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EDITORIAL STAFF 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, will appear at irregular intervals in consecutively numbered issues. Con-tents will emphasize zoology of the southeastern United Stittes, especially North Carolina and adjacent areas. Geographic coverage will be limited to Alabama, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, Virginia, and West Virginia. Subjcd matter will focus on taxonomy and systematics, ecology, zoo-geography, evolution, and behavior. Subdiscipline areas will include general in-vertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and paleontology. Papers will stress the results of original empirical field studies, but synthesizing reviews and papers of significant historical interest to southeastern zoology will be included. Suitability of manuscripts will be determined by the Editorial Board, and ap-propriate specialists will review each paper adjudged suitable. Final ac-ceptability will be decided by the Editor. Address manuscripts and all cor-mspondence (except that relating to subscriptions and exchange) to Editor, Brirnleyona, N. C. State Museum of Natural History, P. 0. Box 27647, Raleigh, NC 2761 1. & In cltatrons please use the full name - Brtmlejano NORTHC AROLINSAT ATEM USEUMOF NATURAHLI STORY NORTHC AROLINDAE PARTMENOFT AGRICULTURE JAMES A GRAHAMC,O MMISSIONER 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 .-6=- 5 c a 3 m 0 h .o ' o : C - 8 - v--, -- 3 -* c Q 0 o-u c -- x: . = 5 5 n rg mo .- 8 * E-i. 5C s s .=- ;-- 2 ; .:- z = o .:- 2 Q e .-G- c 3 + o 2 m .$- .-- L > r c i 0 Q - 5 c - .- - "3 3 m 2 Ci .c-o - 0 u c e! " --3 m 51 c2 ti -a n c 7- - -Y rf ??14""4m"O"""" - n o d o m-r o - o o o l r i -r r o m --o d d --o o o o m o - 2 r o X o d-t 0i 0 0 0 0 0 ~--4o ~IOo"" * 0 2"N?\a o o o c m ?4 0" -Pa -" -mmC---,.mNCrO* d G o o o-~ - ~ o < d o d N j-; ~ t i-i 2 d 4 ~ d ~ 2 4Y‘Dq 1 9?9?-. Y-g" "1 m N o - o o o ~ o o o o ~ m - " ~ O - ~ - * O a ~ m rn mm Y Y- 4 - ; 0 0 2 0 0 0 ~ 0 0 0 0 ~ " O C 1 0 0 d ~ o ~ 0 0 r 09- N"m"- --" *? Y ; d - o o o o o o o o o ~ ~ ~ d 4 0 0 ~ t i 4 0 m 0 m - - rn q c ? m ~ ~ ~ * s r ao a ~ d o o o - t i o o o o o o n ~ o m n a : d ~ - d < O 2 ?4- m -m - or., -a o o o o o o o o o o o ~ = o i,.o i r i o ~ + o o o o Y YIO Y ss-? ??":?- s o o o o - m o o o o o o ~ w aN~ o mN o ~ o o a o ~ 2 C .- 8 * B .s 5 0 P- - N 2 $ -c 3 -. - e2 o ;, a" z. x. ,2 sr., z0; 2 - 5 2 2 - W -,.. 0 z ' 2 9 .- - r. g - .3- + .s m B TI .- .- c, L .- u u h 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 -g c E 00 z-z Fd z 6 .. 3 '<:? .- u 5 z!: 0 0 2 : L L " = :z L L . D.0 o c WI . : rn .. = ?s z s n; m gz z, 0: 5 - .P m .a.- + x x X X X X X X X X X X '.XX X X X - ~ X X X X X X X X :.$ 3 B 5 .I-: 3 & E m6 , 3 e U .- Y O e- -c0 .- 3 i E3 C _ - P, r "'= 0_ Fm, x-- z .'-4 x x x x x x x - X X X X X X X x 2- 2 xx x x x x x x x x x x x x x x x x x x x x & z ; : K - E S ZE 2a3 m b ugk.i ~ 5 Y C -3mm . rs s - :: s? 6 .E - m 4 c.m > w, 2 zr. p rn x -.. .- m z. s .-c- E 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
Object Description
Description
Title | Brimleyana |
Contributor |
North Carolina State Museum of Natural Sciences. |
Date | 1981 |
Subjects |
Zoology--Southern States--Periodicals Ecology--Southern States--Periodicals Natural history--Southern States--Periodicals |
Place | North Carolina, United States |
Time Period | (1945-1989) Post War/Cold War period |
Description | Vol. 4;"The journal of the North Carolina State Museum of Natural History." |
Publisher | [Raleigh, NC : North Carolina State Museum of Natural History] |
Agency-Current |
North Carolina Department of Environmental Quality |
Rights | State Document see http://digital.ncdcr.gov/u?/p249901coll22,63754 |
Physical Characteristics | v. :ill. ;23 cm. |
Collection | North Carolina State Documents Collection. State Library of North Carolina |
Type | text |
Language | English |
Format | Periodicals |
Digital Characteristics-A | 17236 KB; 168 p. |
Digital Collection | North Carolina Digital State Documents Collection |
Digital Format | application/pdf |
Audience | All |
Pres File Name-M | pubs_serial_brimleyana1980v4.pdf |
Pres Local File Path-M | Preservation_content\StatePubs\pubs_serial_brimleyana\images_master |
Full Text | EDITORIAL STAFF 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, will appear at irregular intervals in consecutively numbered issues. Con-tents will emphasize zoology of the southeastern United Stittes, especially North Carolina and adjacent areas. Geographic coverage will be limited to Alabama, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, Virginia, and West Virginia. Subjcd matter will focus on taxonomy and systematics, ecology, zoo-geography, evolution, and behavior. Subdiscipline areas will include general in-vertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and paleontology. Papers will stress the results of original empirical field studies, but synthesizing reviews and papers of significant historical interest to southeastern zoology will be included. Suitability of manuscripts will be determined by the Editorial Board, and ap-propriate specialists will review each paper adjudged suitable. Final ac-ceptability will be decided by the Editor. Address manuscripts and all cor-mspondence (except that relating to subscriptions and exchange) to Editor, Brirnleyona, N. C. State Museum of Natural History, P. 0. Box 27647, Raleigh, NC 2761 1. & In cltatrons please use the full name - Brtmlejano NORTHC AROLINSAT ATEM USEUMOF NATURAHLI STORY NORTHC AROLINDAE PARTMENOFT AGRICULTURE JAMES A GRAHAMC,O MMISSIONER 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 .-6=- 5 c a 3 m 0 h .o ' o : C - 8 - v--, -- 3 -* c Q 0 o-u c -- x: . = 5 5 n rg mo .- 8 * E-i. 5C s s .=- ;-- 2 ; .:- z = o .:- 2 Q e .-G- c 3 + o 2 m .$- .-- L > r c i 0 Q - 5 c - .- - "3 3 m 2 Ci .c-o - 0 u c e! " --3 m 51 c2 ti -a n c 7- - -Y rf ??14""4m"O"""" - n o d o m-r o - o o o l r i -r r o m --o d d --o o o o m o - 2 r o X o d-t 0i 0 0 0 0 0 ~--4o ~IOo"" * 0 2"N?\a o o o c m ?4 0" -Pa -" -mmC---,.mNCrO* d G o o o-~ - ~ o < d o d N j-; ~ t i-i 2 d 4 ~ d ~ 2 4Y‘Dq 1 9?9?-. Y-g" "1 m N o - o o o ~ o o o o ~ m - " ~ O - ~ - * O a ~ m rn mm Y Y- 4 - ; 0 0 2 0 0 0 ~ 0 0 0 0 ~ " O C 1 0 0 d ~ o ~ 0 0 r 09- N"m"- --" *? Y ; d - o o o o o o o o o ~ ~ ~ d 4 0 0 ~ t i 4 0 m 0 m - - rn q c ? m ~ ~ ~ * s r ao a ~ d o o o - t i o o o o o o n ~ o m n a : d ~ - d < O 2 ?4- m -m - or., -a o o o o o o o o o o o ~ = o i,.o i r i o ~ + o o o o Y YIO Y ss-? ??":?- s o o o o - m o o o o o o ~ w aN~ o mN o ~ o o a o ~ 2 C .- 8 * B .s 5 0 P- - N 2 $ -c 3 -. - e2 o ;, a" z. x. ,2 sr., z0; 2 - 5 2 2 - W -,.. 0 z ' 2 9 .- - r. g - .3- + .s m B TI .- .- c, L .- u u h 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 -g c E 00 z-z Fd z 6 .. 3 '<:? .- u 5 z!: 0 0 2 : L L " = :z L L . D.0 o c WI . : rn .. = ?s z s n; m gz z, 0: 5 - .P m .a.- + x x X X X X X X X X X X '.XX X X X - ~ X X X X X X X X :.$ 3 B 5 .I-: 3 & E m6 , 3 e U .- Y O e- -c0 .- 3 i E3 C _ - P, r "'= 0_ Fm, x-- z .'-4 x x x x x x x - X X X X X X X x 2- 2 xx x x x x x x x x x x x x x x x x x x x x & z ; : K - E S ZE 2a3 m b ugk.i ~ 5 Y C -3mm . rs s - :: s? 6 .E - m 4 c.m > w, 2 zr. p rn x -.. .- m z. s .-c- E 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 |
OCLC Number-Original | 4904283 |