scale line = 0.1 mm.
Kings Mountain Milliped Fauna
Xystodesmidae
Boraria stricta (Brolemann, 1896)
Figs. 47-53
The color of 8. stricta, black with yellow paranota, is typical of most
xystodesmid species in the Kings Mountain region. Adults were most
abundant in April, and juveniles were common in October and April.
Large colonies were discovered in wet mud-clay soils lining the banks of
streams throughout the region. Individuals often were captured in tun-nels
beneath shallow layers of detritus. Round molting chambers, simi-lar
to those described for this species by Hoffman (1965), were observed
in the vertical shafts of several tunnels in April (Figs. 52-53). Each cham-ber
was formed of clay attached to exposed plant roots, and inhabited by
a newly molted milliped with its cast exuvium. As can be seen by compar-ing
illustrations (Figs. 45-46, 52-53), the molting chamber of B. stricta is
spherical with an apical "chimney" and is attached at its base, whereas
that of A. erythropygos is round, flattened in a vertical plane, and at-tached
at both ends. These distinctions reflect the different biotopes in-habited
by the species.
Hoffman (1965) reported that the known range of B. stricta
coincided closely with the southern section of the Blue Ridge physi-ographic
province and predicted only slight extensions at the northern
and southern extremities. Discovery of the species in the Kings Mountain
region represents an extension of slightly less than 64 km east into the
Piedmont Plateau. The species was not reported by Shelley (1978) from
the fall zone region, and extensive investigations in the Uwharrie Moun-tains
also have failed to produce it. Hence, the Kings Mountain popula-tion
is the easternmost known and is probably peripheral. Specimens also
have been collected from a number of other Piedmont localities in the
past eight years by Shelley, and since the Piedmont is geologically and
climatically distinct from the southern Appalachians, material from the
entire range was examined to determine if recognition of geographic
races was warranted. Hoffman (1965) noted the homogeneity of B. stricta
gonopods, with only slight differences detected. The lobes of the distal
suhhastate end of the telopodite varied in size relative to each other, and
the degree of bending at midlength of the telopodite and apically on the
prefemoral process varied. These differences were scattered and inconsis-tent,
not conforming to any geographic pattern. The prefemoral proc-cesses
of seven percent of males in the Kings Mountain population
(including material from York County, South Carolina), however, were
apically bifurcate (Figs. 47-48, 51), a condition never before reported for
either the genus or species. These bifurcate males were intermixed with
normal individuals, although there were differences in the apical pre-femoral
bend of the latter (Figs. 49.50).
In summary, no significant geographical variation was observed in
Marianne E. Filka and Rowland M. Shelley 28
B. stricta, and the homogeneity noted previously by Hoffman (1965) also
applies to Piedmont populations. The bifurcate prefernoral process is
new, however, and its occurrence solely in the Kings Mountain popula-tion
may represent a peripheral population effect. This occurs in too
small a sample of the Kings Mountain population, however, to justify
taxonomic recognition. The known range of B. stricta is expanded con-siderably
to include Gaston and Cleveland counties, North Carolina, and
York (Kings Mountain State Park) and Spartanburg (Croft State Park)
counties. South Carolina.
Croatania catawba Shelley, 1977
Fig. 54
Croatania catawba Shelley, 1977:306, Figs. 1-2, 7, 11-12, 16.
Croatania catawba was one of the few millipeds encountered pri-marily
in July; only three adults were found in April and October. The
preference of species of Croatania for hot, dry conditions was discussed
by Shelley (1977), who also presented a description of the habitat at the
type locality in Cleveland County. Individuals collected during the
present study, however, were taken from cool, moist seepage areas under
deciduous leaf piles and from under large, decaying deciduous logs. As
reported by Shelley (1977), adults were typically black with lemon yellow
paranota and a variable yellow stripe along the anterior edge of the
collum. One female displayed an orange tinted collum stripe similar to
that reported by Shelley for two Union County, South Carolina
specimens. No significant gonopodal variation was discerned.
Shelley (1977) suggested that the distribution of C. catawba in North
Carolina might be associated with the Kings Mountain range, which ex-tends
northeastward through a series of hills and ridges to Anderson
Mountain in Catawba County. Except for one Lincoln County specimen
taken in 1952, however, the milliped has not been collected in North
Carolina outside the contiguous ridge portion of the range in Cleveland
and Gaston counties. Croatania catawba is thus essentially restricted to
this small area in North Carolina, and therefore is considered to be a
species of special concern in the state, as defined by Cooper et al. (1977).
Localities. Cleveland Co.-9.3 km S Kings Mountain (town), along
CR 2245, 0.2 km N jct. CR 2288, 9 8, 5 0 , 16 September 1975, R.M.
Shelley and J.C. Clamp (NCSM A450) TYPE LOCALITY; 1.9 km SW
Kings Mountain (town), along 1-85, jct. NC Hwy. 161, E , 10 April 1977,
M. Filka (NCSM A1040), 3, E,7 July 1976, M. Filka and W.W.Thom-son
(NCSM A1048), and 5 o', 9 , 6 juvs., 10 July 1976, M. Filka and
W.W. Thomson (NCSM A1049). Gaston Co.-7.7 km SW Gastonia,
along CR 1131,0.2 km NWjct. CR 1133, 0.9 July 1976, M. Filka and
W.W. Thomson (NCSM A1340); and 7.2 km S Bessemer City, along CR
1125, jct. CR 1106, 2 , 16 October 1976, M. Filka and G. Wicker (NCSM
A1418).
Marianne E. Filka and Rowland M. Shelley
Figs. 55-64. 55-56, Deltotaria lea, left gonopods, ventrolateral views, coxal
apophysis (ca). 57-59, Pachydesmus crassinrlir incursus. 57, telopodite of left
gonopod, lateral view, coxal apophysis (ca), primary tibiotarsus (ptt), second
tibiotarsus (stt). 58-59, distal ends of secondary tibiotarsi, lateral views. 60-64,
Sigmoria latior. 60, telopodite of left gonopod, medial view. 61-62, distal ends of
telopodites, medial views. 63-64, prefemoral processes, medial views. Scale line =
0.1 mm.
known definitely only from the Kings Mountain region (Shelley and
Filka 1979). It is approximately 7 cm long, and dusky brown with yellow
paranota. As with C. catawba, most specimens were found in July, con-centrated
in wet spots such as seepage areas. Shelley and Filka presented
illustrations of gonopodal variation and showed changes in body dimen-sions
that occur with latitude. Individuals of both sexes are larger and
Kings Mountain Milliped Fauna 3 1
more brightly colored in the Kings Mountain region than farther south in
South Carolina, probably a reflection of more favorable environmental
conditions in the former area.
Gonopod comparisons revealed variation in primary and secondary
tibiotarsi (Figs. 57-59, ptt, stt). As reported by Shelley and Filka, the sub-terminal
process of the secondary tibiotarsus was pointed, blunt, or ab-sent.
Since, in North Carolina, P.c. incursus is apparently restricted to the
Kings Mountain region, it is considered to be endangered within the
state, as defined in Cooper et al. (1977).
Localities. Cleveland Co.-6.6 km SW Kings Mountain (town) along
CR 2245 at Dixon Branch Creek, 0.8 km NW jct. CR 2283,2 3, 2 9, 1
juv., 16 August 1975, R.M. Shelley and J.C. Clamp (NCSM A537); 9.3
km S Kings Mountain (town), along NC Hwy. 245, 0.2 km N jct. CR
2288,3 P, 16August 1975, R.M. Shelley and J.C. Clamp (NCSM A541);
9.1 km SW Kings Mountain (town), along CR 2283, 1.3 km NE jct. NC
Hwy. 216, 8, 8 July 1976, M. Filka and W.W. Thomson (NCSM
A1060); and 4.8 km S Kings Mountain (town), along CR 2289, 1.0 km W
NC Hwy. 161, 2 0 , 18 October 1976, M. Filka and G. Wicker (NCSM
A2239). Gaston Co. -8.5 km SW Gastonia, along CR 1122, 1.4 km w jct.
CR 1131, along Crowders Creek, 2 ?: 2 9, 16 August 1975, R.M. Shelley
and J.C. Clamp (NCSM A547); 6.4 km SW Gastonia, along CR 1126,0.8
km S jct. CR 1113, 9, 16 August 1975, R.M. Shelley and J.C. Clamp
(NCSM A549); 7.7 km SW Gastonia, along CR 1131,0.2 km NW jct. CR
1133, 9 , 8 July 1976, M. Filka and W.W. Thomson (NCSM A1091); and
1.9 km W Gastonia, along CR 1106, 2.4 km E jct. CR 1236, d, 16 Oc-tober
1976, M. Filka and G. Wicker (NCSM A2255).
Sigmoria latior (Bralemann)
Figs. 60-64
This was the most common xystodesmid in the region of study.
Adults and juveniles were discovered beneath decidous leaf litter and on
open substrate in July, but only four adults were taken in both April and
October. In North Carolina the species ranges from the northwestern
mountains to the eastern Piedmont, and intergrades of the three sub-species
were reported from McDowell County eastward to Scotland and
Hoke counties (Shelley 1976c), an area which includes the Kings Moun-tain
region.
Shelley (1976~)n oted that all specimens available from south of the
Catawba and Deep-Cape Fear rivers, including intergrades of S. 1. lazior
(Brijlemann) X S. I. hoffmani Shelley, had stripes along the caudal edges
of the metaterga. The nominate subspecies, occurring north of these
rivers, lacked stripes. During our study, however, specimens of both
color patterns were discovered. At Spencer Mountain they exhibited the
striped pattern, whereas around Kings-Crowders ridges the metaterga
32 Marianne E. Filka and Rowland M. Shelley
were black and without stripes. In both areas the stripe and/or paranotal
color was yellow and did not vary through shades of orange-red, as
reported by Shelley for the nominate subspecies and intergrades.
Since this is the first report of unstriped specimens in the zone of in-tergradation,
the gonopods of 18 striped and 13 unstriped males were ex-amined
for possible differences. Depth of the flange and broadness of the
distal curvature of the telopodite varied, but the flange always extended
below the tip of the telopodite (Fig. 60). The subterminal tooth varied in
prominence and was double in two individuals (Figs. 60-62), and the
prefemoral process ranged from simple to bifurcate with variation in the
relative lengths of the components, although the vertical branch was
always larger (Figs. 63-64). All are typical intergrade variations and do
not correlate with either color pattern. Thus, the solid black metatergal
color is interpreted to represent the nominate subspecies trait, just as
some intergrade gonopods more closely resemble those of one subspecies
than the other two.
DISCUSSION
SEASONAVLA RIATIONOF THE FAUNA
Although the Kings Mountain region was not sampled quanti-tatively
and only limited conclusions can be drawn concerning numbers
of species present in each season, the area was studied with sufficient in-tensity
to reflect general trends in seasonal difference~ (Table 3). The
overall abundance of millipeds increased from April to October, with
only two species, Narceus americanus and Auturus erythropygos, present
as both adults and juveniles in all three months. Adults of other species
varied seasonally, with juveniles present simultaneously or in other
months. The more common species that particularly exemplify these
seasonal variations are discussed by month below.
Diplopods were least abundant in April. Adults of Pseudopoly-desmus
branneri and Boraria stricta, and juveniles of Oxidus gracilis,
dominated the fauna, while adults and juveniles of N. americanus, and
adults of Cmnbala annulata and A. erythropygos were moderately abun-dant.
Adults of Ptyoiulus ectenes, Delophon georgianum, and Deltotaria
lea were less common, and juveniles of these species were absent or nearly
so. Only a few specimens of the remaining species were found. Adults of
D. georgianum, P. branneri, B. stricta, and D. lea, and juveniles of Stri-aria
sp., 0 . gracilis, and B. stricta were more numerous in April than in
any other month. Two species, Brachyiulus lusitanus and Tnchopetalum
dux, were collected only in April.
A different group of diplopods dominated the fauna in July. Narceus
americanus, Abacion magnum, 0. gracilis, A. erythropygos, Pachydesmus
crassicutis incursus, and Sigmoria latior were the prevalent adult forms,
while Polyxenus fasciculatur. Ptyoiulus sp., and N. americanus were com-mon
in immature stages. Intermediate numbers of A. erythropygos and
Kings Mountain Milliped Fauna
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Marianne E. Filka and Rowland M. Shelley
S. Iatior juveniles also were present, and the remaining species were
represented only by scattered individuals. Five species, P, fasciculatus. A.
magnum, Croatania catawba, P.c. incursus, and S. latior were more abun-dant
as both adults and juveniles during July than at any other time. Im-matures
of Nopoiulus minutus and Scytonotus granulatus also were most
numerous in July, although juveniles of the former were taken in October
as well. Species more common as adults in July than in April included
Polyzonium strictum, N. americanus, and 0. gracilis. Species less common
as adults during July than in October or April included C. annulata and
B. stricta. Three species - P. ectenes, P. branneri, and D. lea -were ab-sent
from the July collections. Andrognathus corticarius was collected
only in July.
The greatest abundance and diversity of species occurred in October.
Those most abundant as adults were Ptyoiulus ectenes, P. impressus. C.
annulata, P. branneri, and A. erythropygos. Those most common in im-mature
stages were P. ectenes, 0. gracilis, P. strictum, N. americanus, and
A. erythropygos. Morejuveniles of the last three species were encountered
in October than in either of the other months. The first two species were
moderately abundant as either adults or juveniles. Both Teniulus sp. and
Cleidogona medialis were found only in October and as adults. Millipeds
less common in the adult stage in October than in July included N.
amencanus, A. magnum, 0. gracilis, C. catawba, P.c. incursus, and S.
latior.
The species/genus ratio (S/G) for all three months (1.04) was essen-tially
unity (April and July S/G = 1.00, October S/G = 1.05), with the
slightly higher fraction of October reflecting the presence of both species
of Ptyoiulus. Seasonal changes in the faunal composition ratios (or-ders/
families/genera/species, O/F/G/S) from April to October were
more significant than changes in the S/G ratios. One more order, the
same number of families, and one less genus and species occurred in July
(8/12/16/16) than in April (7/12/17/17). The same number of orders,
one more family, four more genera, and five more species occurred in
October (8/13/20/21) than in July. Thus, the spring and summer faunas
were less diverse than the October fauna. These fluctuations reflect varia-tions
in times of maturation and breeding of the different species.
The overall O/F/G/S ratio for the three months combined
(9/16/23/24) showed one more order, three more families, three more
genera, and three more species than occurred in any single month. This
reflects the appearance and disappearance of species during the year,
which is also indicated by the following seasonal trends. Five species -
P. ectenes, C. annulata, P. branneri, B. stricta and D. lea - were more
common in April than in July and again increased in abundance during
October. A different five species - P. strictum. P. ectenes, C. catawba,
P.c. incursus and S. latior -were more common in July than in either of
the cooler months. Three species - P. szrictum. P. ectenes, and P. im-pressus
-were most abundant in October, and two - N. americanus and
Kings Mountain Milliped Fauna 35
A. erythropygos - were common in all three months as both juveniles
and adults. Seven diplopods were encountered rarely (less than ten
specimens) and were collected during only one month (A. corticarius, B.
lusitanus, Teniulus sp., C. medialis, and T. dux), or two months (N.
minutw, and Striaria sp.) These data indicate that milliped faunas should
be sampled on a seasonal basis, a practice not generally followed to date,
and that collections in spring and fall may produce species not available
in summer.
COMPARISONOF FAUNASAN D SIGNIF~CANTCO EN ORTHC AROLINA
Spencer Mountain is separated from the contiguous Kings-Crow-ders
ridge by approximately 15 km of urbanized Piedmont, and as shown
in Table 4 fewer milliped species occur at the inselberg. At both Spencer
Mountain (S/G = 1.00) and Kings-Crowder ridge (S/G = 1.05) every
genus is represented by one species with the sole exception of Ptyoiulus,
for which both species are present at Kings-Crowders ridge. At Spencer
Mountain, however, three less families, seven less genera, and eight less
species (8/11/14/14) were encountered than at Kings-Crowders ridge
(8/14/21/22). The two areas had 12 species in common - P. fasciculatus,
P. ectenes, N. americanus, A. magnum, D. georgianum. C. annulata, 0.
gracilis, P. branneri, S. granulatus, A. erythropygos, B. stricta, and S.
latior. Two species collected only at Spencer Mountain, A. corticarius
and T. dux, were found in such low numbers (Table 3) that their absence
from the Kings-Crowders ridge could well be a collecting artifact.
The same is true of the apparent absence of six species from Spencer
Mountain - P. stricrum, B. lusitanus, N. minutus, Teniulus sp., C.
medialis, and Striaria sp. Of the remaining five species absent from Spen-cer
Mountain, P. impressus, B. stricta, and P.c. incursus have western or
southern ranges that may well end at Kings-Crowders ridge. Two
xystodesmids, C. catawba and D. lea, could occur at Spencer Mountain,
since both were collected from Lincoln County in the 1950s. Their
presence seems doubtful, however, since the extensive searches for
diplopods at Spencer Mountain would surely have revealed these large,
brightly colored, and obvious millipeds. Thus, the absence of these five
species from Spencer Mountain may be real.
In addition to faunal distinctions between the two areas, color pat-tern
variation was noted in S. latior. As discussed in the species account,
specimens from Spencer Mountain displayed yellow paranota and stripes
along the caudal edges of the metaterga, whereas those from Kings-
Crowders ridge had yellow paranota but lacked the metatergal stripes.
No anatomical differences were detected, and both color patterns are
representative of intergrades. This is the first report of S. larior inter-grades
without stripes, a trait characteristic of the nominate subspecies.
The diplopod fauna of the Kings Mountain region is also compared
with the faunas of the eastern Piedmont and Appalachian Mountains
(numerical data for the Great Smoky Mountains) in Table 4, and is
Marianne E. Filka and Rowland M. Shelley
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Kings Mountain Milliped Fauna 37
shown to have a lower S/G ratio and fewer taxa below the level of order
than either. Comparison with the entire eastern Piedmont is somewhat
misleading, however, since the land area investigated by Shelley (1978)
was much larger and contained a greater variety of biotopes than the
Kings Mountain region. A more meaningful comparison is with the three
smaller areas that he sampled in detail - Medoc Mountain and William
B. Umstead state parks, and the hardwood locality near Ellerbe -each
more comparable in size to the Kings Mountain region. The ratios for
these three sites are as follows: Medoc Mountain State Park (5161717,
S/G = 1.00); William B. Umstead State Park (8/12/14/15, S/G = 1.07);
and Ellerbe (7/10/12/12, S/G = 1.00). The Kings Mountain fauna is
higher in each taxonomic category than any of these sites, but their S/G
ratios still reflect the occurrence of essentially one species per genus. Only
Plyoiulus, with P. ectenes and P. impressus in the Kings Mountain region,
and Narcew; with N. americanur and N. annularis in Umstead State Park,
are represented by more than one species at a site. The greater numbers
of taxa in the Kings Mountain region may reflect its mountainous
character, hut the region is still unable to support significantly more than
one species per genus. Compared to the Appalachian Mountains in
general and the Great Smoky Mountains in particular, the Kings Moun-tain
region has fewer taxa in every category (the number of orders for the
Great Smoky Mountains was not reported by Hoffman 1969) and a
much lower S/G ratio. Many Appalachian genera are represented by
more than one species, a reflection of the greater variety of niches af-forded
by the rugged, heterogeneous terrain.
Despite the numerical differences, however, there are similarities
between the Kings Mountain region and the other areas. Eight species of
widespread distribution are common to all three: P. strictum, A. cor-ticariur,
N. americanus, A. magnum, C. annulata, 0. gracilis, P. branneri,
S. granulatus, and one or possibly two species of Sfriaria (taxonomic
problems exist within this genus). Some of the 24 species found in the
Kings Mountain region also occur in one of the others hut not both.
Seven typically Piedmont inhabitants currently unknown from the moun-tains
are shared with the eastern Piedmont - P, fascinclatus, N. minutus,
B. lusitanus, P. ectenes. T. dm. A. erythropygos, and S. latior. Most were
expected in the Kings Mountain region at the outset of the study. Five
species are likewise shared with the Appalachians - P. impressus.
Teniulus sp., C. medialis, D. georgianwn, and B. stricta. Their discovery in
the Kings Mountain region was a complete surprise and a significant
range extension for each. Fifteen species reported from the eastern Pied-mont
by Shelley (1978) were not found in the Kings Mountain region,
although three, Cylindroiulus truncorum (Silvestri), Ophyiulus pilosur
(Newport), and Apheloria tigana Chamberlin, are considered potential
inhabitants. The first two are synanthropic millipeds that could have
been overlooked in our study since we did not sample urban environ-ments.
Apkeloria tigana is so common in the eastern Piedmont and in the
38 Marianne E. Filka and Rowland M. Shelley
more proximal Uwharrie Mountains that it must he considered a
possibility for the Kings Mountain region. Five millipeds known from
both the Appalachian Mountains and eastern Piedmont must also be
considered potential occupants of the Kings Mountain region due to its
location between these two areas. These five species are Polyzonium
rosalbum (Cope), known from Madison and Moore counties (Shelley
1976a, 1978); Cleidogona caesioannulata (Wood), reported from Macon,
Jackson, Transylvania, and Swain counties (Shear 1972) and Granville,
Orange, Durham, and Johnston counties (Shelley 1978); Branneria
carinata (Bollman) cited from Transylvania and Macon counties (Shear
1972), and Wake County (Shelley 1978); Pseudopolydesmus serrafur.
collected in 14 eastern piedmont counties (Shelley 1978), and reported
generally from the mountains (Chamberlin and Hoffman 1958); and
Pleuroloma flavipes Rafinesque, recorded as Zinaria brunnea from
Watauga and Moore counties (Wray 1967) and as Pleuroloma sp. from
Orange and Wake counties (Shelley 1978).
In addition to species shared with the eastern Piedmont and/or Ap-palachian
Mountains, a fourth group of three xystodesmids is unique to
the Kings Mountain region: C. catawba, D. lea, and P.c. incursus. The
last is known in North Carolina only from the contiguous Kings-
Crowders ridge, but the others also have been recorded from Lincoln
County (Shelley 1978; Hoffman 1961), in the area that is the north-eastward
extension of the range to Anderson Mountain, Catawha
County. Croatania catawba and P.c. incursus are more common in South
Carolina and are basically southern forms which extend into North
Carolina along the Kings Mountain range. Together these two millipeds
lend a southern aspect to the Kings Mountain fauna, which is not found
in any other part of North Carolina. Deltotaria lea appears to be endemic
to a narrow section of the Carolinas, ranging from Lincoln County,
North Carolina, to Chester County, South Carolina.
Thus, the Kings Mountain milliped fauna is characterized by its own
species and the transitional ones it shares with the eastern Piedmont
Plateau and Appalachian Mountains, together and separately. Only five
of these species, however, are shared with the Appalachians alone. This,
plus the low diversity and thelowland nature of the fauna militate against
a prior direct topographic connection between the Blue Ridge Front and
the Kings Mountain region. Aside from a general Cretaceous peneplain
there is no geological evidence for such a connection, just as there is no
faunal evidence from the diplopods. Unlike the Appalachians, the Kings
Mountain region does not seem to have ever been a center of milliped
evolution and dispersal. The five Appalachian species in the area may be
relicts of a continuous Pleistocene or pre-Pleistocene distribution, as
suggested by Shelley (1979a) for D. georgianwn. The most significant as-pect
of the Kings Mountain region is its position at the known range
per~pheryo f several diplopods. It is the northern distribution limit of P.c.
incursus and the northeastern of the genus Pachydesmus (Shelley and
Kings Mountain Milliped Fauna
Filka 1979), and two montane millipeds, D. georgianum and B. stricta,
reach their eastern terminus in the area. It is also the easternmost limit
for P. impressus and the genus Teniulw, the southeasternmost known site
for C. medialis, and the western limit for P. ecfenes. The Kings Mountain
region is therefore a unique area in North Carolina, in the southern ele-ments
of its milliped fauna, in being a transitional area between
predominantly eastern and western faunas, and in forming a part of the
range periphery for four genera.
Teulings and Cooper (1977) used the term "cluster areas" to denote
places in North Carolina where species of concern are grouped. Four
rivers systems and four land areas in the Piedmont Plateau Province were
so identified. In a preliminary report, Filka and Shelley (1977) indicated
that, on the basis of its diplopod fauna alone, the Kings Mountain region
also would qualify as a cluster area. Three species considered of concern
in North Carolina occur in the region - P.c. incursus (endangered), and
C. catawba and D. lea (special concern). Moreover, the range peripheries
of P. ectenes, P. impressus, Teniulw sp., C. medialis, D. georgianum, B.
stricta and P.c. incursus lie there. The area also contains a unique
gonopod variant of B. stricta, and is distinguished by southern elements
of its fauna (C. catawba and P.c. incursus). As far as millipeds are con-cerned
the Kings Mountain region is of singular importance to North
Carolina, and investigations of other animal groups may provide further
evidence of its uniqueness. One state park, Crowders Mountain, exists in
the area, and every effort should be made to expand it to include the
deciduous bottomlands where most milliped species occur, including
those now considered of concern in the state. No millipeds were found
during this study in the dry, predominantly pine habitats of the existing
park.
One objective of this study, that of gaining insight into evolutionary
processes affecting millipeds in the southern Appalachians, went unmet.
With only five species in common and a lowland-type faunal diversity,
the Kings Mountain region adds little to current knowledge of milliped
biogeography that might be applied to such an objective. Moreover, none
of the five shared species belong to the xystodesmid tribe Aphelorini,
which is the single most diverse and abundant element of the Appala-chian
fauna. Aside from the ubiquitous Sigmoria latior, which ranges
from the mountains of West Virginia to the Coastal Plain of southern
South Carolina (Shelley 1976c), the great southeastern aphelorine fauna
is absent from the Kings Mountain region.
The study was, however, the first attempt to document seasonal
occurrence of milliped species in a discrete part of the southeast, an en-deavor
that should receive more attention. Seasonal sampling of juveniles
and adults can vield \,ahable information on life histories. for examole. ~
~ ~, . .
and basic biological knowledge of this type has never been determined
for most North American diplopods. Although direct rearing of larvae
and adults, and breeding experiments, would provide the best such infor-mation,
inferences can nevertheless be gained from seasonal collections.
40 Marianne E. Filka and Rowland M. Shelley
ACKNOWLEDGMENTS.-We take pleasure in thanking the per-sons
who helped collect specimens from the area of study, particularly
John C. Clamp, William W. Thomson, and Gerri W. Wicker. The as-sistance
of C.F. Lytle in providing equipment and laboratory space for
Filka is also gratefully appreciated. The specimens of C. medialis from
Deep Gap, Watauga County, North Carolina, were kindly loaned by
Norman I. Platnick, American Museum of Natural History; Shelley's
collecting in Boone's Cave and Crowders Mountain state parks was
done with permission of the North Carolina Department of Natural
Resources and Community Development, Division of State Parks.
Specimens of B. stricta and D. lea collected by Shelley in Kings Mountain
and Croft state parks, South Carolina, were secured through courtesy of
the South Carolina Department of Parks, Recreation, and Tourism. This
research was partly funded by the North Carolina State Museum of
Natural History and by the Department of Zoology, North Carolina
State University, and constituted part of a Master of Science thesis sub-mitted
to the latter institution by Filka.
LITERATURE CITED
Bollman, Charles H. 1887. Descriptions of fourteen new species of North Ameri-can
myriapods. Proc. U.S. Natl. Mus. 10:617-627.
Brimley, C.S. 1938. Insects of North Carolina. N.C. Dep. Agric. Div. Entomol.,
Raleigh. 560 .D D..
~r6lemann,H enri W. 1895. Liste de myriapodes des Etats-Unis, et principale-ment
de la Caroline du Nord, faisant partie des collections de M. Eugene
Simon. Ann. Soc. Entomol. France 65:43-70.
Burney, D.A. 1974. A preliminary interpretive prospectus of the Crowder's
Mountain-King's Pinnacle area of Gaston County, North Carolina. Unpubl.
rept. Div. Parks Rec., State Parks Sec. Raleigh. 14 pp.
Causey, Nell B. 1940. Ecological and systematic studies on North Carolina
myriapods. Ph.D. dissert., Duke Univ., Durham. 181 pp.
. 1943. Studies on the life history and ecology of the hothouse milli-pede,
Orthomorpha gracilis (C.L. Koch, 1847). Am. Midl. Nat. 3:670-682.
Chamherlin, Ralph V. 1940a. On some chilopods and diplopods from North
Carolina. Can. Entomol. 7256-59.
. 1940b. Four new polydesmoid millipeds from North Carolina
(Myriapoda). Entomol. News 51:282-284.
. 1951. On eight new southern milli~eds.G reat Basin Nat. 11:19-26.
, and R.L. Hoffman. 1958. ~hec'klist of the millipeds of North
America. U.S. Natl. Mus. Bull. 212. 236 pp.
C-n-nn-err. ~Joh~n .E .. S.S. Robinson and J.B. Funderbure reds.). 1977. Endaneered ~ and ~ hreatened Plants and Animals of North Faiolina. N.C. State %us.
Nat. Hist., Raleigh. xvi + 444 pp.
Enghoff, Henrik, and R.M. Shelley. 1979. A revision of the millipede genus
Nopoiulw (Diplopoda:Julida:BIaniulidae). Entomol. Scand. 10:65-72.
Filka, M., and R.M. Shelley. 1977. The Kings-Crowders Mountain Region: A
milliped "cluster" area in North Carolina (Diplopoda). ASB Bull. 24(2):29.
Abstract.
Gardner, Michael R. 1975. Revision of the milliped family Andrognathidae in
the Nearctic Region. Mem. Pac. Coast Entomol. Soc. 5:l-61.
Kings Mountain Milliped Fauna 4 1
Hardin, James W., and A.W. Cooper. 1967. Mountain disjuncts in the eastern
Piedmont of North Carolina. J. Elisha Mitchell Sci. Soc. 83:139-150.
Hoffman, Richard L. 1950. Records and descriptions of diplopods from the
southern Appalachians. J. Elisha Mitchell Sci. Soc. 6611-33.
. 1958. Appalachian Cambalidae: Taxonomy and distribution (Dip-lopoda:
Spirostreptida). J. Wash. Acad. Sci. 48:90-94.
, 1961. Revision of the milliped genus Dehotaria (Polydesmida:
Xystodesmidae). Proc. U.S. Natl. Mus. 113:15-35.
, 1962. The milliped genus Scytonotus in eastern North America,
with the description of two new species. Am. Midl. Nat. 67241-249.
, 1965. Revision of the milliped genera Boraria and Gyalostethus
(Polydesmida:Xystodesmidae). Proc. U.S. Natl. Mus. 117305-347.
, 1969. The origin and affinities of the southern Appalachian diplo-pod
fauna. pp. 221-246 in P.C. Holt (ed). The distributional history of the
biota of the southern Appalachians, part I: Invertebrates. Res. Div.
Monogr. 1, Va. Polytech. Inst. Blacksburg. 295 pp. . 1974. A new polydesmid milliped from the southern Appalachians
with remarks on the status of Dixidesmus and a proposed terminology for
polydesmid gonopods. Proc. Biol. Soc. Wash. 87:345-350.
Hunt, Charles B. 1967. Physiography ofthe United States. W.H. Freeman & Co.,
San Francisco. 480 pp.
Keeton, William T. 1960. A taxonomic study of the milliped family Spiro-bolidae
(Diplopoda:Spiroholida). Mem. Am. Entomol. Soc. 11. 146 pp.
Keith, A. 1931. Geologic Atlas of the United States, Gaffney-Kings Mountain
Folio, South Carolina-North Carolina. U.S. Geol. Survey #222. 13 pp. + 4
maps.
Kesel, Richard H. 1974. Inselbergs on the Piedmont of Virginia, North Carolina,
and South Carolina: Tyues and characteristics. Southeast. Geol. 16:l-30.
Shear. Willi;im A. 1972. ~ t u d / r irn the millipcd order Chordcumida (Diplopoda):
A revifion of rhc fam~lv Cleidoconidae and a reclassification of the order
Chordeumida in the new wor ldr~ul l .M us. Comp. Zool. 114:151-352.
Shelley, Rowland M. 1976a. Two new diplopods of the genus Polyzonium from
North Carolina, with records of established species (Polyzoniida:Poly-zoniidae).
Proc. Biol. Soc. Wash. 88:373-382.
. 1976h. A new diplopod of the genus Cleidogona from North
Carolina (Chordeumida:Cleidogonidae). Fla. Entomol. 59:325-327.
. 1976~. Millipeds of the Sigmoria larior complex (Polydesmida:
Xystodesmidae). PFOC. Biol. SOC. Wash. 89:17-38.
. 1977. The milliped genus Croatania (Polydesmida:Xystodesmi-dae).
Proc. Biol. Soc. Wash. 90:302-325. . 1978. Millipeds of the eastern Piedmont region of North Carolina,
U.S.A. (Diplopoda). J. Nat. Hist. 12:37-79. . 1979a. A revision of the milliped genus Delophon. with the proposal
of two new tribes in the subfamily Abacioninae (Ca1lipodida:Caspio-petalidae).
Proc. Biol. Soc. Wash. 92533-550,
, 1979b. A synopsis of the milliped genus Cambala, with a description
of C. minor Bollman (Spirostreptida:Camhalidae). Proc. Biol. Soc. Wash.
92:551-571.
, and M. Filka. 1979. Occurrence of the milliped Pachydesmus crassi-cutis
incursus Chamberlin in the Kings Mountain region of North Carolina
Marianne E. Filka and Rowland M. Shelley
and the Coastal Plain of South Carolina (Polydesmida:Xystodesmidae).
Brimleyana 1:147-153.
Stuckey, Jasper L. 1965. North Carolina: Its Geology and Mineral Resources.
N.C. Dep. Cons. Devel.. Raleigh. 550 pp.
Teulings, Robert P., and J.E. Cooper. 1977. Cluster Areas. pp. 409-431 in
J.E. Cooper, S.S. Robinson, and J.B. Funderberg (eds.). Endangered and
Threatened Plants and Animals of North Carolina. N.C. State Mus. Nat.
Hist., Raleigh. xvi + 444 pp.
Wray, David L. 1967. Insects of North Carolina, Third Supplement. N.C. Dep.
Agric. Div. Entomol., Raleigh. 181 pp.
Accepted I5 September 1980
Electrophoretic Analysis of Three Species of Necturus
(Amphibia: Proteidae), and the Taxonomic Status
of Necturur lewisi (Brimley)
RAYE . ASHTONJ, R.A ND ALVINL . BRASWELL
North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh. North Carolina 27611
AND
SHELDONI. GUTTMAN
Department of Zoology,
Miami University, Oxford, Ohio 45056
ABSTRACT. Electrophoretic :~nalyseso f 10 Necnrru~m aruloruz from
Minnesota. 10 from biassachusetts. and I from the Mills River. Hen-derson
county, North Carolina, here compared with those of 20
Necrums lewisi and 8 unspotted Nectum punctatus from the Neuse
River drainage and 8 spotted N. puncratus from Naked Creek, Robeson
County, North Carolina. Evaluation of 17 loci showed that the three
samples of N. maculosus were indistinguishable (Nei's D = 0.000) while
N. Iewisi were unequivocally different from N. punctatus at four loci and
from N. mnculosus at six loci. The two N. punctalus populations were in-distinguishable
from each other but were distinguishable from N.
manrlosur at 6 loci. These data indicate that N. maculosus. N. lewisi and
N. punctatus are distinct, long isolated species.
INTRODUCTION
Necturus lewisi is one of several endemic species of vertebrates and
invertebrates found in the Tar and Neuse River drainages of North
Carolina. This waterdog was originally described by Brimley (1924) as a
subspecies of Necturus maculosus because of the "spotted larvae". Viosca
(1937) briefly described the previously unknown striped larvae of N.
lewisi and used its medium size and overall spotting as the apparent
criteria for elevating it to full species status. Ashton and Braswe11 (1979)
compared N. lewisi hatchlings and striped larvae with larvae of N.
maculosus and N. punctatus, and found that the striped larvae of N. lewisi
were quite distinctive. No electrophoretic studies in the genus have been
reported. Our study compared electrophoretic data for all three of these
Necturus species, in an attempt to evaluate the taxonomic status of N.
lewisi.
METHODS AND MATERIALS
Ten N. maculosus from Minnesota were obtained from Nasco, Fort
Atkinson, Wisconsin and ten from Massachusetts were purchased from
Connecticut Valley Biological Supply Company, Southampton,
Massachusetts. One additional N. maculosus was collected in the Mills
River, Henderson County, North Carolina. Twenty N. lewisi and eight
unspotted N. punctatus were captured in the Neuse River drainage. Eight
Brimlnjana No. 1: 4146. December 1980. 43
44 Ray E. Ashton, Jr., Alvin L. Braswell, Sheldon I. Guttman
spotted N. punctarus were collected from Naked Creek, PeeDee River
drainage, in the Sandhills region of Robeson County, North Carolina.
Animals were killed in the laboratory and an organ homogenate pre-pared
from the heart, liver, rinsed stomach and upper part of intestine,
and kidney and tongue of each. The specimen remains are housed in the
North Carolina State Museum collection. The tissues of individual
animals were then homogenized in an equal volume of 2?0 2-
phenoxyethanol and centrifuged at 25,000 g at 4' C for 45 minutes. The
supernatant of soluble proteins was then decanted and stored at -70" C
until used a maximum of 48-hours following preparation.
The 17 loci coding for proteins consistently resolved are as follows:
malate dehydrogenase (NAD-dependent) (Mdh-I); indophenol oxidase
(Ipo-I); or-glycerophosphate dehydrogenase (a-Gpdh-I); isocitrate
dehydrogenase (NADP-dependent) (Idh-I); phosphoglucomutases, three
loci (Pgm-I, Pgm-2, Pgm-3); glutamate oxalate transaminases, two loci
(Got-I, Got-2); glutamate dehydrogenase (Gdh-I); phosphoglu-coisomerase
( P g i malic enzyme, two loci (Me-I, Me-2); 6
phosphogluconate dehydrogenase (6-Pgdh-I); sorbitol dehydrogenase
(Sdh-I), glyceraldehyde-3-phosphate dehydrogenase (G-3-pdh-I); and
lactate dehydrogenase (Ldh-2).
Techniques of horizontal starch gel electrophoresis and protein
staining were similar to those described by Selander et al. (1971). with the
following modifications: Idh, Pgm, Mdh, Gdh, and Me were examined
with their continuous tris-citrate buffer (pH 8.00); 6-Pgdh, Got, Sdh and
G-3-pdh were demonstrated with the tris-borate-EDTA buffer of Ayala
et al. (1973). Staining methods for Gdh, G-3-pdh and Sdh were as
described by Brewer (1970). All gels were 12.55 starch (Electrostarch Lot
#307).
~ e n e t i cin ferences from electrophoretic results are based on the pat-terns
being consistent with known molecular configurations for the pro-teins
analysed, i.e. two-banded patterns are observed for the
heterozygotes for a protein that is a monomer and three-banded patterns
are observed for a dimeric heterozygote. The genes coding for each en-zyme
are represented by italicized abbreviations.
If several forms of the same enzyme are present and each is con-trolled
by a separate gene locus, the hyphenated numeral serves to dif-ferentiate
the loci. The enzyme with the greatest anodal migration is
designated one, the next two, and so on. When allelic variation occurs,
the allele with the greatest anodal migration is called a, the next b, and so
on.
RESULTS
The two N. maculosus samples were essentially identical genetically
(genetic distance, D = 0.000; Nei 1972). One heterozygote was found at
each of the two loci (Pgm-2, Idh-I) in the Massachusetts sample, the only
heterozygotes found.
Nectum Electrophoresis 45
The only variants found in the N. lewisi sample were at the Got-1
locus. Six individuals were heterozygous for the a and b alleles, one was
homozygous for b.
Two unspotted N. punctotus were each heterozygous at single loci
(Pgi-1, Pgm-2). The only difference between the spotted and unspotted
samples was in the frequency of the Pgi-1 alleles (Table 1).
Table I. Fixed genetic differences in three Nectum species
Locus Allele
N. [ewisi N. moculosus N. punctatus
spotted unspotted
Mdh-l b a b b
The three species were unequivocally different at four (N. lewisi vs.
N. punctotur) or six (N. maculosus vs. N. lewisi or N. punclatus) of the
seventeen loci investigated; alleles were not shared at these loci (Table 1).
Nei's standard genetic distance estimates between each pair of species
(Table 2) are indicative of a long history of isolation of the gene pools.
Electrophoretic examination of one N. moculosus from the Mills River,
Henderson County, North Carolina confirms the genetic distinction
found between the larger samples of allopatric N. maculosus and N.
lewisi.
Table 2. Standard genetic distance (D) between species of Nectunrs examined.
I. N. lewisi 0.0
2. N. maculosus 0.435 0.0
3. N.puncmnrs(unspotted) 0.348 0.435 0.0
4. N. punctotus (spotted) 0.339 0.426 0.040 0.0
46 Ray E. Ashton, Jr., Alvin L. Braswell, Sheldon I. Guttman
DISCUSSION
Electrophoretic analysis of ten individuals from each of two popula-tions
of N. maculosus and one individual from a third population showed
that they were indistinguishable using this technique. Necturus lewisi and
N. punctatus, however, were highly distinguishable from each other and
from i. maculosus, indicating that each species has been genetically
isolated for some time. Two populations of N. puncfatus, the spotted
form inhabiting the Sandhills region of North Carolina and the uni-formly
gray-black form inhabiting the Neuse River, were in-distinguishable
from each other.
In conclusion, the specific status of N. lewisi is confirmed by elec-trophoretic
data as well as by the distinct larvae described by Ashton and
Braswell (1979). Further, N. punctafm appears to have been reproduc-tively
isolated from sympatric N. lewisi and from allopatric N. maculosus
for a considerable period of time, and spotted N. punctatus from the
PeeDee River drainage (North and South Carolina) appear on the basis
of electrophoresis to be genetically similar to the unspotted populations
of the Neuse River system.
ACKNOWLEDGMENTS.-The authors wish to express their ap-preciation
to field technicians Angelo Capparella, Paul Freed, and Jerry
Reynolds, and laboratory assistants Gary Trakshel, Kim Haikyong, and
Ernie Flowers. We also thank John E. Cooper for his critical review of
the manuscript. This project was in part supported by funds from a U.S.
Fish and Wildlife Service (Office of Endangered Species) cooperative
agreement with the North Carolina Wildlife Resources Commission.
LITERATURE CITED
Ashton, Ray E., Jr. and A.L. Braswell. 1979. Nest and larvae oftheNeuse River
Waterdog, Necturur lewisi (Brimley) (Amphibia: Proteidae). Brimleyana
.1. .1. 5. -7 2
Ayala, Francisco J., D. Hedgecock, G. Zumwalt and J. Valentine. 1973. Genetic
variation in Tridacna maxima, an ecological analog of some unsuccessful
evolutionary lineaaes. Evolution 27:177-191.
Brewer, George. 19701 Introduction to isozyme techniques. Academic Press,
New York. 186 pp.
Brimley, Clement S. 1924. The Water Dogs (Necturw) of North Carolina.
J. Elisha Mitchell Sci. Soc. 40(3-4):166-168.
Nei, Masatoshi. 1972. Genetic distance between .o oo, ulations. Am. Nat.
105283-292.
Selander, Robert K., M.H. Smith, S.Y. Yang, W.E. Johnson and J.B. Gentry.
1971. Biochemical polymorphism and systematics in the genus Perornyscur.
I. Variation in the old-field mouse (Peron~yscuspolionorus)S. tud. Genet. VI.
Univ. Texas Publ. 7103:49-90.
Viosca. Percy, Jr. 1937. A tentative revision of the genus Necrum, with descrip-tions
of three new species from the southern Gulf drainage area. Copela
1937(2):120-138.
Accepted 14 Ocfober 1980
Vertebrates of the Okefenokee Swamp
JOSHUAL AERMB, .J. FREEMANL,A URIEJ. VITT
Museum of Natural History and
Department of Zoology
JOSEPH M. MEYERS
Institute of Ecology
AND
LLOYD LOGAN
Museum of Natural History and
Department of Zoology
University of Georgia, Athens, Georgia 30602
ABSTRACT.-Four hundred nineteen vertebrate species and sub-species
are known from the Okefenokee Swamp region of Georgia and
adjacent Florida. These include 36 fishes, 37 amphibians, 66 reptiles,
232 birds, and 48 mammals. The vertebrates occurring in the
Okefenokee represent a typical southeastern Atlantic Coastal Plain
fauna. There are no endemic species. Eleven species, recognized as
threatened or endangered under state and/or federal guidelines, occur
in the swamp.
INTRODUCTION
The Okefenokee Swamp region of southeastern Georgia and adja-cent
Florida contains an extremely diverse vertebrate fauna. However,
with the exception of biological surveys conducted by Cornell University
in the early decades of this century, that fauna has received little atten-tion.
At present there exists no comprehensive information on the ver-tebrates
of the swamp. Most of the available literature is semipopular,
anecdotal, or, at best, outdated.
Accurate faunal information is essential to understanding the Oke-fenokee
Swamp ecosystem. The long term value and credibility of the
systems ecology studies presently being undertaken in the swamp will, in
large part, be determined by the extent to which base level natural history
information can be incorporated into definitive analyses and models.
Base level faunal surveys provide information on species diversity and
patterns of habitat use that are crucial for biogeographic and systematic
research. Furthermore, comprehensive faunal studies serve also as dated
testaments to species composition and distribution within specific
habitats, which are crucial for enviromental impact assessments
associated with management practices.
For these reasons, we have undertaken vertebrate faunal surveys
within the Okefenokee Swamp and surrounding uplands. We report here
the results of these surveys. We present, too, a review of pertinent
historical foundations of our present knowledge of the swamp's ver-tebrate
fauna, a comparison of the fauna with that of adjacent south-eastern
regions, and a preliminary analysis of habitat distributions of ver-tebrates
known to occur within the swamp.
Brimlcyana No. 4: 47-73. December 1980. 47
48 Joshua Laerm, et al.
GENERAL HABITAT CHARACTERISTICS
The Okefenokee Swamp is one of the largest freshwater wetlands in
the United States. Situated in Charleton, Clinch, Echols and Ware coun-ties,
Georgia, and Baker and Columbia counties, Florida, the Oke
fenokee watershed includes both swamp (189,000 ha) and surrounding
uplands (181,000 ha). It lies within the humid subtropical climatic zone
(Trewartha 1968) and is characterized by warm moist springs, hot wet
summers, warm dry falls, and cool moist winters. Weather is
predominantly influenced by tropical maritime air masses from the Gulf
of Mexico and the tropical Atlantic Ocean in spring, summer and fall,
but by continental air masses in winter. Annual precipitation averages
100-150 cm (Hunt 1972).
The Okefenokee consists of a variety of vegetational habitat types.
Plant specimens are on file at the University of Georgia Herbarium.
a. Two prairie habitat types are identified, comprising approx-imately
21% of the swamp. (1) Aquatic macrophyte prairies are
dominated by emergent, floating-leaved, and submerged hydrophytes
such as white water lily, Nymphaea odorata; yellow water lily, Nuphar
luteum; neverwet, Orontium aquaticum; floating heart, Nymphoides
aquaticum; yellow eyed grass, Xyris smalliana; pickerel weed, Pontederia
cordata; redroot, Lachnanthes caroliniana; and bladderwort, Utricularia
spp. (2) Grass-sedge prairies are characterized by various species of
sedges, Carex; panic grasses, Panicum; and beak rush, Rhynchospora, as
well as broomsedge, Andropogon virginicus; giant chain fern, Woodwardia
virginica; and Sphagnum moss.
b. Shrub swamps cover approximately 34% of the swamp and are
predominated by hurrah bush, Lyonia lucida; fetter hush, Leucothoe
racemosa; titi, CyriNa racemiflora; sweet spire, ltea virginica; pepper
bush, Clerhra alnifolia; and dahoon, Ilex cassine.
c. Blackgum forests cover less than 6% of the swamp. Blackgum,
Nyssa sylvatica var. biflora, with a small amount of dahoon and pond
cypress, Taxodium ascendens, dominates the canopy, with red maple,
Acer rubrum, and dahoon the predominant understory plants.
d. Bay forests also cover less than 6% of the swamp. Loblolly hay,
Gordonia Iasianthus; red bay, Persea borbonia; and sweet bay, Magnolia
virginiana, are the predominant canopy species although occasional pond
cypress, hlackgum, and slash pine, Pinw elliottii, are seen.
e. Mixed cypress forests are characterized by pond cypress domi-nated
canopy and subcanopy, but loblolly bay, dahoon, and blackgum
are frequently scattered in the subcanopy. This and the following habitat
make up approximately 23% of the swamp.
f. Pure cypress forests are limited in extent but consist almost en-tirely
of a cypress canopy with a sparse subcanopy or understory.
g. There are approximately 70 islands in the swamp and they ac-count
for roughly 12% of the area. Vegetation is dominated by loblolly
pine, Pinus taeda: slash pine; longleaf pine, Pinus palustris; water oak,
Okefenokee Swamp Vertebrates 49
Quercus niger: and live oak, Quercus virginiana.
The uplands surrounding the swamp are intensively managed pine
forests. Historically, the area was dominated by longleaf and slash pine
with an understory dominated by saw palmetto, Serenoa repens; small
gallberry, IIex glabra; and various forbs and grasses. Fire was the major
factor maintaining successional stages (Monk 1968). Today the uplands
are dominated by slash pine plantations with a similiar understory
managed by prescribed periodic burns. Remnants of hardwood and
mixed hardwood-pine forests are very limited but occur in scattered loca-tions
on some islands and at the periphery of the swamp. Management
for pine, including prescribed burns, is responsible for the virtual absence
of hardwoods in the uplands.
FISHES
HISTORICAFLO UNDATIONS
Scientific collections of fishes in the Okefenokee Swamp span 68
years. The earliest significant collections were undertaken in 1912 by
A.H. Wright and Francis Harper, both from Cornell University. The ac-count
of Palmer and Wright (l920), based primarily on these collections,
represents the only published information on fishes of the swamp. Subse-quent
collections, resulting from various museum expeditions and the ac-tivities
of Okefenokee National Wildlife Refuge (ONWR) personnel,
were made by R.A. Chesser in 1922, R.T. Berryhill in 1924, T.
Reichelderfer in 1935, M.S. Verner, Jr. in 1936, B. Cadbury in 1937, C.B.
Obrecht and M. Godfrey in 1941, H.A. Carter in 1941-1942, Southern
Piedmont and Coastal Plain Survey in 1941, T. Rodenberry in 1941, and
R.J. Fleetwood in 1947. Collecting activities ceased in the 1950s and
began again in the 1960s (E. Cypert in 1960, 1963; T. Cavender in 1965;
and M.W. Bohlke in 1966), and have continued to the present (B.J.
Freeman, 1978-1980). Additional studies in the southeastern lower
Coastal Plain (Gassaway 1976; Holder and German 1977) contributed
much to existing knowledge of swamp ichthyofauna. Voucher specimens
of significant collections are deposited in the National Museum of
Natural History, Philadelphia Academy of Natural Sciences, Cornell
University, University of Georgia Museum of Natural History, and Uni-versity
of Michigan Museum of Zoology.
COMPARISOWNI TH REGIONAFLA UNA
The ichthyofauna of Okefenokee Swamp consists of 36 species
representing 13 families (Table I). The most remarkable character of the
fauna is the absence of minnows (Cyprinidae). The remaining fish fauna
is not substantially different from adjacent southeastern drainages.
Average faunal resemblance values (Ramsey 1965) were computed for
Okefenokee Swamp and major drainages in the area. Values can range
50 Joshua Laerm, et al,
from 0 to I, with 0 indicating no species in common and 1 indicating all
species in common. The river systems compared were the Suwannee
River (from Fargo, Georgia to its junction with the Alapaha River); the
Alapaha River (a Suwannee River tributary); the Withlacoochee River (a
Suwannee River tributary); the St. Mary's River, and the Satilla River.
The values ranged from a minimum of .84 for the Withlacoochee to a
maximum of .90 for the St. Mary's. The Alapaha, Suwannee, and Satilla
were intermediate, with resemblance values of .86, 26, and .88, respec-tively.
These differences are due entirely to absence from the swamp of
minnows, which otherwise are widely distributed in adjacent drainages.
Their absence appears to be due to substantially lower pH values in the
swamp.
Swamp pH ranges from 3.1 to 4.2; pH values for surrounding
streams (where minnows occur) range from 4.8 to 6.9. In a study of
Carolina bay lakes in North Carolina, Frey (1951) noted that in two lakes
with a pH of 4.3 there were no minnows, while lakes with higher pH
values (up to 5.9) had some minnows present. These were Notropis
chrysoleucas, N. chalybaew, and N. petersoni - three of the species that
occur near the Okefenokee Swamp. Comparing minnow distributions
with pH shows that appearance of minnows in the St. Mary's River coin-cides
with a pH of 4.8 or higher. The pH values for surrounding streams
are even higher. Although detailed pH data for these streams are not
available (especially for the Suwannee River section) the general pattern
suggests that increasing acidity might limit, or at least influence, minnow
distributions. This possibility deserves more critical attention.
HABITATD ISTRIBUTIOONF THE FISHES
The 36 species of fish occurring in the swamp are distributed in a
heterogeneous series of aquatic habitats that can be broadly classified as
lake, aquatic prairie, and stream.
Lakes are open bodies of water of .25 ha or larger with depths of .5
m or more. The bottom is generally unconsolidated peat, which may have
a depth of .3 m to greater than 1 m; some lakes, however, have hard sand
bottoms. The margins are heavily vegetated with rooted and floating
aquatic plants as well as submergent vegetation. The topography around
the lakes grades into aquatic prairie (when the water levels are not low)
composed of a variety of rooted aquatic macrophytes, floating vegeta-tion,
sedges, and small shrubs. Water depth may range from several cm
to over I m. Current in these two areas varies from none in the lakes to
noticeable in the prairies. Streams generally have noticeable to moderate
current, consolidated banks, and sandy bottoms. Some aquatic vegeta-tion
and backwater areas are at the water margins.
The streams are located primarily in the northwest part of the
swamp and on some of the islands. The prongs of the Suwannee River
and the Suwannee Canal also provide stream habitat. Elements of the
prairie habitat, i.e. heavily vegetated areas, can be found bordering lakes
Okefenokee Swamp Vertebrates 5 1
and streams as well as in the large. open expanses of the swamp.
Aquatic habitats are not discrete units in the swamp but are graded
and sometimes mixed. The distribution of fishes reflects this. The habitat
associations of the swamp ichthyofauna indicates the fishes are rather
uniformly distributed (Table 1). Comparison of water current preferences
among the species does, however, indicate some degree of habitat
segregation. Noturus leptacanthus and Percina nigrofasciata will
generally be found in water with noticeable to moderate current. Umbra
pygmaea. Fundulus chrysotus, Fundulus cingulatus, Fundulus lineolatus,
Leptolucania ommata, Heterandria formosa, Elassoma evergladei and
Elassoma okefenokee generally are found in areas with no current but
with abundant aquatic vegetation. The remaining fishes occur in areas
with water currents ranging from none to noticeable. This wide range of
current tolerances helps explain the overlap observed in fish distributions
across obvious physically different habitats.
Table I. List of fishes of the Okefenokee Swamp. Based on museum records and
data from Dahlberg and Scott (1970), Gasaway (1976), Holder and
German (1977), and personal observations (B.J. Freeman). Scientific
and common names based on Bailey et al. (1970). L = lake, P= prairie,
S = stream.
SPECIES
ORDER SEMIONOTIFORMES
Family Lepisosteidae
Lepisos~arsplatyrhincur. Florida gar
ORDER AMllFORMES
Family Amiidae
Amia calva. Bowfin
ORDER ANGUILLIFORMES
Family Anguillidae
Anguilla roslrata. American eel
ORDER OSTEOGLOSSIFORMES
Family Esocidae
Esox americanus, Redfin pickerel
Esox niger, Chain pickerel
Family Umbridae
Umbrapygmaea, Mudminnow
ORDER CYPRINIFORMES
Family Catostomidae
Erimyzon sucella. Lake chubsucker
Minyrremamelanops, Spotted sucker
HABITAT
PREFERENCE
L P S
L P S
L P S
L P S
L P S
L P S
L P S
L S
52 Joshua Laerm, et al,
SPECIES
ORDER SILURIFORMES
Family Ictaluridae
Icralurusnatnlis. Yellow bullhead
Ictalurusnebulo~us. Brown bullhead
Icralurus ounctatus, Channel catfish
~orurus~yr inuTsa.d pole madtom
Notum lepracanrhus, Speckled madtom
ORDER PERCOPSIFORMES
Family Aphredoderidae
Aphredoderussayanus. Pirate perch
ORDER ATHERINIFORMES
Family Cyprinodontidae
Fundulus chrysotus. Golden topminnow
Fundulur cingulatus, Banded topminnow
Fundulur lineolatus, Lined topminnow
Leprolucania ommafa, Pygmy killifish
Family Poeciliidae
Gambusiaafflnis, Mosquitofish
Hererandria formosa. Least killifish
~ a m i~lt~h e r i i i d a e
Labidesthes sicculus, Brook silverside
HABITAT
PREFERENCE
L P S
L P S
L P S
L P S
L P S
L P S
L P S
L P S
L P S
ORDER PERCIFORMES
Family Elassomidae
EIassomaevergladei. Everglades pygmy sunfish L P S
Elassoma okefenokee, Okefenokee pygmy sunfish L P S
Family Centrarchidae
A cantharcuspomofis, Mud sunfish L P S
Cenrrarchusmacroprerus, Flier L P S
Enneacanthus chaetodon, Blackbanded sunfish L P
Enneacanthusgloriosus, Bluespotted sunfish L P S
Enneacanthus obesus. Banded sunfish L P S
Lepomis gulosus, Warmouth L P S
Lepomismacrochirus, Bluegill L P S
Lepomismarginarus, Dollar sunfish L P S
Lepomispuncrarus. Spotted sunfish L P S
Microprerussalmoides. Largemouth bass L P S
Pomoxis nigromacuiatus. Black crappie L S
Family Percidae
Etheosromo fusiforme, Swamp darter L P S
Percina nigrofmciara, Blackbanded darter S
AMPHIBIANS AND REPTILES
HISTORIFCOAULND ATIONS
Serious investigations of the herpetofauna of Okefenokee Swamp
began in 1912 with the first in a series of surveys conducted by Cornell
University. Prior to this only anecdotal accounts of the reptiles and
Okefenokee Swamp Vertebrates 53
amphibians are known (Fountain 1901 [cited in Wright and Funkhouser
19151; Reese 1907). At least three herpetologists participated in the Cor-nell
collections: A.H. Wright, W.D. Funkhouser, and S.C. Bishop. Dur-ing
the same period (and possibly with the same expedition) F. Harper
began recording observations on some of the reptiles and amphibians. In
two summary publications (Wright and Bishop 1915; Wright and
Funkhouser 1915), 9 chelonians, 6 saurians (actually 7, as Wright and
Funkhouser had 2 species of Ophisaurus), 21 serpents and 1 crocodilian
were recorded. This represents less than half the currently known fauna.
Harper (1934) discussed aspects of the ecology and behavior of several
Okefenokee reptiles and amphibians based on his visits, and numerous
short papers on aspects of the biology of Okefendkee species, mostly
authored by A.H. Wright, appeared in various scientific journals. Many
of the observations on anurans in Wright (1932) and Wright and Wright
(1949) were based on Okefenokee studies. Since these early visits to the
swamp (up to about 1946), there has only recently been a renewed in-ierest
in its herpetofauna. Several southeastern herpetologists made small
collections in the area, including W.T. Neill and F.L. Rose, but the
collections made by C.H. Wharton and his students at Georgia State
University are by far the most extensive. Additional surveys have been
conducted by L. Vitt and J. Laerm. Significant collections of Okefenokee
material can be found at Cornell University, Florida State Museum,
National Museum of Natural History, University of Georgia Museum of
Natural History, and University of Michigan Museum of Zoology.
COMPARISOWNI TH REGIONALFA UNA
The Okefenokee Swamp contains a diverse herpetofauna of 103
species and subspecies including 2 crocodilians, 15 chelonians, 38 ser-pents,
11 saurians, 16 urodeles, and 21 anurans (Table 2). The present
herpetofauna can be considered a typical southeastern Atlan.tic Coastal
Plain fauna (see Conant 1975). There are no species endemic to the
swamp. In general species diversity within the swamp and surrounding
uplands is greater than in similar sized areas in the adjacent southeastern
Atlantic Coastal Plain, primarily because of the high habitat diversity
associated with the swamp. However, the high species diversity can also
be attributed to the fact that at least 20 species of reptiles and amphibians
reach the limit of their natural range in the region of the swamp (see Co-nant
1975). Thus, the faunal diversity is somewhat greater in the
Okefenokee region in comparison to other Atlantic Coastal Plain
localities to the immediate north or south.
HABITATD ISTR~BUTIOOFN THE AMPHIBIANASN D REPTILES
Unlike the other vertebrates, most amphibians and reptiles in the
Okefenokee are not usually associated with a particular vegetational
habitat (blackgum swamps, for example) but rather seem to be associated
with structural habitats (water courses, sandy bottoms, etc.) Thus, it
54 Joshua Laerm, et al.
serves little purpose togroup species by vegetation habitats recognized by
biologists. Ecological distribution of the herpetofauna can, however, be
summarized in terms of general habits of the animals. Many species, for
example, are entirely aquatic and use most if not all aquatic habitats in
the swamp. Other categories also are useful in respect to ecological dis-tribution
of the reptiles and amphibians. For descriptive purposes, the
herpetofauna is partitioned into six "ecological" groups (Table 2): 1) En-tirely
aquatic species are those that spend nearly all of their lives in water;
2) Semi-aquatic species are those that spend a major part of their lives in
water, but may often be found on land (does not include species entering
water only for breeding); 3) Fossorial species are those that spend most of
their lives underground (they may become surface active for breeding or
limited foraging); 4) Terrestrial species are those most often encountered
on the surface and that spend most of their active time there; 5)
Terrestrial-arboreal species may spend nearly as much time in arboreal
habitats as on the surface; 6) Arboreal species are those that spend nearly
all of their lives in vegetation (some of these may enter water to breed, or
lay eggs on the ground).
Of the Okefenokee Swamp herpetofauna, 25 (24.3%) species are en-tirely
aquatic, 21 (20.4%) are semiaquatic, 10 (9.7'70) are fossorial, 29
(28.2%) are terrestrial, 9 (8.7%) are terrestrial-arboreal, and 9 (8.7%) are
arboreal. Most turtles are either aquatic or semiaquatic, most lizards
tend to be terrestrial, terrestrial-arboreal or arboreal, most snakes are
terrestrial (but there are large numbers of species in other groups), most
salamanders are aquatic, semiaquatic or fossorial, and frogs (including
toads) tend to be semiaquatic or arboreal (Table 2).
Table 2. List of amphibians and reptiles of the Okefenokee Swamp. Based on
museum records and data from Wright and Funkhouser (1915), Wright
and Bishop (1915), Wright (1932), Harper (1934), Wright and Wright
(1949), and personal observations (L. Vitt, J. Laerm). Most scientific
and all common names based on Collins et al. (1978). Aq = aquatic.
Ar = arboreal, F = fossorial, Sa = semi-aquatic, T = terrestrial.
T-Ar = terrestrial-arboreal.
SPECIES
CLASS AMPHIBIA
ORDER ANURA
Family Bufonidae
Bufo quercicus, Oak Toad
BuJo terresrris, Southern Toad
Family Hylidae
Acrisgryllusdorsalis. Florida Cricket Frog
Hyla chrysoscelis. Gray Treefrog
HABITAT
Okefenokee Swamp Vertebrates
SPECIES
Hyla cinerea cinerea, Green Treefrog
Hyla cruciferbarirnmiana, Southern Spring Peeper
Hyla fernoralis, Pine Woods Treefrog
Hyla gratiosa, Barking Treefrog
Hylasquirella, Squirrel Treefrog
Limnaoedus ocularis. Little Grass Frog
Pseudacrisnigrita nigrita, Southern Chorus Frog
Pseudacrisornata, Ornate Chorus Frog
Family Microhylidae
Gastrophiyne carolinensis, Eastern Narrow-mouthed Toad
Family Pelohatidae
Scaphiopur holbrooki holbrooki, Eastern Spadefoot Toad
Family Ranidae
~ o n aare olala aesopur. Florida Gopher Frog
Rona coresbeiana. Bullfrog
Rona clamitanr clonrirans. Bronze Frog
Rana grllio. Pig Frog
Ronn hpckscheri. R~kerF roe
Rana utrinrlnria; ~ o u t h e r n ~Ee o ~Farrodg
Rnna virgatipes, Carpenter Frog
ORDER CAUDATA
Family Ambystornatidae
Ambystoma cinwlatum. Flatwoods Salamande~
Amh~rromao pocum. Marbled Salamander
Amhysromo ralpord~um.M ole Salamander
Ambysroma tigrinwn. Tiger Salamander
Family Amphiumidae
Amphiwna means. Two-toed Amphiuma
Family Plethodontidae
Desmognathusfuscusauriculatur. Southern Dusky Salamander
Eurycen bislineata cirrigera, Southern Two-lined Salamander
Eurycea quadridigitata, Dwarf Salamander
Plethodonglutinosurglutinosur, Slimy Salamander
Pseudotriton montanurJ7oridanur. Gulf Coast Mud Salamander
Stereochilusmarginatus2, Many-lined Salamander
Family Salamandridae
Notophrhalmusperstriafus. Striped Newt
Norophthalamus viridescens louisianensis, Central Newt
ORDERTRACHYSTOMATA
Family Sirenidae
Pseudobranchusstriarurspp.', Dwarf Siren
Siren intermedia intermedia. Eastern Lesser Siren
Siren lacertina, Greater Siren
HABITAT
Ar
Ar
Ar
Ar
Ar
Sa
Sa
Sa
F, Sa
F
F
Aq
Aq
Aq
Aq
Sa
Sa
56 Joshua Laerm, et al.
SPECIES HABITAT
CLASS REPTlLIA
ORDER CROCODILIA
Family Alligatoridae
Alligaiormississippiensis, American Alligator Sa
Caiman scleropsd, Spectacled Caiman Sa
ORDER SQUAMATA
Family Anguidae
Ophisaurus aftenuafuslongicaudus, Eastern Slender Glass Lizard T
Ophisaurus compressus, Island Glass Lizard T
Ophisaurus ventralis, Eastern Glass Lizard T
Family lguanidae
AnoIis carolinemis. Green Anole Ar
Scelouorus undulatus undulatus. Southern Fence Lizard T-Ar
~amil~'~cincidae
Eumeces egregiussimilis, Northern Mole Skink F
Eumeces fasciafus, Five-lined Skink T-Ar
Eumeces inexpectatus, Southern Five-lined Skink T-Ar
Eumeces lariceps, Broad-headed Skink Ar
Scincella laterale, Ground Skink T
Family Teiidae
Cnemidophom sexfineatus sexlineatus, Six-lined Racerunner T
Family Colu bridae
Cemophora coccinea copei, Northern Scarlet Snake F
Coluber constrictor uriaous. Southern Black Racer T-Ar
Diad~phicpunrratu~~u~crarSuoru.t hern Ring-necked Snake T
Drymarchon corarscouperi. Indino Snake T
~ l u ~ghuteta rn guitata.-~orn~ n i k e
Elaphe obsoleta quadrivitiata, Yellow Rat Snake
Elaphe obsoletespiloides, Gray Rat Snake
Farancia abacura abacura, Eastern Mud Snake
Farancia er-v iroaramma. Rainbow Snake
H~r~rodonplaryrhinoFs.a stcrn HognoseSnske
fleterodon ~.imlriS, oulhcrn Iloenose Snake
LampropeIris caNigasfer rhomb~maculata. Mole Snake
Lnmuroueltisnetulur~etulus, Eastern Kinnsnake
~arnpro~el r i r~elu/usb. r ruxlJulonsd ana, intergrade k~ngsnake
Larnpropelrir rria~rgrrlume lapsoides. Sc3rlet Kingsnake
Muv1ic6nhi.i flaeelium ,fla n..e ilum. Eactern Coauhwhin
.~erodia'c)c~o~t>nJlorida~nal .i r i d Ga r een Water k a k e
.Verudia eryrhrogarterer)rhro~arrerR, ed-bellled Water Snake
Nerudia/asciara fasciara, Ronded Water Snakc
Nrrod~a/ascrarapiclivenrris.F lorida Water Snake
.\"Prodto laxispiioia. Brown Water Snake
Opheodrysaecri\w. Rough Green Snake
Pituophis~nelanoleucr~rmugiruFs.l orida Pine Snake
Rrqina al l~niS. triped Swamp Snake
Regina r;,ida rlgrda, EYS~I.C 1I3 1 ~ ~ 5U)', ,lel Sl~ake
Okefenokee Swamp Vertebrates 57
RhadinaeaJ7avilata. Pine Woods Snake T
Seminatrixpygaeapygaea, North Florida Black Swamp Snake
Storeria dekavi vicla. Florida Brown Snake
Aq
T
Sforeria occi~rromocr~laroub mrra, Florida Red-Rcllicd Sn;+ke T
Thamnoohissaurirussarkeni. Fastern Rlbbon Snake T
~ h amn o ~ hsiirr a1i.i rirralis, Eastern Garter Sn:*ke
Vi.r~iniaslriurulaR. ough Earth Snake
~i&inia valeriae va1eri;e. Eastern Smooth Earth Snake
Family Elapidae
Micrurus fulvius fulviw Eastern Coral Snake
Family Viperidae
Agkistrodon piscivorus conanti. Florida Cottonmouth
Croralusadamanteuf, Eastern Diamondback Rattlesnake
Crotalushorridus atricaudatuf. Canebrake Rattlesnake
Sistrurusmiliarius barbouri. Dusky Pigmy Rattlesnake
ORDER TESTUDINATA
Family Chelydridae
Chelydra serpenlinaserpentina, Common Snapping Turtle
Macroclemys femmincki, Alligator Snapping Turtle
Family Emydidae
Chrysemysnelsoni, Florida Redbelly Turtle
Deirochelvs reticularia reticularia. Eastern Chicken Turtle
- .
fseudemys i=~hr>semjsj scr&tascripta. Yellowbelly Slider
Terrapene carolina bauri, Florida Box Turtle
Terrapene carolina carolina, Eastern Box Turtle
Family Kinosternidae
Kinosternon bauripaimarum. Striped Mud Turtle
Kinosternonsubrubrum subrubrum. Eastern Mud Turtle
Stemotherusminor minor, Loggerhead Musk Turtle
Sfernotherus odorafus, Stinkpot
Family Testudinidae
Gopheruspolyphemus, Gopher Tortoi