INFORMATION TO USERS
The most advanced technology has been used to photo graph and reproduce this manuscript from the microfilm master. UMI films the original text directly from the copy submitted. Thus, some dissertation copies are in typewriter face, while others may be from a computer printer.
In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyrighted material had to be removed, a note will indicate the deletion.
Oversize materials (e.g., maps, drawings, charts) are re produced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each oversize page is available as one exposure on a standard 35 mm slide or as a 17" x 23" black and white photographic print for an additional charge.
Photographs included in the original manuscript have been reproduced xerographically in this copy. 35 mm slides or 6"X 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order.
[re Accessing the World’s Information since 1938
300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA
Order Number 8820372
Revision of the genusPseudopolydesmus Attems, 1898 and its relationships to tlie North American genera of the family Polydesmidae Leach, 1815
Withrow, Charles Phillip, Ph.D.
The Ohio State University, 1988
UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106
PLEASE NOTE:
in all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark V
1. Glossy photographs or pages.
2. Colored illustrations, paper or _____print
3. Photographs with dark background____
4. Illustrations are poor copy______
5. Pages with black marks, not original copy.
6. Print shows through as there is text on both sides of page.
7. Indistinct, broken or small print on c-evera!___ pages
8. Print exceeds margin requirements_____
9. Tightly bound copy with print lost______in spine
10. Computer printout pages with indistinct print,
11. Page(s)______lacking when material received, and not available from school or author.
12. Page(s)______seem to be missing in numbering only as text follows.
13. Two pages num bered . Text follows.
14. Curling and wrinkled pagest/^ 15. Dissertation contains pages v/ith print at a slant, filmed as received/
16. Other ______
REVISION OF THE GENUS PSEUDOPOLYDESMUS ATTEÎ'ÎS, 1898
AND ITS RELATIONSHIPS TO THE NORTH
AMERICAN GENEE^A OF THE FAMILY
POLYDESMIDAE LEACH, 1815
Presented in partial fulfillment of the Requirements for
the Degree Doctor of Philosophy in the Graduate
School of The Ohio State University
By
Charles Phillip Withrow, B.S., M.S.
*****
The Ohio State University
1988
Reading Committee; pproved By
Barry D. Valentine
John L. Crites Advisor
David H. Stansbery Department of Zoology
Charles A. Triplehom Copyright by
Charles Phillip Withrow
1988 To My Parents ACKNOWLEDGEMENTS
I would like to thank my advisor during my career at Ohio
State University, B a n y D. Valentine, for his advice, course assistance, counseling, stimulation, and understanding.
I would also like to thank Dr. Richard L. Hoffman, of Radford
University for the ej^sure and introduction to the interesting field of diplopodology and for suggesting this topic.
The stimulation, comradery, and help of graduate students in both departments of zoology and entomology has been well appreciated.
I would like to thank the Department of Zoology and the
Graduate School of Ohio State University for their continued financial stroport t h r o u ^ toy studies.
Finally I would like to acknowledge the support of my parents,
Robert C. and Jean P. Withrow, for their tolerance of ity early adventures into science. Their guidance and support of ïty decisions in life is unrepayable.
11 VITA
March 13, 1955 ...... B o m - Gallipolis, Ohio
1977 ...... B. S., Zoology, Marshall University, Huntington, West Virginia
1979 ...... M.S., Biological Sciences, Marshall University, Huntington, West Virginia
1979 ...... Teaching Associate, Department of Biological Sciences, Marshall University, Huntington, West Virginia
1981-87 ...... Teaching Associate, The Ohio State University, Columbus, Ohio
HJBIICATIONS
1978 Millipedes of Cabell County, West Virginia. W. Va. Acad. Sci. 38:48-50.
FIELDS OF STUDY
Major Field: Diplopoda Systzematics and Biogeography
1 1 1 TABLE OF CONTENTS
Page
DEDICATION...... i
ACKNOWLEDGEMENTS...... ii
VITA...... iii
LIST OF TABLES...... vi
LIST OF FIGURES...... viii
LIST OF MAPS...... xii
INTRODUCTION...... 1
MATERIALS AND METHODS...... 6
Materials...... 6
Generalized Methods...... 7
Phylogenetic and Classification Methods...... 10
RESULTS...... 15
Family Doratodesmidae...... 16
Family Polydesmidae...... 19
Subfamily Mastigonodesminae...... 21
Key to North American Genera...... 24
Subfamily Scytonotinae...... 26
Speodesmus...... 28
Scytonotus...... 32
Utadesmus...... 37
Bidentoaon...... 41
iv Coronodesmus...... 45
Sipeorthus...... 48
Idahodesmus...... 51
Siabfamily Polydesminae...... 55
Polvdesmus...... 57
Pseudopolydesmus...... 64
Subfamily E^anerchodinae...... 126
CLADISTIC ANALYSIS...... 129
Character States...... 129
Classification of Polydesmidae...... 166
ZOOGBOGRAmY...... 167
The Vicariance Model...... 167
Distribution Patterns...... 169
Historical Zoogeography...... 172
BIBLIOGRAHiY...... 173
APPENDIX A: TABLES...... 188
APPENDIX B: FIGURES...... 205
APPENDIX C: MAPS...... 254
APPENDIX D: IDCALETY DATA...... 264 U S T OF TABLES
Tables Page
1. Systematics of the Family Polydesmidae...... 189
2. List of generic character states for the Family Polydesmidae...... 190
3. Data set for the Family Polydesmidae and genera of Scytonotinae...... 192
4. Manhattan distance matrix for Family Polydesmidae...... 194
5. Patristic distance matrix for the Family Polydesmidae...... 195
6. Hamoplasy matrix for the Family Polydesmidae... 196
7. List of specific character states for the genus Pseudopolydesmus...... 197
8. Data set for genus Pseudopolydesmus...... 198
9. Length and width measurements of Pseudopolydesmus species, males and females, with range, mean, standard deyiation, and number measured...... 199
10. W/L (width/length) and WL (width X length) measurements of Pseudopolydesmus species, males and females, with range, mean, standard deyiation, and number measured...... 200
11. Distance from telopodite tip to base of mesial and ectal (E^-^) processes as a percentage of total telopodite length...... 201
12. Manhattan distance matrix for the species in the genus Pseudopolydesmus...... 202
13. Patristic distance matrix for the species in the genus Pseudopolydesmus...... 203
yi 14. Hamoplasy matrix for the species in the genus Pseudopolydesmus...... 204
Vll U S T OF FIGURES
Figures Page
1-3. Dorsum, dorsal view, male. 1. Cerastelachvs cavemicola; 2. Mastigonodesmus vignai; 3. Epanerdhodus bidens...... 206
4-6. Dorsum, dorsal view, male. 4. Speodesmus echinourus; 5. Scytonotus granulatus; 6. Utadesmus henriensis...... 207
7-9. Dorsum, dorsal view, male. 7. Bidentogon helferorum; 8. Coronodesmus bituberculatus; 9. Speorthus tugahbius...... 208
10-12. Dorsum, dorsal view, male. 10. Idahodesmus dentatus; 11. Polvdesmus inconstans; 12. Pseudopolydesmus serratus...... 209
13-15. R i ^ t gonopod, lateral view. 13. Cerastelaolivs cavemicola : 14. Mastigonodesmus vignai; 15. Epanerdhodus bidens...... 210
16-18. left gonopod of Speodesmus edhinourus. 16. medial view; 17. lateral view; 18. ventral view...... 211
19-20. Dorsal view. 19. Pseudopolydesmus erasus, male; 20. Scytonotus granulatus male and female 212
21-23. R i ^ t gonopod of Scytonotus granulatus. 21. medial view; 22. lateral view; 23. ventral view...... 213
24-26. R i ^ t gonopod of Utadesmus henriensis. 24. medial view; 25. lateral view; 26. ventral view...... 214
27-29. R i ^ t gonopod of Bidentogon helferorum. 27. medial view; 28. lateral view; 29. ventral view...... 215
30-32. R i ^ t gonopod of Coronodesmus bituberculatus. 30. medial view; 31. lateral view; 32. ventral view...... 216
viii 33-35. R i ^ t gonopod of Speorthus tugahbius. 33. medial view; 34. lateral view; 35. ventral view...... 217
36-39. Right gonopod of Idahodesmus dentatus. 36. medial view; 37. lateral view; 38. ventral view; 39. sternum III, posterior view, female.. 218
40-42. Ric^t gonopod of Polvdesmus inconstans. 40. medial view; 41. lateral view; 42. ventral view...... 219
43. Cladogram for the Family Polydesmidae including the genera of Scytonotinae...... 220
44-45. Head and tergites 1-4, dorsal view. 44. Scytonotus granulatus; 45. Pseudopolydesmus serratus...... 221
46-47. Diplosegment lateral view, midboc^. 46. Pseudopolydesmus serratus; 47. Scytonotus granulatus...... 222
48-49. Diplosegment, cross section, midbody. 48. Pseudopolydesmus serratus; 49. Scytonotus granulatus...... 223
50-52. Dorsal setae, lateral view. 50. Pseudopolydesmus caddo; 51. Bidentogon helferorum; 52. Speodesmus echinourus...... 224
53-54. Legs, posterior view, male. 53. Utadesmus henriensis. ri^t, third leg; 54. Scytonotus granulatus. right posterior leg, 13th segment, also ventral view...... 225
55-56. Claw, posterior view. 55. Pseudopolydesmus canadensis; 56. Scytonotus granulatus...... 226
57-58. Antennae, anterior view. 57. Pseudopolydesmus serratus; 58. Scytonotus granulatus...... 227
59-60. Midbody sternum, ventral view. 59. Pseudopolydesmus serratus; 60. Scytonotus granulatus...... 228
61-62. Posterior sternum of 7th segment, posterior view, male; 61. Pseudopolydesmus canadensis; 62. P. minor...... 229
IX 63-64. Synœxostemtrai, posterior view. 63. Pseudopolydesmus serratus; 64. Scytonotus granulatus...... 230
65-66. Sternum III, posterior view, female. 65. Polvdesmus inconstans; 66. Pseudopolydesmus erapnis...... 231
67-68. Cÿphopod, lateral view. 67. Pseudopolydesmus serratus; 68. Scytonotus granulatus...... 232
69. Midbody leg of Pseudopolydesmus serratus. anterior view, male and female, semidiagrammatic, setae emitted...... 233
70-73. Right gonopod of Pseudopolydesmus canadensis, male; 70. mesal view; 71. lateral view; 72. ventral view; 73. in situ...... 234
74-77. Dorsum, dorsal view, midbody. 74. Pseudopolydesmus pinetorum; 75. P. tallulanus; 76. P. canadensis; 77, P. caddo...... 235
78-79. Female sternum III, ventral view. 78. Pseudopolydesmus canadensis; 79. P. minor 236
80-81. Ip>iproct, lateral view. 80. Pseudopolydesmus pinetorum; 81. P. serratus...... 237
82-85. Right gonopod of Pseudopolydesmus spp., Groiç) canadensis, lateral view. 82. P. pinetorum; 83. P. tallulanus; 84. P. erasus; 85. P. collinus.. 238
86-89. R i ^ t tibiotarsus of Pseudopolydesmus spp., Group canadensis, medial view. 86. P. pinetorum; 87. P. tallulanus ; 88. P. erasus; 89. P. collinus...... 239
90-93. R i ^ t tibiotarsus of Pseudopolydesmus spp., Groiç) canadensis, ventral view. 90. P. pinetorum; 91. P. tallulanus; 92. P. erasus; 93. P. collinus...... 240
94-97. R i ^ t gonopod of Pseudopolydesmus spp., Groiç» serratus, lateral view. 94. P. serratus; 95. P. paludicola; 96. P. caddo; 97. P. minor..... 241
98-101. R i ^ t tibiotarsus of Pseudopolydesmus spp., Group serratus, medial view. 98. P. serratus; 99. P. paludicola; 100. P. caddo; 101. P. minor 242
X 102-105. R i ^ t tibiotarsus of Pseudopolydesmus spp., Groiç) serratus, ventral view. 102. P. serratus; 103. P. paludicola; 104. P. caddo; 105. P. minor...... 243
106-110. R i ^ t cyphcpod of Pseudopolvdesmus spp Grotp canadensis, lateral view. 106. P pinetorum; 107. P. tallulanus; 108. P. erasus; 109. P. canadensis; 110. P. collinus.. 244
111-115. R i ^ t cypbopod of Pseudopolvdesmus spp Groiç) canadensis, ventral view. 111. P pinetorum; 112. P. tallulanus; 113. P. erasus; 114. P. canadensis; 115. P. collinus.. 245
116-117. R i ^ t cyphcpod of Pseudopolvdesmus spp., Groijp serratus, lateral view. 116. P. caddo; 117. P. minor...... 246
118-120. R i ^ t cyphopod of Pseudopolvdesmus spp., Group serratus, ventral view. 118. P. caddo; 119. P. minor; 120. P. serratus..... 247
121. Cladogram for the genus Pseudopolvdesmus...... 248
122. Graph of length measurements of Pseudopolvdesmus species, male and female 249
123. Graph of width measurements of Pseudopolvdesmus species, male and female 250
124. Graph of W/L measurements of Pseudopolvdesmus species, nale and female...... 251
125. Graph of IW measurements of Pseudopolvdesmus species, male and female...... 252
126. Graph of distances from telopodite tip to bases of mesial (Mi_4 ) and ectal processes as a percentage of total telopodite length...... 253
XI LIST OF MAPS
Maps Page
1. Distribution of families Polydesmidae and Doratodesmidae...... 255
2. Distribution of Scytonotinae...... 256
3. Distribution of Pseudopolvdesmus...... 257
4. Distribution of Pseudopolydesmus pinetorum 258
5. Distribution of Pseudopolvdesmus tallulanus and P. erasus...... 259
6. Distribution of Pseudopolvdesmus canadensis— . 260
7. Distribution of Pseudopolvdesmus collinus 261
8. Distribution of Pseudopolvdesmus serratus 262
9. Distribution of Pseudopolydesmus paludicola. P. caddo and P. minor...... 263
Xll INTRODÜCnON
The family Polydesmidae is a moderately large milliped taxon of the Superfamily Polydesmoidea in the Order Polydesmida. The family has a Holarctic distribution encortpassing 24 genera and approximately 220 species. The systematics of the family is presently in a state of disorder, primarily because of a lack of investigators; this is especiailly true of the Palearctic fauna vÆiich will not be addressed in this study. One Palearctic genus,
Polvdesmus Latreille, 1802, vdiich has a wide distribution west of central Asia, is presently composed of 26 subgenera, and hundreds of taxa, including many subspecies and forms. In several cases, taxa are distinguished only by the number of segments. Only in western
Europe is there any degree of comprehension in the classification.
The North American fauna is only moderately better understood, chiefly due to fewer taxa. The first modern listing of the North
American taxa by Chamberlin and Hoffman (1958) had 11 genera and 50 species. According to Hoffman's revised classification (1979), this has decreased to 6 genera and 35 species. This decrease was 2 primarily caused by the réévaluation of the works of Richard V.
Chamberlin vho, thro u ^ the 1930's and 40's, described a large number of taxa, over half of %hich are now known to be synonyms. The genera of Polydesmidae listed by Chamberlin and Hoffman are;
Antriadesmus Loomis, 1943, Brachvdesmus Heller, 1858, Chaetaspis
Bollman, 1887, Dixidesmus Chamberlin, 1943, Polvdesmus Latreille,
1802, Pseudopolydesmus Attems, 1898, Scytonotus Koch, 1847,
Speorthus Chamberlin, 1952, Speodesmus Loomis, 1939, Tidesmus
Chamberlin, 1943, and Utadesmus Chamberlin and Hoffman, 1950.
Hoffman (1979) made the following changes: Chaetaspis and Tidesmus were placed in the siçerfamily Trichopolydessmoidsa; Chaetaspis in the family Macrostemodesmidae, and Tidesmus in the family
Trichopolydesmidae. He also suggested the following new synonyms:
Chaetaspis f=Antriadesmus), Pseudopolvdesmus (=Dixideanus), and
Polvdesmus f =Brachvdesmus^. Also the genus Bidentogon (Buckett &
Gardner, 1968), which was originally placed in the family
Fuhrmannodesmidae, was included.
As of this stucty, the North American genera of the family
Polydesmidae are: Pseudopolvdesmus. with 9 ^)ecies; Scytonotus. with
11 species; Speodesmus with 7 species; Speorthus. with 1 species;
Utadesmus. with 2 species; Bidentogon. with 2 species; Coronodesmus new genus, with 2 species; and Idahodesmus new genus, with 1 species. Also, the Old World genus Polvdesmus is now well established in the Nearctic, with 7 species.
This paper deals with the taxonomic problems present in the dominant eastern North American genus Pseudopolvdesmus. and with the 3 interrelations of all the Holarctic genera. In the past, haphazard definitions of taxa at hi^er levels and the multiplication of taxonomic species has led to many misidentified forms.
The family Polydesmidae was erected by leach in 1815. Prior to
1898, viien the generic name Pseudopolvdesmus was erected 1:^ Attems, all North American polydesmids were placed in the externally similar
Palearctic genus Polvdesmus. The generotype for Pseudopolvdesmus is
Polvdesmus canadensis Newport, 1844, by monotypy. From this early beginnir^ the taxonomic position of Pseudopolvdesmus underwent few major changes, although generic associations came and went with increased understanding of milliped systematics.
Chamberlin and Hoffman (1958) list 28 names in the genus
Pseudopolvdesmus; there are now over 30. Despite this accumulation, no revisional stu(^ has been attenpted. The names are; canadensis
(Nevport, 1844) ; pensvlvanicus [sic] and alaucescens (Koch, 1847) ; branneri and nitidus (Bollman, 1887) ; minor and pinetorum (Bollman,
1888); euthetus. paroicus. planicolens. scopus. neoterus. echinogon. and natchitoches (Chamberlin, 1942); hubrichti. svlvicolens. penicillus. modocus. humilidens. conlatus. and tallulanus.
(Chamberlin, 1943); erasus (Loomis, 1943); christianus (Chamberlin,
1946) ; catskillus (Chamberlin, 1947) ; caddo Chamberlin, 1949; paludicolus [sic] Hoffman, 1950; phanus (Chamberlin, 1951); qausodicrorhachus (Johnson, 1954); bidens. Loomis, 1959; and collinus Hoffman, 1973. Mich suspected synonyity can be blamed on
Chamberlin. The general opinion is that he rarely if ever examined types. A second problem, shared by almost all milliped taxa below 4
the generic level, is the lack of distinctive iroiphological
characters with the exception of the male gonopods. A third more
technical problem deals with labelling and relative positioning of
the gonopodal processes. Depending on the orientation of the
drawing, processes are hidden or appear to change location. Hoffinan
(1973) addressed the latter problem by setting vç) a gonopodal
formula for the naming and identification of the processes. In the
same paper he stated, without further elaboration, that in
Pseudopolvdesmus many species could be synonymized.
Very little has been done with the ecology of these millipede.
The only Nearctic polydesmoid millipeds studied are a species of
Eurvurus. of the family Xystodesmidae by Miley (1927), and the life history and ecology of Orthomorpha gracilis (Roch), of the family
Strongylosomatidae by Causey (1943). Other ecological studies only
include species in habitat lists (Bailey, 1919, Branson & Batch,
1971; Shelley, 1978; Withrow, 1978; etc.). In eastern North America, the genus Pseudopolvdesmus is the most common native polydesmid.
Individuals are usually found under decaying leaves and loose bark throughout the year, but are common during spring and fall. They can easily be obtained by hand collecting, sifting, or using a Berlese
funnel. This is because millipeds are rarely seen by the typical nature observer, they are economically unimportant, difficult to work with, and (to many) just plain unexciting. They are however very important ecologically in breaking down detritus, thereby returning nutrients to the ecosystem. A habitat study (O'Neil, 1967) dealt with preferences of millipeds in different maple-oak 5 microhabitats in central Illinois. He states that 66.7% of his sairple size Pseudopolvdesmus serratus occurs under siçierficial wood of logs and 20.8% on the outer surface of logs under the bark. Based on my observations^ O'Neil's data are not typical over the wide geographical range of the genus. Individuals are found in most niches althou^ dependent on tenperature and humidity. Mass migrations of Pseudopolvdesmus serratus have been observed (Ramsey,
1966). Essentially nothing is known about growth, behavior, and metabolism of North American taxa. There is information on the secretions of Pseudopolvdesmus species by many workers, summarized by Blum (1981). Pseudopolvdesmus species have been found to produce the typical, polydesmoid products such as formic, isovaleric, acetic, and benzoic acids, mandelonitrile benzoate and hydrogen cyanide.
Because of the phylogenetic closeness of Pseudopolvdesmus to
Polvdesmus. studies of European taxa may be informative; for example. Blower (1970) looked at growth rates, Kime and Wauthy
(1984) examined habitat preference based on soil coiposition, and
Baker (1978) investigated population dynamics. MMERIAIS AND METHODS
MATERIALS
This stu£^ was based on the examination of over 8,000 specimens borrowed from the following institutions and individuals.
The abbreviations used in the text are also included. The listing of specimens can be found in ^pendix D.
AMNH American Maseum of Natural History, New York, Dr. Norman Platnick.
ANSP Academy of Natural Sciences, Lhiladelphia, Mr. Donald Azuma.
EMNH British Museum of Natural History, London, Dr. Paul Hillyard.
d W C author, personal collection.
EMNH Dayton Museum of Natural History, Ohio, Mr. Gary A. Coovert.
LMNH Field Museum of Natural History, Chicago, Dr. John Retliley.
ESCA Florida State Collection of Arthropods, Gainesville, Dr. G. B. Edwards. MCZC Museum of Ctonparative Zoology, Harvard University, Cambridge, Massachusetts, Dr. H. W. Levi.
MHNG Mjseum d'Histoire naturelle, Geneve, Switzerland, Dr. Hauser.
MCJIC Marshall University Invertebrate Collection, Huntington, West Virginia, Dr. Donald Tarter.
NCMH North Carolina Museum of Natural History, Raleigh, Dr. EolarKÎ M. Shelley.
OSEM Ohio State University Collection of Insects and Riders, Columbus, Dr. Charles Triplehom.
OSZM Ohio State University Museum of Zoology, Columbus, Dr. Carol Stein.
RIHC Dr. Richard L. Hoffitian, Radford University, personal collection.
IMMC Texas Memorial Miseum, UNiversity of Texas, Austin, Dr. James Reddell.
USNM National Maseum of Natural History, Washington D.C., Dr. John Coddington.
WASC Dr. William A. Shear, Haitpton-Sydney College, personal collection.
In addition, I have drawn heavily on the literature, especially for familial characters.
GENERALIZED METHODS
Several morphological structures need to be defined. The paranotum(a) is the lateral projection of a tergite or diplosegment.
The gnathochilarium is the fused labium and maxillae. The most diagnostic tool in milliped systematics is goncpod morphology. The 8
gonqpods are specially rccdified legs (in polydesmids, the anterior
pair of the seventh diplosegment) used for the transfer of sperm
from male to female. They are large, easily removable, species
specific, and provide numerous characters (Fig. 70). The gonopod
components are the gonocoxae, prefetmor, femur, and tibiotarsus. The
last is probably the fused tibia and tarsus of the original leg. The
term telqpodite refers to the prefemur, femur, and tibiotarsus,
Vihile the acrcpodite is the terminal, non-grooved structure. On the
medial surface of the femur, a femoral groove extends anteriorly
from under the endomerite. The endomerite is a hirsute projection
distal to the external openim of the seminal groove on the
posterior surface. Within the femur there is a seminal groove vhich
may be enlarged into a seminal vesicle. The tibiotarsus may have
various ^ines or processes.
In females, the cyphopods (Fig. 67) are the corresponding
sperm receiving structures, located between the second and third
segments. These paired structures were examined althou^ they are
usually internal, small, and not as diagnostic as the gonopods. In
the polydesmids, the cyphopods are composed of a pair of large valves margined with macrosetae. lateral and medial setae are also
present. A small sperm receptacle is located on the posterior edge.
The venter of the second segment excluding its single pair of legs,
is referred to as the ^ncoxostemum (Fig. 63).
Specimens were examined in 70% isopropyl alcohol using an AO
Spencer binocular microscope with 15X to 9 OX magnification and
incident light. Gonopods were positioned on obsidian sand for easier 9
observation. Whole body measurements were determined by mechanical
calipers (accurate to 0.1mm), viiile smaller measurements (such as
antennomeric and podcmeric lengths) were obtained with an ocular
micrometer at 90X (accurate to 0.01mm). Individual components (ex.
gnathochilarium) were sometimes cleared with 10% KOH or typsin,
temporarily slide mounted, and examined with an Olympus compound
microscope using transmitted light. Surface sculpturing and setae
were examined with specimens temporarily removed from alcohol and
allowed to air dry. Specimens for scanning electron microscopy were
dissected and ultrasonically cleaned, mounted on stubs, allowed to
air dry,- and then coated with gold, and pallidium. Specimens were
examined with a Hitachi S-500 SEM.
A gonopod, usually the riiÿit one, was dissected from the
seventh segment with watchmakers forceps and hooked minuten pins,
cyphopods were usually studied by first breaking the specimens between the 2nd and 3rd segments, vhich also permitted examination
of the syncoxostemum and sternum III.
Gonopodal measurements proved especially difficult because of the highly variable nature of telopodite curvature. Total length was measured in lateral view along the curve from the tip of the
acropodite to the distal tip of the prefemur. A series of chordal measurements was taken from the base of one process to the next proximally. These were used to determine the distance of each process from the tip and combined to obtain total telopodite length.
Three or four repetitions of each gonopod measurement were recorded and averaged for each of 10 specimens. The process lengths were 10 measured from mi(%ioint on the telqpodite to the distal tip, in lateral or medial view; depending on visibility. The lateral view was more accurate for process E 2 because of process orientation.
When possible, specimens from all areas and geographic extremes were used. These measurements are moderately reliable and provide an idea of relative location and degree of variability. Table 11 gives sizes and lengths based on percentage of telopodite length. The process formula follows Hoffman (1974).
Figures were drawn with the binocular microscope fitted with a
10 X 10 ocular reticle.
ÎHYIÛGENEITC AND CLASSIFICATION METHODS
Except vdien dealing with asexual organisms, a species can be defined as a population of interbreeding individuals vdiich gives rise to viable offering vdiile maintaining their similarity and relatedness. Therefore breeding and reproductive data are the best and most conclusive l^pes of information for determining spéciation.
Since this information is rarely available, most mo d e m systematics is based on morphological similarities and differences between species.
This study is based on the principles of cladistics and phylogenetic systematics developed by Hennig (e.g. 1966). Wiley
(1981) goes into great detail on phylogenetic theory and application. Cladistic studies are virtually unknown in milliped systematics. Investigations by Enghoff (1981, 1984) represent the 11 only major works errploying cladistic techniques. Most milliped systematists (e.g. Hoffianan, 1979) still employ the evolutionary systematics of Sitrpson and Mayr (e.g. Mayr, 1969).
Phylogenetic systematics is based on the theory of moncphyly.
A monophyletic taxon is defined as a groip of Recent species, ihich includes all descendants and only the descendants of a single ancestral species. These taxa are indicated by autapomorphies
(uniquely derived similarities) shared by their members. Also, there is a decreased importance placed on convergences and parallelism (or homoplasies), vhich are independently derived similarities.
Syitplesiomorphy is a state vdiere similarities inherited from a single ancestral species occur in more than one taxon.
The major problem of phylogenetic systematics is the determination of the polarity of character states between the plesicmorphic or primitive state and apomorphic or derived state.
Characters consist of a groip of states considered variants of the same thing, usually morphological structures. Character states are the observed conditions in a taxon or grotp of taxa. Jong (1980) details how to determine character polarity. The most frequently used method is by outgroip comparison. The polarily of a character vhich is shared with the outgroup (the believed closest relative) is considered plesicmorphic. Another way of stating this is that the single shared ancestor possessed the same character state observed in the primitive members of each taxon. Each taxon is then examined to determine the plesicmorphic or apomorphic status of each character. ^ convention, the data sets utilize 0 to define the 12 plesicmorphic state vÆiile a 1 designates the apomorphic state. Data sets for the taxa and their states are set for use in conputer analysis for obtaining phylogenetic trees. These dichotomous branching trees, commonly referred to as cladograms, are manufactured using parsimonious methods to develop the sirrplest and most conservative patterns. This reduces the number of observed homoplasies. These trees are then used to set vp a classification scheme.
The Siperfamily Polydesmoidea includes five families,
Polydesmidae, Haplodesmidae, Doratodesmidae, Opisotretidae, and
Cryptodesmidae, with the latter being the most distantly related to the others. Hoffman (1979) stated that the Family Doratodesmidae was the closest relative to the Family Polydesmidae, based on overall gonopodal similarities. I have accepted this opinion and have used the Family Doratodesmidae as the outgroup for the Family
Polydesmidae vdien determining plesiomorphic states. The outgroip for
Pseudooolvdesmus was determined to be the genus Polvdesmus. The characters examined and their states are listed on Tables 1 & 2.
Data sets in this stuc^ are listed in Appendix A and the actual cladograms are given in Appendix B.
The trees generated in this stuc^ were obtained by PAUP
(Phylogenetic Analysis Using Parsimony) version 2.4, developed by
David L. Swofford. This study enploys Wagner parsimony. The assunptions of Wagner parsimony are: 1, ancestral states are unknown; 2, independent evolution of characters; 3, independent evolution of different lineages; 4, changes 0>1 are as probable as 13
1>0; 5, both types of charges are a priori irrprobable over time; 6,
evolutionary events such as retention of polymorphism are less
probable than 0>1; 7, that evolution rates are slow enouÿi that two
changes over a long period of tine are less probable than one change
over a short period of time (Felsenstein, 1981).
In determining the most parsimonious tree a nurnber of analysis
tests were performed throu^ PAUP, the most important measurement
being tree length. The calculation of tree length is the summation
of all OIUs (operational taxonomic units) or taxa differences between each node of the generated tree. The generation of the trees was ly the use of closest selection procedure. Each currently urplaced OTU was added, in turn, to every possible position on a
tree, and the length required for each of these new trees computed.
The OTU whose connection added the least to the additive length of the existing tree is then chosen for the next addition. The parameters used in obtaining tree length were; the use of Farris optimization (Farris, 1970) ; the root was set as the outgroup,
localized branch swapping with the generation of 50 individual trees; and no character state weighting.
The remaining calculations were derived from the numbers generated for three matrices. The first was the Manhattan distance matrix.
The second matrix was the patristic or path-length distance matrix. The patristic distance is the sum of the intemodal distance connection between two pairwised endpoints (OTUs) divided by the number of characters examined. Missing data were calculated as the 14
mean distance. This matrix was used for the computation of the two
f-values. The f-value of Farris (1972) is the sum of all deviations
of the path-length across distant pairs of taxa. When there are no
missing data this number is equal to the homcplasy of Farris' 78
program. A normalized f-value (f (n) ) was also calculated. It was
derived by the division of the f-value (Farris) by the sum of
pairwise Manhattan distances. This is the same as the deviation
ratio of Wagner 78. The f-values give comparative values as to the
degree of parisomy in the tree. The lower the value the hi^er the
degree of parisomy. Values tend to be lower in larger trees due to
the 'usual' increase of homoplasy. These measurements can help to
identify modified or miscoded characters, and also identify highly
variable or missing OTOs.
The third matrix deals with homoplasy. For each pair of taxa,
homoplasy is equal to the patristic distance minus the character
difference (Manhattan distance). Missing data are replaced by the
mean patristic distance. The lower each value, the lower the degree
of homoplasy.
A final calculation is the generation of the Consistency Index
(Cl), defined by Kluge and Farris (1969). The consistency index is
the sum over all characters of the 'range' of each character
(equivalent to the minimum tree length computed for that character
only) divided by the total tree length for all characters. This is a neasurement of the deviation of the tree from a perfect fit to the
data. The range of Cl is between 0 and 1, with 1 having no 15
œnvergencies. Therefore, the hi^er the index the more parsimonious the tree.
TWO additional tests were examined. These dealt with a variation of the Wagner parsimony. The first was DELTEAN or delayed transformation optimization. The running does not change the tree length, endpoints, or the consistency index, it can change the maJœtç) of the intemodal placement of characters. The test optimizes the ratio of parallelism to reversal, it favors two 0>1 changes over
0>1 change followed by a 1>0 change. This tends to reduce the f- value and increase the number of convergences and parallelisms.
The second test is the reverse of DEdRAN, or ACCTRAN, accelerated transformation optimization. Here the ratio of reversal is maximized.
The programs were run on an IBM PC portable. RESUETS
To understand the North American polydesmid fauna, an analysis of the family was needed to determine relationships of the major taxonomic units. Due to the inadequate descriptions, vdiich are prevalent in milliped systematics, an effort was made to describe those taxa not being revised. These descriptions at the family and subfamily levels were based on literature and specimens examined.
Because this represents the first revision of the genus
Pseudopolvdesmus. full morphological descriptions were included.
FAMILY DORATODESMIDAE Cook, 1896
(Figs. 1, 13; Map 1)
Doratodesmidae Cook, 1896. Brandtia 5:19.
DIAGNOSIS: Small to medium (4 to 25mm). Segments numbering 19 or 20. Capable of involution. Length to width ratio 15-32%. Tergal
16 1 7 width to h e i ^ t ratio 1.3-1.45. Body width to height ratio 1.1-1.25.
Brown, yellow or vdiite. Head mostly smooth with microsetae, usually with large tubercles dorsal to antennal sockets. Frons flat and glabrous. Mandible quadrate laterally. Gnathochilarium quadrate.
Antennae short and very clavate. Interantennal distance small.
Metazonites with regular pattern of small tubercles. Body massive and triangular in cross section. Metaterga (usually 5 through 19) with conspicuous rç>ri^t, median projections, and laterally trilobed, large medial lobe with ozopore. Anterior terga directed cephalad. Tergites 1-4 with three irregular rows of 6-8 usually very large tubercles, vhich may be setaceous. Second tergite flabellately enlarged, covering head laterally. Collum oval and small, narrower than head. Paranota poorly developed or extensive and directed ventrad. Peritremata non-existent. Poriferous arrangement normal.
Ozopores moderately large, usually located medially and away from lateral edge on anterior and medial segments, becoming more posteriolateral on caudal segments. Dorsal setae absent. Milliped able to involute. Sterna narrow and unmodified. Vasa deferentia open flush on surface of second coxae. Legs slender and long without modifications in male. Leg length to body length 10.9%. Length relationships of podomeres usually 4<7<1<5<2<3<6. Claws short.
Sphaerotrichomes absent. Hypoproct small, transverse, smooth, and not produced medially. Paraprocts normal with 2 pairs of marginal setae set fairly close to dorsal end. Ipiproct tuberculate, non- hirsute, and may be eu^janded caudad in males. 18
Gonopod aperture large, slightly transversely oval, lateral edges elevated. Gonopod (Fig. 13) sinple, moderately thin, arcuate, capable of moving parallel to boc^, and attached to aperture by merrbrane. Gonocoxae large, in contact medially, and subtriangular with prominent ventral depression to accommodate telopodite reflexion. Depression usually with large dorsolateral field of macrosetae. Prefemur small, thin, and hirsute with a prominent spiniform process covering site of cannular attachment. Femur long with caudad process and posterior row of macrosetae. Cingulum distal on femur, may be very large, forming a hirsute shelf. Seminal groove running distally on medial surface to cingulum, then crossing laterad to small setal pad or setose knob (=endomerite), or may continue to distal tip. Endomerite moderate to non-existent and flush with femur. Seminal vesicle absent. Tibiotarsus regular with large esqjanded or spiniform processes and attached ventrally to gonocoxae.
cyphopod elongate with large operculum with few lateral setae and few marginal macrosetae.
CONTENT: Nine genera, Doratodesmus Cook, 1896, Eutrichodesmus
Silvestri, 1910, Pauroplus Chamberlin, 1945, Eucondvlodesmus Miyosi,
1956, Dimorphodesmus Murakami, 1966, Cerastelachvs Hoffman, 1977,
Ascetophacus Hoffman, 1977, Scolopopvce Hoffman, 1978, and
Seminarchus Hoffman, 1978, and a total of thirteen species. This family can probably be divided into at least 2 subfamilies. 19
The type species in this family was Doratonotus armatus Pocock
(1894). The genus Doratodesmus was subsequently proposed by Cook
(1896) to replace Doratonotus vÆiich was preoccupied in fishes.
Cook proposed the family but without diagnosis. Because of its ability to involute, Verhoeff (1941) placed this family along with the Families Sphaeriodesmidae, cyrtodesmidae, and Oniscodesmidae into a new Suborder Sphaerosomita. This was immediately rejected by the serious workers in the field.
DISTRIBUTION: Southeast Asia (Map 1). Presently known from
Malaysia, Sumatra, Java, Japan, and New Guinea. Several genera are confined to caves, and cilthou^ troglodytic modifications have not been reported, this may be due to insufficient collecting.
FAMILY POLYDESMIDAE Leach, 1815
(Map 1)
Polydesmides leach, 1815. Trans. Linn. Soc. london 11:391. EiçKDlydesminae Attems, 1898. Denkschr. Akad. Wiss., Wien (roath.- naturwiss. Classe) 67:287. Scytonotidae (in part) Cook, 1911. Proc. U. S. Nat. Mus. 40:147. Mastigonodesmidae (in part) Attems, 1914. Arch. Naturg. abt. A 80(4):273.
DIAGNOSIS: Small to large (4-36mm). Segments numbering 19 or
20. I/W ratio 8-22. Tergal width to height ratio approximately 1.1-
1.3. Body width to height ratio 1.0-1.1. Usually brown, rarely red, white, or black. Head smooth, either covered with microsetae or with 20 few moderate filifonn setae. Frons smooth, flat or bulging. Mandible angled laterally. Gnathochilarium either quadrate or elongate.
Antennae usually long and slender but may be short and clavate.
Interantennal distance large. Ifetazonite with regular pattern of small pits. Terga either with small tubercles or transverse rows of flattened polygonal bosses, these sometimes with setae. Anterior terga siitple, directed either laterad or cephalad. Collum large, usually wider than head although smaller in some taxa. Paranota variable, may be ventrally, or laterally directed. Peritremata either thin or thickened. Paranotal edge usually serrate. Poriferous arrangement for 19-segmented millipeds 5, 7, 9, 10, 12, 13, 15, 16,
17, and 18, for 20-segmented millipeds 5, 7, 9, 10, 12, 13, 15, 16,
17, 18, and 19. Ozopores small, opening dorsally or dorsolaterally and either flush with peritremata or elevated. Sterna wide and unmodified. Vasa deferentia large, quadrate, and well separated, legs usually slender and long. leg length to boc^ length 14-23%. length relationships of podomeres usually 1=7<4<5<2=3<6. Male legs sometimes more robust than females, usually unmodified, and with ventral sphaerotrichomes on various podomeres. Female legs unmodified and without sphaerotrichomes. Hypoproct small and oval.
Paraprocts with 2 pairs of setae with margin thickened. Epiproct truncated and projecting sli^tly.
Gonopod aperture large, subcordate, lateral edge slightly elevated with transverse internal shelf on posterior margin. Gonopod usually single, hinged, capable of moving only parallel with body.
Gonocoxae large, subtriangular, with prominent ventral depression to 21
accxmmodate telopodite reflexion. Seminal groove originates mesally,
runs dorsally, then laterally into an internal distal vesicle, usually opening onto a setaceous pad (endomerite). Prefemur moderately hirsute. Femur long with caudally directed distal processes. Tibiotarsus elongate with caudad projections.
Syncoxostemum large, smooth, curving posteriolaterally, and sometimes lobed ventrally. Sternum III may be modified, cyphqpod usually oval but may be elongate, valves large, with long, stout, marginal macrosetae. Operculum small. Receptacle absent.
GOIfrEbTT: Tifenty-four genera including approximately 374 species and subspecies. The Palearctic Region has 16 genera and approximately 345 species and subspecies; the Nearctic Region has 9 genera and 29 species. The genera are arranged into 4 subfamilies.
Mastigonodesminae, Scytonotinae, Ipanerchodinae, and Polydesminae.
DISTRIBUTION: Holarctic (Map 1), including the extreme northwestern tip of Africa. Introduced species have been established in the southern Hemisphere. Species are concentrated in the
Appalachian Mountains, Alps-Balkan Region, and Japan.
Subfamily Mastigonodesminae Attems, 1914
(Figs. 2, 14; Map 1)
Mastigonodesmidae Attems, 1914. Arch. Naturg. Abt. A 80(4) :273. 22
DIAGNOSIS: Small (less than 10mm). Body with 19 or 20 segments.
I/W ratio 10.0-11.5%. IW between 6.5 and 28.3 ram.^. Tergal width to height ratio approximately 1.33. Boc^ width to height ratio approximately 1.00. Brown or depigmented. Head subglobose, covered with microsetae. Frons smooth and bulging. Mandible rounded laterally. labrum projecting very slightly ventrad. Gnathochilarium quadrate. Antennae short and very clavate, with small dorsal tubercle on 7th antennomere. length relationships of antennomeres
1<7=4<5=2<3<6. Antennomeric shapes: 1 quadrate, 2-3 cylindrical, 4-6 variously clavate, 7 subconical. Interantennal distance small.
Ifetazonites with small pits. Anterior terga laterad. Terga (Fig. 2) arched, with 3 rows of seta-bearing tubercles, the setae long and filiform. Anterior terga directed laterad. Collum small, narrower than head. Paranota poorly developed, dentate laterally, and ventrally with posterior margin wider than anterior margin.
Peritremata absent. Poriferous arrangement normal. Ozopores small, opening posteriodorsally, not elevated. Sterna unmodified. Vasa deferentia small. Legs short (14-18% of body length). Length relationship of podomeres 1<7=4<5=2=3<6. Legs unmodified, although more robust in male than female. Claws thin, long, and tapered.
Trochanter and prefemur with sphaerotrichomes, tarsus with long filiform setae. Hypoproct oval. Paraprocts typical, hirsute, and with thin margins. Epiproct sli^tly truncate with few long setae.
Aperture transversely oval with a sternal remnant. Gonopods
(Fig. 14) large, massive and arcuate, but not extending over 6th 23
segment. Gonocoxae large with small distal modifications, extending
laterad over edge of aperture. Prefemur small, hirsute, and may have
small lateral processes. Femur non-existent. Endomerite long, thin,
ending subterminally on the tibiotarsus, and convoluted
(=pseudoflagella of Strasser, 1974) with small terminal vesicle.
Tibiotarsus sittple, massive, long with bend, or curved, attached laterally to the gonocoxae, and may have several terminal caudal processes. Syncoxostemum not narrowed, thickened. Sternum III unmodified. Cyphopod oval without anterior plate.
CONTENT: Two genera, Masticfonodesmus Silvestri, 1898
( =Eumastigonodesmus Brolemann, 1916; =Schedoleiodesmus Silvestri,
1898), with 11 species and 2 subspecies, and Heterocookia Silvestri,
1898 f=Cookia. Attems, 1898, =Haplocookia. Brolemann, 1916) with 4 species are known.
The placement of the mastigonodesmines in the family
Polydesmidae was originally by Hoffman (1979). Previously they were usually listed as a separate family. Hoffman listed the genera individually and stated that the genus lyfastioonodesmus probably deserved subfamilial classification.
DISTRIBUTION: Palearctic (Map 1). The subfamily
Mastigonodesminae is distributed around the western rim of the
Mediterranean Sea with Mastioonodesmus in Italy, Sardinia, Sicily, and the Pyrenees, and Heterocookia in Tunisia and Algeria. Some species are trogloc^rtic with some modification for cave habitation. 24
The two remaining subfamilies of Polydesmidae include Nearctic
r^resentatives and are treated here in more detail. They and their
included genera can distinguished as follows:
Key to North American Genera of Polydesmidae
la. Dorsum without tubercles; paranota horizontal; ozopores not elevated; male legs unmodified; seminal groove looped with large terminal vesicle; tibiotarsus with caudally directed processes...... Subfamily.Polydesminae...... 2
lb. Dorsum with tubercles; paranota ventrally directed; ozopores usually elevated; male legs variously modified; seminal groove strai^t without terminal vesicle; tibiotarsus sinple...... Subfamily Scytonotinae...... 3
2a. Posterior row of quadrate bosses on tergites enlarged; body with 19 or 20 segments; endomerite flush with femur and sometimes with processes...... Polvdesmus
2b. Posterior row of quadrate bosses not enlarged, body with only 20 segments; endomerite projecting from femur and without processes...... Pseudopolvdesmus
3a. Gonopod hiiÿily dendritic with large lateral process...... Idahodesmus
3b. Gonopod not highly branched, siirple or with small regular processes...... 4
4a Gonopod branched; dorsal setae hooked; leg sphaerotrichomes large; femoral processes absent...... Scvtonotus
4b. Gonopod not branched; dorsal setae not hooked; leg sphaerotrichomes small; femoral processes present...... 5
5a. Endomerite without processes; anterior denticles on gonopods present; legs unmodified; tibiotarsus branched...... 6 5b. Endomerite with processes; anterior denticles absent; legs variously modified; tibiotarsus unbranched...... 7 25
6a. Antennae long (18-25% of total boc^ length) ; gonopod with basally directed lateral spine...... Speodesmus
6b. Antennae short (15-18%) ; gonopod without basally directed lateral spine; tibiotarsus with numerous terminal processes...... Coronodesmus
7a. Endomerite flattened, with numerous setae; without femoral setae...... Utadesmus
7b. Endomerite elongate, without numerous setae; femoral setae present...... Bidentoaon 26
Subfamily Scytonotinae NEW STAIUS
(mp 2)
Scvtonotus Koch, 1847, ^ Kirit. Rev. Insect. Deutschlands 3:57.
DIAGNOSIS: Small to medium (4-20ram). Body with 19 segments.
W/L ratio 8-18%. IW between 8.6 and 58.5mm^. Tergal width to height ratio approximately 1.0-1.1. Body width to height ratio approximately 1.0. Usually brown but may be depigmented. Head usually smooth and covered with numerous microsetae. Mandible usually rounded laterally. Gnathochilarium rectangular.
Interantennal distance large. Antennae usually long and may be either filiform or clavate. Length relationship of antennomeres
7<1=4<2<5<3=6. Metazonite with small pits. Terga well arched, never flattened, with seta-bearing tubercles instead of quadrate bosses.
Dorsal and lateral setae usually clavate. Anterior terga directed cephalad. Collum transversely oval ard small, narrower than head.
Paranota poorly developed, directed ventrad, either rectangular or trapezoidal, and dentate laterally. Peritremata usually thick.
Poriferous arrargement normal. Ozopores large, opening dorsolaterally or dorsoposteriorly, and may be elevated. Sterna unmodified although wider in female. Vasa deferentia small. Anterior pair of legs slightly separated from posterior pair. legs short and variously modified. Cla\fl®, short, thickened at base, and constricted distally. Various podomeres with sphaerotrichomes. Hypoproct small and non-hirsute. Paraprocts with at least 4-6 setae and thin margin. 27
%)iproct rugose, sometimes deflexed, setae short and usually with
numerous setae.
Gonopods usually not arcuate, usually directed cephalad
without caudad process, aperture oval. Gonocoxae small sli(ÿitly
triangular to rectangular. Prefemur typical and usually non-hirsute.
Femur usually short. Endomerite small. Seminal groove not looped nor
terminating in small vesicle. Endomerite small. Tibiotarsus long,
may be branched, and attached to the gonocoxae laterally.
Cyphopod oval. Syncoxostemum thin, small, and spined
laterally, with medial hooks.
CONTENT: There are presently 7 genera, Speodesmus. Scvtonotus.
Utadesmus. Bidentoaon. Coronodesmus. Speorthus. and Idahodesmus. and
26 species.
DISTRIBUTION : Nearctic (Map 2). Most genera are from western
North America, the exception is Scvtonotus. vhich includes several eastern North American species. Except for Scvtonotus. most taxa have limited distribution in pockets of stable and protected environments. This subfamily probably represents primitive taxa with relict populations. Several genera also appear to be troglodytes with various degrees of modification. The presence of troglodytic species in regions unsuitable for epigian populations supports the hypothesis of primitiveness. 28
Speodesmus Loomis, 1939
(Figs. 4, 15-18, 52; Map 2)
Speodesmus Loomis, 1939. Bull, ly&is. Coitp. Zool. 86:187.— Hoffman, 1979. Classification of Diplopoda, p. 173. Speobius Chamberlin, 1952. Ent. News 63:12; lapsus. Specodesmus Chamberlin, 1959. Proc. Biol. Soc. Washington 72:74; Lapsus.
GENEROTÏPE: Speodesmus echinourus Loomis, 1939, by monotypy.
DIAGNOSIS; Small to medium, 20 segments, vÆiite or colorless.
Tergites with tubercles bearing filiform setae. Anterior margin narrower than posterior margin. Paranota curved ventrally. Denticles present. Ozopores not on enlarged swellings, opening dorsally. Legs and sterna unmodified. Gonopods straight or curved with posterior femoral ^ines. Tibiotarsus with posteriorly directed spine.
Endomerite flush with tibiotarsus, bearing a thin, curved process.
Seminal groove straight. Females thinner, with thin legs and without sphaerotrichomes. Syncoxostemum membranous, narrow basally. Sternum
III unmodified, cyphopod oval, with few valvular macrosetae, setae absent laterally and medially.
DESCEÎIPITON: Small to medium (8.5-19ram) polydesmids, 20 segments, not including head. W/L ratio 7.4-12.7% and WL 6.4-
40.5rara^.
Color in life viiite or unpigmented, although strongly sclerotized. 29
Head large, smc»th, broader than collum. Genae quadrate.
Cranial suture distinct, extending anteriorly to normal antennal position. Antennae filiform, long, 18-25% of body length, and extending to middle of 4th tergite. Antennomere lengths in
increasing sequence 1<7<4<5<6<2<3. First antennomere rectangular,
2nd and 3rd elongate, 4th rounded ventrally with single large distoventral seta, 5th and 6th clavate, 7th typical with 4 terminal sensillae, antennomeres 5, 6, and 7 with dorsodistal patches of bacilliform setae.
Facial setae numerous, fine, and sitiple.
Mouth components typical except gnathochilarium elongate, I/W approximately 110%.
Metaterga flat, with 4-5 posteriorly curving rows of seta- bearing tubercles, sulcus not distinct, setae not filiform, but peglike (Fig. 52). Collum broadly oval, smaller than head, posterior margin straight, with three rows of setae, lateral denticles absent.
Paranota (Fig. 4) small, curved, anterior margin of tergite narrow, becoming wider posteriorly. T/W ratio approximately 1.4 for paranota and 1.0 for tergites (at midbody). Lateral edges slightly curved, with four or five small denticles, 4 occurring on poriferous segments. Denticles with peglike setae. Ozopores moderate, not elevated, located on posterior margin of paranotum, opening dorsal.
Ozopore formula typical.
Epiproct conical with 4-13 pairs of setae. Hypoproct triangular but truncated, with 7-16 pairs of setae. Paraprocts smooth, margin thin, setae absent. 30
S t e m a flat, unmodified. Legs long, slender, 24-30% of total body length. Coxa small, subquadrate; trochanter moderate, slightly barrel-shaped; prefemur sli^tly curved and clavate; femur and tibia short and subquadrate; tarsus sli^itly curved and elongate; claw short, sinple, and curved. Trochanter, prefemur, and femur usually with large sphaerotrichomes.
Gonopods, in situ, moderately eiqxDsed, curving toward body, parallel, orientated along axis of boc^, capable of independent movement, and extending slightly past anterior margin of 7th sternum.
Gonopod aperture subelliptical with I/W ratio 60-70%. Gonopods large (Figs. 16-18). Gonocoxae large, sli^tly smaller than telopodite, subglobular, flattened mesially, with small ventral cavity. Prefemur small, moderately setaceous. Femur moderate, straight or sli^tly curved, with single anterior seta, and posteriolateral patch of small teeth. Tibiotarsus with large, posteriorly directed spine. Endomerite flush with tibiotarsus, with thin elongate curved process. Cannula connected to telopodite under small ledge. Seminal groove strai^t, not terminating in a vesicle.
Female similar to male except much more slender, legs much shorter and slenderer. Syncoxostemum membranous, narrowed basally and divergent anteriomedially. Sternum III unmodified.
cyphopods oval, with few valvular macrosetae, and setae absent laterally and medially. 31
CONTENT: There are 7 species of Speodesmus of vAiich only 3 are described, Speodesmus echinourus Loomis, 1939, S. bicomourus
Causey, 1959, and S. aouiliensis Shear, 1984. The remaining four species are tentatively based on the unpublished morphometric study by Elliott (1976).
DISTRIBUTION: The genus is centered in the Karst region of the
Balcones Fault Zone of westcentral Texas (Map 2). Elevation of the region is between 450 and 900 meters. The drainage basin includes the Nueces, San Antonio, Guadalupe, and Colorado Rivers. This is primarily a troglodytic genus.
NOTES: Elliott (1976) did an unpublished morphometric study of specimens from central Texas. Althou^ extensive, he didn’t examine cyphopods, and he did not give parameters of any of the characters he used in his principal component analyses.
Elliott stated that bicomourus had 3 sister species, all undescribed. These were based on differences of the following characters: body size, segment 3 denticle number, hypoproct setal number, claw length, setal (bacilliform?) number on sixth antennomere of male and female, height of segment 3, and the following gonopod characters; medial angle of lateral process, tooth number on accessory branch, lateral process length, angle alpha, lateral process width, and number of lateral denticles.
He also determined that echinourus had one sister species, also undescribed. This was based on differences of the following 32 characters; body size, setal number on sixth antennomere, hypoproct, denticle number on third segment of males and females, second segment width of females, and sphaerotrichome size on male legs.
Gonopod characters were angle alpha, lateral process width, lateral process length, medial angle to lateral process, accessory branch tooth number, number of lateral denticles.
Shear (1974b) described a Mexican species, pecki. but in a later paper (Shear, 1975) moved it to the genus Sumidero in the
Family Fuhrmannodesmidae.
Scvtonotus Koch, 1847
(Figs. 5, 20, 21-23, 44, 47, 49, 54, 55
58, 60, 64, 68; Map 2)
Scvtonotus Koch, 1847, In Krit. Rev. Insect. Deutschlands 3:57.— Cook and Cook, 1894, Ann. New York Acad. Sci. 8:233.— Hoffman, 1947, Proc. Biol. Soc. Washington 60:139. Stenonia Sager, 1856, Rroc. Acad. Nat. Sci. Philadelphia. 8:109 (generotype: S. hispida Sager, 1856, by monotypy). lasiolathus Loomis. 1943, J. Washington Acad. Sci. 33:318 (generotype: L. vircfinicus Loomis, 1943, by monotypy).
ŒNEROTYPE: Scvtonotus scabricollis Koch, 1847 r= Polvdesmus qranulatus Say, 1821] by subsequent designation of Bollman,
1893:151.Synonymy of scabricollis Koch with cfranulatus Say by
Bollman, 1893:146. 33
DIAGNOSIS: Small to medium, 19 segments, brown or light cream
colored. Tergites with seta-bearing tubercles. Setae of adults
hooked, and filiform; juvenile with clavate setae. Paranota small and
curved ventrolaterally. Ozopores usually enlarged, opening
dorsolaterally. Male tibia may be modified. Sterna unmodified
although wider in males. Gonopods arcuate with flexible tibial arm
with processes. Endomerite small, seminal groove sinuous, with small
terminal bulb. Female paranota on tergites 5-11 may be reduced.
Syncoxostemum base membranous, separated distally, with cephalad
lateral spines. Sternum III unmodified. Cyphopod oval, without
anterior plate, ventral lobes small and numerous, receptacle small
with several large cephalad directed setae.
DESCRIPTION: Small to medium (7 to 2Oram) polydesmids, with 19
segments not including head. W/L 10.7-14.6% and WL 18.0-58.5mm^.
Color in life, either l i ^ t brown or cream colored. Venter
lighter.
Head smooth. Genae quadrate, projecting laterally. Cranial
suture distinct. Antennae (Fig. 58) well separated, 20-25% of total
head width, sli^tly clavate, base thicker than in Pseudopolvdesmus,
approximately 18-24% of total body length, extending to anterior
margin of 4th segment. Antennameric length in increasing sequence
7<1<4<2<5<6<3. Antennomeres 1-4 rectangular, 5 and 6 slightly
clavate, and 7 typical with four terminal sensillae present.
Antennomeres 5, 6, and small swelling on 7 with small patches of
bacilliform setae located dorsodistally. 34
Mouthparts typical except gnathochilarium elongate, I/W
approximately 110%.
Face with numerous moderate filiform setae.
Metaterga (Fig. 5) convex, not quadrate, with 4 or 5
transverse series of seta-bearing tubercles. Each tubercle with long
hooked setae. In juveniles setae irregularly clavate. Usually 4
pairs of anterior tubercles located on anterior half of tergum
between stricture and transverse sulcus, immediately behind sulcus
intermediate set with 5 pairs, and along posterior margin 6 pairs
present. Collum small, oval, or with anteriolateral edge slightly
produced cephalad, with numerous of setiferous tubercles in 3 to 5
rows. Anterior edge of collum flaired. Metazonite with medium sized
pits.
Paranota curved ventrally (Fig. 49), W/T ratio l. 46 and
tergite ratio 1.0, without serrations but 7-8 lateral projections
present, each usually with hooked setae. Paranota quadrate
laterally. Ozopores usually elevated with enlarged peritremata (Fig.
47), opening dorsolaterally on posterior half of paranota. Ozopore
formula 5, 7, 9, 10, 12, 13, 15, 16, and 17. Paranota of segments 2-
4 directed cephalad (Fig. 44), of other segments progressively more
caudad; posterior margins with setae and convexities usually less
developed than those on anterior segments.
Sterna smooth, penes small. Anterior and posterior sterna
(Fig. 60) of male in several species wide but otherwise unmodified.
Male legs enlarged (Fig. 54). Leg length to body 17-22%. Coxa
clavate; trochanter bowed on both ends; prefemur elongate; femur 35
short; tibia rectangular or in some species with 1 or 2 irregular
projections on legs 13-20; tarsus curved and long; claw moderate,
constricted basally (Fig. 56). Sphaerotrichomes on tarsus, small and
numerous with small patches on remaining segments except coxa.
%)iproct truncate and rugose, with at least 8 pairs of setae.
Hypoproct small, rounded, with a pair of posteriolateral setae on
small swellings. Paraprocts smooth, with two pairs of lateral setae,
lips thin.
Gonopods, in situ, moderately ej^xDsed, usually not inserted,
parallel, oriented along medial axis of b o ( ^ , and capable of
independent movement, not extending over anterior margin of 7th
segment (Figs. 21-23).
Gonopod aperture oval, with I/W ratio between 60 and 76%,
posterior shelf large, anterior shelf small, raised laterally.
Gonocoxae moderate, smaller than telopodite, with large ventral
concavity for retraction of gonopods. Ventrolateral edge with two
setae. Prefemur without processes, very hirsute, and long medially.
Femur short and either thick (eastern species) or thin (western
species), with 1-2 mesial macrosetae, and 1-4 large cephalad
directed setae. Telopodite arcuate with two flexible pieces. Based
on Hoffman (1962), Scvtonotus has up to 2 lateral (x & y) and/or
lateral (s & t) and endomerite (a) processes. Cannula connected to
prefemur under ledge. Endomerite small and may have a small distal process. Seminal groove sinuous, extending anterior to level of
endomerite. Tibiotarsus simple, sometimes with distally directed
tooth-like setae. 36
Female structurally dissimilar depending on species. Paranota
5-11 variously reduced (Fig. 20), the segments appearing almost
cylindrical. Antennae shorter and more clavate. Sterna wider than
male. legs short and unmodified but slightly weaker than males,
sphaerotrichomes not as distinct. Syncoxostemum membranous, coxae
narrowed, separated from sternum, with ventrolateral spine (Fig.
64). Apodemes long. Sternum III rounded but unmodified.
cyphopods small, axis longitudinal, moderate, with 25-30 small
macrosetae (Fig. 68). Flattened ventromedially, receptacle present,
small not elongate, with 2-5 very long cephalad directed setae,
lateral and medial surface with few setae.
CONTENT: The eleven described species include seven in the
northwest and four in the east. The western species are: amandus
Chamberlin, 1910, bercorothi Chamberlin, 1911, columbianus
Chamberlin, 1920a, insulanus Attems, 1931, orthodox Chamberlin,
1925, piaer Chamberlin, 1910, and simplex Chamberlin, 1941. The
eastern species are: in Grorç) Granulatus: qranulatus (Say, 1821) and
australis Hoffman, 1962, and in Groiç) Virginicus: virginicus
(Loomis, 1943b) and michauxi Hoffman, 1962.
DISTKEBUnON: Nearctic (Map 2). Two separate populations
occur, one in the Pacific Northwest, from southern Alaska south to
Utah and Idaho, and the other in northeastern North America, New
York south to North Carolina, west to Iowa and Missouri. Available data suggest that the disjunction may be real. 37
NOTES: Tÿpe materials of Polvdesmus granulosus (Say, 1821),
and two of its synonyms: S. scabricollis (Kcch, 1847), and S.
laevicollis (Koch, 1847) are lost. Say (1821) described the first
species of Scvtonotus. Koch (1847) described the genus but failed to
designate a l^'pe from the three described species of vhich
granulosus was not one. Bollman (1893) (posthumously) states
incorrectly (page 146) that Koch designated granulosus as the type.
On page 151, however, he states that scabricollis is the type of the
genus.
Cook and Cook's redescription (1894) of S. granulosus was very
extensive and provides a basis for the modem use of the name.
Lasiolathus Loomis (1943) vras based on juvenile specimens of
Scvtonotus virginicus and was synonymized into granulosus by Hoffman
(1947). Hoffman (1962) recognized S. virginicus to be a separate
species. Stenonia hispida was synonymized by Bollman (1893) with
granulatus.
Scvtonotus cavemarum Bollman was made a synonym of S.
granulosus by Chamberlin and Hoffman (1958). The identity of S.
nodulosus Koch is uncertain because the type is presumed lost and
the description is inadequate.
Utadesmus Chamberlin and Hoffman, 1950
(Figs. 6, 24-26, 53; Map 2)
Utadesmus Chamberlin and Hoffman, 1950. Chicago Nat. Hist. Misc. 71:2. 38
GENEROTYPE: Brachvdesmüs henriensis Charaberlin, 1930, by
original designation.
DIAŒOSIS: Small, 19 segments, brown. Tergites tuberculate.
Paranota small, ventrally curving. Ozopores elevated, opening
dorsoventrally. Legs and sterna unmodified. Gonopods sirrple,
gonocoxae slightly expanded laterally, prefemur small, femur long
and thin, tibiotarsus sittple with small lateral tooth. Endomerite
large and flattened. Syncoxostemum with large lateral spines and
small medial pair of hooks. Sternum III ventrally lobate. cyphopod
small, oval, with diminished macrosetae. Receptacle with several
long, anteriorly-directed setae.
DESCKEPITON: Small (5-17mm) polydesmids, 19 segments not
including head. W/L 13-19% and WL 12.4-24.6mm2.
Color in life earth brown, venter and legs lighter.
Head smooth and globose. Genae quadrate, projecting laterally.
Cranial suture short but distinct, terminating dorsal to antennae.
Antennae well separated, set medially on face, antennae short, 12-
16% of body length, extending to midline of tergite 3, and very
clavate. Antennomeric lengths in increasing size 7<1<5<2=4<3<6.
First antennomere small, quadrate, 2-4 elongate, and 5, 6, and 7 markedly clavate, each with patches of bacilliform setae and with 4 terminal sensillae.
Head covered with numerous microsetae.
Mouthparts typical with slightly elongated gnathochilarium. 39
Metaterga (Fig. 6) arched, not horizontal, with three
transverse rows of tubercles, each with a single peg-like seta.
Anterior row with 10 tubercles and separated from medial and posterior rows vSiich have 10 and 8 tubercles respectively. Collum
oblong, narrower than head, with 3 rows of seta-bearing tubercles,
collum not flaired.
Paranota curving ventrally, not expanded laterally. W/T ratio
1.5 for paranota and 1.1 for terga. Anterior paranota directed cephalad directed. lateral edge quadrate. Denticles with peg-like setae, somevôiat variable. Non-poriferous segments with 6 lateral denticles \diich, on the posterior segments, are reduced and directed posteriorly. One to several small non-setaceous teeth may be present anterior to anteriomost denticle. Poriferous segments with 3 denticles anterior and 2 denticles posterior to ozopores, althouÿi several very small denticles may be associated with ozopores.
Denticles reduced on posterior segments. Ozopores large, elevated, located slightly posterior to midline of tergite, opening dorsally, arrangement typical. Posterior paranota large and posteriorly directed.
Epiproct smooth and truncated with approximately 8 pairs of setae. Hypoproct round with single pair of setae. Paraprocts with thin lips and two lateral pairs of setae.
Sterna flat, unmodified, with few setae. legs (Fig. 53) short,
10-14% of total body length. Coxae quadrate, trochanter rectangular, femur quadrate, tibia quadrate, tarsus elongate and thin, claw 40 constricted distally. Sphaerotrichomes large on tarsus, tibia, and femur.
Gonopods (Figs. 24-26), in situ, lying partly out of aperture, gonocoxae ejqxDsed, telopodite directed anteriorly, along midline and extending to midway on the sixth segment. Posterior pair of legs on sixth segment only sli^tly displaced laterally.
Gonopod aperture wide, I/W ratio approximately 60%, kidney shaped with large medially directed, posterior shelf. Gonocoxae large, globose, and anterior margin quadrate. Prefemur small and hirsute. Femur elongate, very sli^tly curved, uribranched, without setae, and distally with large flattened endomerite. Seminal groove strai^t. Tibiotarsus long, spine-like, with small proximolateral spine.
Female similar to male, except legs thinner and without sphaerotrichomes. Syncoxostemum with anteriorly directed lateral spines, a small pair of medial hooks, and anteriomedially, a small lobe between coxae. Coxae slightly ejpanded medially. Sternum III ventrally Idbate.
cyphopod oval, small, with few diminished macrosetae.
Receptacle with several long anteriad directed setae.
CONTENT: Two species, henriensis Chamberlin, 1930 and hoffi
Chamberlin and Hoffman, 1950. Undoubtably more species will turn up with adequate collecting. 41
DISTRIBUTION : Central New Mexico and northcentral Utah, on the
Colorado Plateau (Map 2). Probably extends into southern Viycming in
the San Juan, Sangre de Cristo, Medicine Bow, and Sawatch mountain
ranges.
NOTES; The holotype of U. henriensis has a gonopod missing.
The type of U. hoffi also has only one gonopod and lacks the
anterior half of the body. The separation between these two species
is sli^t with differences occurring in lateral spine shape and
femur curvature. Packard (1877) described Polvdesmus cavicola from a
northern Utah cave. Because the Packard material has never been
located and his description lacks a gonopod description or drawing,
its identification will remain in doubt. Chamberlin and Hoffman
(1958) placed cavicola in the genus Brachvdesmus ; however, I ej^ct
it to be a Utadesmus.
Bidentoaon Buckett and Gardner, 1968
(Figs. 7, 27-29, 51; Map 2)
Brachvdesmus. Chamberlin, 1918a, Pomona Coll. J. Ent. and Zool. 10(1):9, not Brachvdesmus. Heller, 1858. Bidentoaon Budœtt and Gardner, 1968, Pan-Pacific Ent. 4:198.
GENEROTYPE: Bidentoaon helferorum Buckett and Gardner, 1968, by monotypy. 42
DIAGNOSIS: Small, 19 segments, brown. Tergites tuberculate.
Paranota small, curving ventrally. Ozopores elevated, opening dorsoventrally. Male third pair of legs with tibia swollen and prefemur slightly swollen. Remaining legs and sterna unmodified.
Gonopod directed cephalad, gonocoxae large and variously lobed.
Prefemur small and hirsute. Femur strai^t with elongate row of setae. Tibiotarsus bifurcate. Endomerite long, with straight seminal groove, few setae. Syncoxostemum menbranous, with lateral spines.
Sternum III unmodified, cyphopod small and oval.
DESCRIPTION; Small to medium (<12mm) polydesmids, with 19 segments not including head. W/L 14.1-17.9% and WL 8.6-21.8mm^.
Color in life, brown, venter and legs lighter.
Head smooth and broad. Genae quadrate, projecting laterally.
Mandible laterally rounded. Cranial suture short but distinct, terminating dorsal to antennae. Antennae well separated, set medially on face, antennae short and clavate, 12-18% of body length, extending to anterior edge of 4th tergite. Antennomeric lengths in increasing sequence 7<1<4<2<5<6<3. First antennomere quadrate, 2-4 rectangular, 5 and 6 slightly clavate, and 7 typical with 4 terminal sensillae, antennomeres 5, 6, and small knob on 7 with small patches of bacilliform setae.
Face covered with numerous microsetae.
Mouthparts typical except for elongation of gnathochilarium.
Metazonite (Fig. 7) with small pits. Metaterga convex, with 3 transverse rows of small tubercles, each with a single peg-like seta 43
(Fig. 51). Anterior row with 10 tubercles, separated from medial and
posterior rows v M c h have 10 and 8 tubercles, respectively. Collum
oblong, rounded laterally, smaller than head and remaining tergites,
setae-bearing tubercles present, margins not flaired.
Paranota curving ventrally. W/T ratio approximately 1.3 for
paranota and 1.0 for tergites. lateral edges quadrate with 3 or 4
denticles, 4th denticle directed posteriorly, and occurring only on
poriferous segments. Anterior paranota anteriorly directed. Ozopores
small, elevated, opening dorsolaterally, located on posteriolateral
comer, arrangement typical. Posterior paranota progressively more
caudally directed.
ïpiproct smooth, sli^tly deflexed, with numerous setae.
Hypoproct small, with pair of seta-bearing swellings. Paraprocts
smooth, with two pairs of lateral setae, lips small.
Sterna cruciately inpressed, otherwise unmodified, with few
setae. Posterior 6th sternum with deep depression. Legs short, 15.4%
of total body length. Coxa short and globose; trochanter elongate
and bowed; prefemur short and rectangular; femur short; tibia moderately short and rectangular; tarsus long and very slightly
curved; claw moderate with tip sharply tapered. Male with tibia of third leg twice as large as remaining legs, femur also swollen.
Gonopods, in situ, partly hidden, gonocoxae moderately exposed, telcpodite directed anteriorly and extending to sixth
sternum, capable of independent movement. Posterior pair of legs of segment 6 displaced laterally. 44
Gonopod aperture oval with I/W ratio 67.4%. Gonopods elongate and small (Figs. 27-29). Gonocoxae moderately large, round, with distal edge variously lobed, ventrally large, dorsally small.
Prefemur small and hirsute. Femur straight, long, with irregular row of 10-20 elongate, caudally directed setae. Cannula attached to prefemur under small ledge. Seminal groove curved but not looped.
Tibiotarsus with two caudal branches; a thin, mesal fork, and a large, lateral fork with seminal groove terminating close to distal tip, with fine setae (vestigial endomerite?).
Female similar to male except legs thinner, unmodified, and without sphaerotrichomes. Syncoxostemum coxae small, separated from sternum; sterna overlap medially and are slightly lobate, with laterally distally directed spines. Sternum III with small, central, anteriorly directed swelling.
cyphopod oval, small, without anterior plate, but with several long lateral setae. Receptacle small with long setae.
CONTENT: Two species, Bidentoqon helferorum Buckett and
Gardner, 1968 and B. califomicus (Chamberlin, 1918a) are known.
More species are ej^>ected.
DISTRIBUTION: The genus is limited to the coastal regions of northern and central California.
NOTES: The original description by Buckett and Gardner was well done, although they placed this genus in the family 45
Vanhoeffeniidae (=Fuhrniarinodesmidae) viiich has no North American
representatives. This arrangement was continued by Shear (1972) # 1 0
redescribed and redrew Brachvdesmus califomicus. Its placement in
the family Polydesmidae was corrected by Hoffinan (1979).
Coronodesmus NEW GENUS
(Figs. 8, 30-32; Map 2)
Brachvdesmus (Brachvdesmus), Causey, 1954, Pan-Pacific Ent. 30(3):224 (not Keller, 1858).— Loomis, 1960, J. Kansas Ent. Soc. 33(2) :57.— Chamberlin and Hoffman, 1958, 212:65, not Brachvdesmus Heller, 1858.
GENEROTYPE: Brachvdesmus vosemitensis Causey, 1954, by present
designation.
DIAGNOSIS: Small, 19 segments, red-brown. Tergites with 3 rows
of tubercles, anterior row with filiform setae cephalad and
posterior rows with setae caudad. Paranota directed ventrally with
anteriorly directe! denticle. Ozopores elevated, opening
dorsolaterally. Male sterna and third pair of legs modified,
hypoproct and paraprocts hirsute. Gonopods straight except
tibiotarsus bent anteriorly, with posterioventrally directed processes and anterior patch of swellings or small teeth. Seminal groove straight, without terminal vesicle. Endomerite small. Female
legs thin, unmodified, without sphaerotrichomes. Syncoxostemum membranous with small lateral spines. Sternum III unmodified, cyphopods oval, with few valvular macrosetae, and a few lateral and medial setae. 46
DESCRIPTION; Small (5.5-12iran) polydesmids, with 19 segments not includirg head. W / L 8-18%, and WL 5.9-15.4ram^.
Color red-brown. Venter lighter.
Head moderately smooth. Genae quadrate, projecting laterally.
Cranial suture faint, not extending to antennae. Antennae well separated, clavate, short, 15-18% of body length, extending to posterior margin of third segment. Antennomeric length in increasing size 1<4<7<2=5<6<3. First antennomere rectangular and short, 2 slightly elongate, 3 elongate and clavate, 4 short and subquadrate,
5 and 6 moderate and slightly clavate, and 7 typical with terminal sensillae; antennomeres 5, 6, and 7 with small patches of bacilliform setae located dorsodistally.
Facial setae short and numerous.
Mouthparts typical, with elongate gnathochilarium, I/W approximately 110%.
Metaterga arched, each with 3 transverse rows of tubercles: anterior row with 8 tubercles each with an anteriorly directed filiform seta, and located on anterior half of tergum between stricture and transverse sulcus; intermediate row immediately behind sulcus with 8 tubercles each with a posteriorly directed filiform seta; posterior margin (row) with 12 tubercles each with a posteriorly directed filiform seta (Fig. 8). Collum large, about equal to head, rounded anteriorly, with 5 uneven rows of tubercles, and posteriolateral angle rounded.
Paranota curved ventrally, W/T ratio 1.4, tergal ratio 1.0, 4 or 5 well defined seta-bearing denticles, 5 occurring on poriferous 47
segments with anteriorly directed denticle. Paranota 2-5 anteriorly
directed. Ozopores moderately large, elevated on posteriolateral
c o m e r , opening dorsolaterally on segments 5, 7, 9, 10, 12, 13, 15,
16, 17, and 18. Posterior paranota large, progressively more
caudally directed.
Epiproct truncate, with approximately 8 pairs of setae.
Hypoproct rounded, with numerous setae. Paraprocts highly setaceous
with 20-30 long filiform setae scattered over surface. Lips small.
Stema flat, unmodified, except 10th segment with anterior
sternum with 2 small triangular lobes, posterior sternum with two
long conical lobes projecting caudally. Legs short, typically shaped
except third pair with tarsus sli^tly swollen and femur very
swollen. Sphaerotrichomes present on tarsus, tibia, and femur.
Gonopods (Figs. 30-32), in situ, moderately exposed, not
inserted, parallel, separated, not particularly movable, and not
extending past anterior margin of 7th sternum.
Gonopod aperture elliptical with I/W ratio 60-72%. Gonocoxa
large, about equal to telopodite, with small ventral concavity.
Anterior edge smoothly lobed. Cannula connected to prefemur under small ledge. Prefemur small and hirsute. Femur straight, thin basally, e>q)anding distally, with lateral, distally directed process. Seminal groove straight, terminating in small endomerite without vesicle. Tibiotarsus sli^tly bent anteriorly, with several large posteriorly directed ^ines. Anterior surface with, a patch of swellings or small siitple teeth. 48
Female similar to male but without sternal and leg
modifications. Sphaerotrichomes absent. Syncoxostemum membranous
and sli^tly produced ventromedially, lateral spines small. Sternum
III unmodified.
cyphopods oval, with few macrosetae along valvular edge. Few
lateral and medial filiform setae present. Cyphopod plate rounded, with few lateral and medial setae.
CONTENT: Two described species, Coronodesmus vosemitensis
(Causey, 1954) and Ç. bituberculatus (Loomis, 1960), and a third undescribed species.
DISTRIBUTION : Coronodesmus has been collected in central
California around San Francisco and in Yosemite National Park.
NOTES: It has been noted by several authors that these species were not in the genus Brachvdesmus. but no one has officially made the nomeclatural change.
Speorthus Chamberlin, 1952
(Figs. 9, 33-35; Map 2)
Speorthus Chamberlin, 1952, Ent. News 63:12.— Loomis, 1960, J. Kansas Ent. Soc. 33:66.— Shear, 1969, Psyche 76:134. Speodesmus. Shear, 1974, Occasional Papers California Acad. Sci. No. 112:6, not Loomis, 1939.
GENEROTYPE: Speorthus tucfanbius Chamberlin, 1952, by monotypy. 49
DIAGNOSIS: Small, 20 segments, pale or colorless. Antennae
short and clavate. Tergites with 3 rows of tubercles. Ozopores
moderately large, elevated, opening dorsolaterally. Male stema and
legs unmodified. Gonopod arcuate, without posteriorly directed spine
or dentate field. Endomerite dendritic, lateral process with
proximally directed teeth. Female legs thin and unmodified.
Syncoxostemum membranous and thin basally. Stemum III unmodified.
Cyphopods oval, small, with a very small smootli cyphopod plate with
few valvular macrosetae.
DESCRIPTION: Small (8.0-9.5mm) polydesmid, with 20 segments
not including head. W/L 11-13.5% and WL 8.2-10.8mm^.
Color in life pale or colorless, although well sclerotized.
Head large. Genae quadrate, projecting laterally. Antennae
short and clavate, 11-16% of body length, extending to midline of
4th tergite. Antennomere length in increasing sequence
1<7<2<3=4<5<6. First antennomere short and subquadrate, 2-4 ^ort, rectangular with 3 and 4 rounded ventrally, 5 short and clavate, 6 large and clavate, 7 typical, with 4 terminal sensillae.
Facial setae short and numerous.
Mouthparts typical, gnathochilarium elongate.
Metatergite arched, tuberculate, with 4 or 5 rows each with
10-14 tubercles, each tubercle with a peg-shaped seta (Fig. 9).
Collum broadly oval, slightly narrower than head, with 3 irregular rows of setae and two lateral denticles. 50
Paranota small, curved ventrally, T/W ratio 1.4 and tergal
ratio 1.0. lateral edge with 4 or 5 denticles, each with single
seta. Lateral margin quadrate. Ozopores moderately large, elevated,
opening dorsolaterally on segments 5, 6, 8, 9, 10, 12, 13, 15, and
16. Posterior margin with large lateral serrations.
Epiproct with approximately 8 pairs of setae. Hypoproct small,
non-hirsute, and truncate. Paraprocts with 2 pairs of seta-bearing tubercles, lip slight.
Stema flat, unmodified. Legs long and enlarged, 17-22% of total body length. Coxae large and globular; trochanter thick and barrel-shaped; prefemur thick and curved; femur short and quadrate, tarsus long and curved, and claw long and constricted. Femur and tibia each with single dorsodistal seta. Only tarsus with
sphaerotrichomes.
Gonopods, in situ, es^sed, parallelling body axis, extending past anterior margin of 7th stemum.
Gonopod aperture oval with I/W ratio 68-72%. Gonopods long and thin (Figs. 33-35). Gonocoxae moderate, smaller than telopodite, with small, deep ventral concavity, anterior edge monolobed.
Prefemur moderate and hirsute. Cannula thin, connected to prefemur beneath ledge. Femur not thin, curved with a single anteriorly directed seta. Seminal groove straight without terminal vesicle.
Tibiotarsus simple and elongate. Ehdcmerite large and dendritic, with long lateral process.
Female similar to male; however, more slender, head appearing proportionally larger. Legs shorter, more slender but without 51 sphaerotrichomes. Syncoxostemum membranous and thin basally.
Stemum III unmodified.
cyphopod oval, small, with few valvular macrosetae, and with a very small smooth cyphopod plate.
CONIENT: One species, Speorthus tucfanbius Chamberlin.
DISTRIBUTION : Speorthus tucfanbius occurs in caves in central and southern New Mexico and the western tip of Texas in the Capitan-
Sacramento-Guadalupe mountain systems. The caves are in karst and gypsum plains, elevation 1,800 to 2,450 meters.
NOTES: Loomis (1960) suggested that Speorthus and Speodesmus may be related. Shear (1974a) placed Speorthus with Speodesmus and placed Speodesmus in the family Fuhrmannodesmidae, based on gonopod characteristics. Buckett and Gardner (1968) mentioned that a number of similarities are shared with Bidentoaon.
Idahodesmus NEW GENUS
GENEROTYPE: Idahodesmus dentatus new species.
DIAGNOSIS & DESCRIPTION: See below under I. dentatus.
DISTRIBUTION: Northem Idaho. 52
Idahodesmus dentatus NEW SPECIES
(Figs. 10, 36-39; Map 2)
DIAŒJOSIS: Small, 20 segments, vàiite. Tergites slightly tuberculate. Paranota small and curving ventrally. Ozopores elevated, opening dorsolaterally. legs and stema unmodified.
Gonopod large, elongate, with siirple tibiotarsus, femur with 3 large dendritic lateral processes and medially with band of teeth extending the length of femur, and distally with elongated endomerite with 2 spines. Seminal groove looped. Prefemur small, and gonocoxae large with anterior directed spine. Female with wider stema and thinner legs. Syncoxostemum spined with coxa mgose and hirsute. Stemum with small bifurcated process, cyphopod slightly elongate with few valvular macrosetae, 20-25 large lateral and medial setae, without dorsal plate.
TYPES; Holotype and allotype deposited in USNM; 2 male and 1 female paratypes deposited in North Carolina Museum of Natural
History; all collected from the U. S. Forest Service caitpground, l.dmi. E. of Harvard, Latah Co., Idaho, IX-17-1978 by A. K. Johnson.
DESCRIPTION: Small (5-9mm) polydesmids, with 20 segments not including head. W/L 11.2-13.4% and W L 4.9-8.2rnm^.
Color in life, probably vhite. 53
Head small, globose. MarKÜble quadrate laterally. Cranial
suture indistinguishable. Interantennal distance small. Antennae set
medially on face, long, sli^tly clavate, 18-26% of body length, and
extending to posterior margin of 3rd tergite. length relationship of
antennomeres 7<1=4<2<5<3<6. First antennomere small and quadrate, 2
globose, 3 elongate and slightly clavate, 4 rounded ventrally, 5
short and clavate, 6 long and enlarged, and 7 typical with 4
terminal sensillae. Small patches of bacilliform setae located
dorsolaterally on antennomeres 5, 6, and small knob on 7.
Head covered with numerous microsetae.
Mouttçarts normal except gnathochilarium elongate.
Metaterga arched, not horizontal, with 3 transverse rows of
small tubercles, each with single filiform seta. Anterior row with
10 tubercles, separated from remaining rows #iich have 10 and 8
tubercles respectively (Fig. 10). All dorsal setae directed caudad.
Collum semicircular, with 3 rows of setae, not flaired, and with
small lateral denticles.
Paranoiza curving ventrally, not e)ganded laterally. W/T ratio
approximately 1.3 for paranota and 1.0 for tergites. Anterior paranota 2-4 directed cephalad. lateral edges rounded, with six
denticles on poriferous segments and five on non-poriferous
segments. Small teeth or serrations present anterior to anterior
denticle. Ozopores small, opening dorsolaterally and elevated.
Poriferous arrangement typical. Posterior paranota progressively reduced and directed posteriorly. 54
Hypoproct small with single pair of seta-bearing swellings.
Paraprocts with 4-6 pairs of setae, lips thin. Epiproct rugose,
sli^tly deflexed, with numerous setae.
Stema unmodified, with many short setae. Posterior half of
6th sternum sli^tly di^laced laterally and moderately deep. Legs
unmodified and short, 16.2% of total body length. Coxae, short and
quadrate; trochanter, moderately long and quadrate; prefemur,
elongate and subquadrate; femur, enlarged and rounded dorsally;
tibia, moderately short and quadrate; tarsus, long and slightly
curved; claw, short and constricted. Legs with numerous setae and
sphaerotrichomes.
Gonopod, in situ, lying out of aperture, gonocoxae exposed,
telopodite directed anteriodorsally, and extending to anterior
margin of 6th stemum.
Gonopod aperture oval and wide, with I/W ratio 52.5%. Gonopod
large (Figs. 36-38). Gonocoxae large, globose, with large anteriorly
directed spine. Cannula attached under base of coxal spine. Prefemur
large, hirsute. Femur with medial band of small, posteriorly
directed, dendritic teeth, laterally with three major processes,
proximally with small, siitple dendritic process, the central process
large with medially scimitar-shaped branch with small teeth on outer
edge and two smaller laterally directed branches, one with two
spines and 4 dendritic branches, and distal process spine-like with two smaller spines. Endomerite inserted distally on femur, elongate, 55
with short medial bifurcated process and large terminal process.
Seminal groove strai^t. Tibiotarsus simple^ curving laterally.
Female similar to male, except stemum wider, legs thinner,
and without sphaerotrichomes. Syncoxostemum rou^ and hirsute,
lateral spines present. S t e m u m III (Fig. 39) modified.
cyphopods elongate with few marginal macrosetae, although
with 20-25 large lateral and medial setae, anterior plate absent.
DISTEÎIBÜTION; Ehown only from the vicinity of the type
locality. Idaho: Latah Co., 1.5mi. E. of Harvard, USFS campground,
(NCMH). l^roximate elevation between 450 and 900 meters.
Subfamily Polydesminae Leach, 1815, NEW STATUS
(Table 1)
Eupolydesminae, Attems, 1898. Denkschr. Akad. Wiss. Wien (Math, naturwiss. Classe) 67:416.
DIAGNOSIS: Small to large (7-36ram). Boc^ usually with 20
segments, sometimes with 19. W/L ratio between 8-22%. LW between 10 and 165 ram?. Tergal width to height ratio approximately 1.3. Body width to height ratio approximately 1.0. Usually brown with
occasional red, vhite, and black. Head smooth with few filiform
setae. Frons flat or sli^tly globose. Mandible quadrate laterally.
Gnathochilarium quadrate. Antennae long and filiform, rarely clavate. Length relationship of antennomeres 7<1<2<6=5<4<3.
Antennal shape usually long and filiform. Interantennal distance 56
large. Metazonites with small pits. Terga only sli^tly arched,
flattened, with three rows of smooth quadrate bosses. Dorsal setae,
if present, filiform. Paranota laterally directed. Collum large,
wider than head. Paranota well developed, laterally smooth or
dentate sometimes with setae. Peritremata thick. Poriferous
arrangement normal. Ozqpores small, opening dorsolaterally, not
elevated. Sterna in male may be sli^tly modified. Vasa deferentia
large. legs long, thin, and unmodified and sometimes more robust in males. Length relationship of podomeres 7<1<4<5<2<3<6. Claws short,
thin, and equally tapered. Tarsus, tibia, and femur with
sphaerotrichomes. Hypoproct small, sli^tly oval. Paraprocts typical with thin margins. Epiproct small, with short setae, and may be deflexed.
Gonopods siitple, arcuate, with caudally projecting processes.
Gonopodal aperture transversely oval. Gonocoxae large, triangular, variously lobed anteriorly, with ventral depression. Prefemur large, hirsute, and unmodified. Femur long, curved, with small distal cingulum. Seminal vesicle looped, ending in large terminal bulb.
Endomerite midway up tibiotarsus, large, and may have processes.
Tibiotarsus unbranched, attached ventrally to gonocoxae, and with various caudad projecting processes.
Syncoxostemum unspined, thickened, and massive. Sternum III may be modified, cyphopods may oval or kidney-shaped with moderately large anterior plate. 57
CONTENT : Presently includes 11 genera (Table 1) and 220
species. The status of many Central Asia genera is uncertain. The
subfamilial designation b y Attems (1898) included Polvdesmus.
Brachvdesmus. Archipolvdesmus. and Pseudooolvdesmus.
DISTEÎIBÜTION; Holarctic. Eastern North America
fPseudopolvdesmus), Asia, and the northwestern tip of Africa, most
prevalent in Europe. No species shows any trogloc^rtic modifications.
Polvdesmus latreille, 1802
(Figs. 11, 40-42, 65)
Polvdesmus Latreille, 1802, Histoire naturelle des crustacés et des insectes. 3:44.— Attems, 1940, Das Tierreich 70:3.
GENEEOTÏPE: Julus complanatus Linnaeus, 1761, by monotypy.
DIAGNOSIS: Small to large, 19 or 20 body segments, brown, sometimes dull vhite, red, or black. Antennae usually filiform but may be sli^tly clavate. Tergites quadrate, each with polygonal bosses including a pair of lateral bosses, four medial bosses, and three rows of internal bosses. Paranota large and horizontal.
Ozqpores not elevated, opening dorsally. Male sterna and legs unmodified. Female legs unmodified, thin, without sphaerotrichomes.
Gonopods arcuate with enlarged endomerite usually with at lecist one process, seminal groove looped with large terminal vesicle. 58
Synœxostemum ventramedially modified but not enlarged, base not expanded laterally. Sternum III variously lobed. Cyphopod elongate, short macrosetae only on posterior edge of valves, anterior plate variable, usually small. Lateral and medial setae present.
DESCRIPTION: Small to large (7-36rnm) polydesmids, with 19 or
20 segments not including head. W/L approximately 11-18% and WL 10-
IBOmm^.
Dull colors prevalent, usually brown, althou^ vhite, brown, and black may occur. Venter, legs, and genae usually lifter.
Antennomeres darker distally with terminal antennomere darkest.
Head smooth. Genae sli^tly triangular, projectirg laterally.
Cranial suture distinct, not extending to antennal insertion.
Antennae well separated, set medially on face, short, slightly clavate, reported to be filiform in some species, approximately 12-
18% of total bo(^ length, extending to midpoint of third segment.
Length relationship of antennomeres 7<1<2=3<4<5<6. First antennomere short and rectangular, antennomeres 2-4 short and subquadrate, antennomeres 5 and 6 clavate with 6 elongate, and 7 laical short, with 4 terminal sensillae, antennomeres 5 and 6 and small knob on 7 with small distinct patches of bacilliform setae dorsally.
Facial setae numerous, filiform, and small.
Mouthparts typical, with quadrate gnathochilarium.
Metaterga essentially flat, not tuberculate, each with three well defined transverse internal rows of quadrate convex bosses, anterior row usually with two bosses located on anterior half of 59
tergum between stricture and transverse sulcus, immediately behind
sulcus intermediate row with 4 small bosses, posterior margin with 6
toothed projections (Fig. 11). Ifedially to the three internal rows
are an anterior and posterior boss. Also a single lateral boss is
present. Setae, usually filiform, may be associated with bosses.
Collum wider than head, rounded, unflared, and may have denticles
and setae.
Paranota flat, expanded laterally, dorsally with one large
medial boss and two elongated lateral bosses. Paranotal W/T ratio
1.6-1.9 and tergal ratio 1.0-1.2. Lateral edge strai^t with 3 or 4
minute denticles, 4 occurring on poriferous segments. Denticles may
each have a single filiform seta. Ozopores small, not elevated,
located on posterior half of paranota, opening dorsally on small
rounded convex bosses, in 20-segmented species, on segments 5, 7, 9,
10, 12, 13, 15, 16, 17, and 18 and in 19-segmented species on
segments 5, 7, 9, 10, 12, 13, 15, 16, and 17. Posterior paranota progressively more caudally directed.
Epiproct smooth, truncate, with 8 pairs of setae, sometimes deflexed. Hypoproct small, moderately rounded, with a pair of small posteriolateral seta-bearing knobs. Paraprocts smooth with moderately thick lips.
Sterna cruciately impressed, but otherwise unmodified, each quadrant usually without setae. Sterna between leg pairs 3, 4, 5, and 6 sli^tly produced into small, blunt, paramedial lobes. Legs long, approximately 17-24% of total boc^ length. Coxae small and subquadrate; trochanter moderate with dorsal bulge; prefemur long 60
and curved ventrally; femur subquadrate; tibia elongate; tarsus very
long and curved sli^tly ventrally; claws short, curved, and
regularly tapered. Sphaerotrichomes on femur, tibia, and tarsus.
Gonopods, in situ, moderately e^gosed, partly retractile,
caudally directed, parallel to each other, oriented along median
axis of body, and capable of independent movement althou^ connected by a thin membrane. Prefemur and gonocoiae extend over sternum between posterior legs of sternum 7, causing legs to be displaced
laterally. Tibiotarsus and femur curving ventrally, extending
anterior to micÿoint of sternum 6.
Gonopod aperture subelliptical to circular, I/W ratio between
50 and 90%. Gonopod arcuate (Figs. 40-42). Gonocoxae large, sli^tly smaller than telopodite, with large, open, ventral concavity for gonopod retraction. Ventrolateral edge without setae. Anterior edge variously lobed. Prefemur hirsute and posteriorly directed. Seminal groove looped, extending to endomerite, terminating in a large terminal vesicle. Tibiotarsus usually single but may be extensively modified. Caudal surface with at least one processes. Acropodite generally scythe-shaped, variously bent. Endomerite usually small, flush with femur, and a small lateral process may be present.
Female simi.lar to male. Sterna flat without lobes. Legs less robust, shorter. Tarsus usually Sorter, femur smaller, prefemur not curved nor as massive, and trochanter not ejqjanded. Sphaerotrichomes may be absent. Syncoxostemum fused, massive, posteriorly flattened, enlarged ventramedially, not ejqpanded laterally, and curved slightly 61 posteriolaterally to protect c^^opods. Sternum III with median lobe
(Fig. 65).
Cyphopods usually withdrawn internally behirai second pair of legs, large, oval, and with posteriolateral projecting macrosetae on ventral edges of valves, flattened medially with membranous connection, lateral and medial surfaces with various numbers of elongate filiform setae. Receptacle small, closely associated with posterior edge of valves.
CONTENT: The number of species in the genus Polvdesmus may approach 200. It is probable that a revisional study will reduce this number. These "species" are presently divided into 26 subgenera mostly confined to the Old World. In the United States the following introduced species have been reported: Polvdesmus fEubrachvdesmus) anqustus Latzel, P. (Polvdesmus) complanatus (Linnaeus), P.
(Nomardhus) denticulus Koch, P. (Polydesmus) inconstans latzel, P.
(Hormobrachium) racovitzai Brolemann, P. (Eubrachvdesmus) superus
Latzel.
DISTRIBUTION: Native to the Palearctic region. The Nearctic representatives are introduced, with several now established throu^out many areas of primarily urban North America. Several species have also been introduced into other temperate and semitropical regions throughout the world. 62
NOIES: Several new species have been described from the New
World, all of #iich have subsequently been determined to be synonyms
or placed in other genera.
Those in other genera are: Brachvdesmus califomicus
Chamberlin, 1918 (= Bidentoaon^, B. hastingsus Chamberlin, 1941 (=
Iheatodesfflus) . B. vosemitensis Causey, 1954, and B. bituberculatus
Loomis, 1960 (both = Coronodesmus).
The following species have previously been synonymized with
Polvdesmus inconstans Latzel, 1884: Polvdesmus testi Bollman, 1888,
from Indianapolis, Indiana, P. socamius Chamberlin, 1910, from Salt
lake City, Utah, P. hortus Williams and Hefner, 1928, from Athens
Co., Ohio, P. proncmeutes Chamberlin, 1942b, from Fort Collins,
Colorado, and P. vheeleri Causey, 1950, from Grand Forks, North
Dakota.
The following species have previously been synonymized with
Brachvdesmus superus Latzel: P. dux Chamberlin, 1940, from Durham,
North Carolina, P. pallidus Loomis, 1939, from Charleston, West
Virginia, and P. gladiolus Williams and Hefner 1928, from Allen Co.,
Ohio. Brachvdesmus has recently been defined as a subgenus of
Polvdesmus (Hoffman, 1979).
One species name has had a varied and troubled history.
Polvdesmus moniliaris Koch (1847) has been used for different species in eastern North America by several authors because of a poor description, lack of a locality, and the loss of the type.
Pseudopolvdesmus serratus. P. collinus. and Polvdesmus inconstans have at one time or another been identified as moniliaris as have 63
several unrecognizable taxa. The positive identification of moniliaris is still in doubt.
The Palearctic genera of the subfamily included:
Archiopolvdesmus. Attems, 1898; Polvdesmus. Latreille, 1802;
Jaxartes, Veriioeff, 1930; Kiroisdesmus. Lohmander, 1933;
Schizoturanus. Verhoeff, 1931; Schizomeritius. Verhoeff, 1931;
Turanodesmus. Lohmander, 1933; Usbekodesmus. Lohmander, 1933;
Cretodesmus. Strasser, 1974; and Serradium. Strasser, 1974. The dominant genus is Polvdesmus with over 200 species recorded; the relationships of the remaining genera vAiich total 25 species is uncertain. They may very well be synonymized with Polvdesmus.
The most recent revision of Polvdesmus was by Attems (1898), prior to the description of many of the species. Verhoeff (1941), reorganized the superfamily and in the process violated virtually every principle of logical systematic procedure. For exairple, he invalidly proposed most of the 26 subgenera. His higher classification was based on various misconceptions and errors viiich was swiftly disregarded by other workers. However, his lower classification, at generic and specific levels, was blindly accepted, despite the fact that Attems did not designate types; this is a major stumbling block, to the understanding of the genus.
Attems' (1940) massive work on the complex was impressive in his determination but also lead to much confusion. In most cases, his descriptions and figures were just reediting of earlier or the original works. He took little or no time to understand or examine the specimens in detail. Many of the earlier genera, subgenera under 64
Attems, were and still are based principly on differences in segment number.
The first major step in cleaning qp the genus Polvdesmus will be the simplification and standardization of gonopod terminology. A list of gonopod processes from Attems' (1940) revision includes:
terminal: s
medial: p, q, i, lo, f, 1, a, F, Z^, Z2
prefemur: a
lateral: T, I 2 , k, z, n, A, ha, b, u, d, c, Q, r, h
The proliferation of medial and lateral processes is due to the recognition of many minor variations.
Since 1940, other authors have described new species and have added more new names for these processes. A system similar to that in Pseudopolvdesmus needs to be set vp to handle this problem. A complete, modem revision of this genus will take many years.
Pseudopolvdesmus Attems, 1898
(Figs. 12, 19, 45, 46, 48, 50, 55, 57, 59, 63
66, 67; Map 3)
Polvdesmus of American authors prior to 1943 (not Latreille, 1802). Pseudopolvdesmus Attems, 1898, Denkschr. Akad. Wiss., Wien, 67:270; 1940, Das Tierreich, 70:139.— Chamberlin, 1943c, Bull. Univ. Utah, biol. ser., 8(2): 17. Dixidesmus Chamberlin, 1943, Bull. Univ. Utah, biol. ser., 8(2) : 18.- - Chamberlin and Hoffman, 1958, U. S. Nat. Mrs. Bull. 212:65. (Type species: D. tallulanus Chamberlin, by original designation.) Synonymy by Hoffman, 1974. 65
GENEEXDTYPE; Polvdesmus canadensis Newport, 1844, b y jmonotypy.
DIAGNOSIS: Small to large, 20 segments, brown or black, sometimes with red paranota. Antennae usually elongate and filiform, sometimes sli^tly clavate. Tergites quadrate, with connected medial bosses but without tubercles. Paranota large and horizontal.
Ozopores not elevated, opening dorsally. Ozopore formula normal.
Male sterna sli^tly modified. Legs unmodified. Gonopods arcuate.
Gonocoxae large, triangular with ventral concavity. Prefemur moderate, hirsute, without processes, attached to posterior edge of gonocoxae. Femur short, curved, terminating at large endomerite.
Tibiotarsus elorgate, usually sli(ÿitly curved, with simple caudalad processes. Seminal groove looped, with large terminal vesicle at endomerite. Female legs thin, unmodified, without sphaerotrichomes.
Syncoxostemum basally thickened, fused, and distally separated; lateral spines absent. Sternum III unmodified, cyphopods kidney shaped, dorsal plate present, macrosetae moderate, numbering between
10 and 18. Operculum small. Receptacle absent.
CONTENT: Two groi:ps totaling nine species can be distinguished by gonopodal differences: Group Serratus, consists of serratus. paludicola. caddo. and minor, and Group Canadensis consists of pinetorum. tallulanus. erasus. canadensis, and collinus.
DESCRIPHON: Small to large (9-32mm) polydesmids, with 20 65
segments not including head (Fig. 19). W/L 12.2-22.0% and WL 38.0-
164.8mm^.
Color in life usually brown or black metaterga althou^
paranota may be red. Venter, legs, and genae usually li ^ t brown.
Antennomeres darker distally, with 7th darkest.
Head smooth. Epicranial suture distinct, not extending to
antennae. Vertex and frons equally large. Gena with large lateral projection. Parietal sclerite small, adjacent to Organ of Tomosvary.
Mandible with large cardo and stipes, triangular, projecting
laterally, with incomplete suture dorsally. Antennae well separated,
set medially on face, usually filiform, 10-25% of body length, and
extending to posterior edge of 4th tergite (Fig. 57). Antermomeric
lengths in increasing sequence 7<1<6<4=2<3<5. First antennomere
rectangular, 2-5 moderately long, subquadrate, and usually very
sli^tly clavate, 6 elongate, and 7 quadrate with 4 terminal sensillae. Antennomeres 5, 6, and 7 normal, each with a small dorsodistal patch of bacilliform setae.
Facial setae few, long, and filiform. Distribution as follows: subantennal, several; frontal numerous and evenly spaced; clypeal 3 pairs; labral 3 pairs; genal 9-13 pairs; postantennal none.
Labrum with 3 small, blunt teeth, medial tooth largest.
Mandible with one external, broadly-rounded, single-cusped tooth; 6 round, flat, single-cusped internal teeth, 6 single-toothed pectinate lamellae, and molar plate with several deep furrows with anteriorly curved monocusped fringe teeth. Epipharynx with large, medial, teardrop-shaped keel without teeth. Lateral keels very 67
dentate, almost touching medial keel anteriorly. Retrorse lobe present. Retinaculum arched. Hypcpharynx with anterior teeth.
Gnathochilarium typical but relatively short, quadrate, W/L
ratio approximately 100%. Stipes large, projecting caudally, lateral
edge flaired slightly. Setae numbering 9-20. Anterior palps with 9-
20 macrosetae. lamellae linguales small and elongate; palps with 7-
11 scattered setae and two large dissimilar anteriomedial macrosetae. Mentum triangular, extending slightly caudal to stipes.
Cardo small, triangular. Erebasilare elongate. lypostoma short.
Metaterga (Fig. 12) essentially flat, not tuberculate, each with three transverse rows of medial quadrate convex bosses, anterior row usually with two bosses located on anterior half of tergum between stricture and transverse sulcus intermediate row posterior to sulcus, with 4 bosses, and posterior margin with 6 bosses, with a single elongate lateral boss. On caudal segments posterior row projects si^tly caudally beyond margin. Filiform setae (Fig. 50) may be associated with these quadrate areas.
Erozonite completely covered in small pits. Collum moderately large, wider than head, rounded, not flaired, and sometimes with several lateral denticles and setae.
Paranota expanded laterally and horizontally, T/W ratio approximately 1.6 for paranota and 1.0 for tergites. lateral edges curved, with 3 or 4 minute denticles, 4 occurring on poriferous segments. Denticles may have one or two filiform setae. Ozopores small, not elevated, located on posterior half of paranotum, opening dorsally, on segments 5, 7, 9, 10, 12, 13, 15, 16, 17, and 18. 68
Peritremata thicker on poriferous segments. Posterior paranota progressively more caudally directed.
Epiproct smooth, truncated, with 8 pairs of setae, and may be deflexed. Hypoproct small, rounded, with single pair of small seta- bearing posteriolateral swellings. Paraprocts smooth, with two pairs of lateral setae; lips thin to moderate.
Sterna cruciately inpressed, each quadrant usually moderately setose (Fig. 59). Sterna between legs 3, 4, 5, 6 , 9, 11, and 13, produced into paired prominent, angular, blunt, paramedial lobes, tegs long, 14-25% of total boc^ length. Coxa small and subquadrate; trochanter moderate and clavate; prefemur curved and massive; femur lorg and curving; tibia subquadate; tarsus elongate; claw (Fig. 55) moderately long, sittple, and tapered gradually. Tarsus, tibia, and femur of males with sphaerotrichomes, and large ventral mammulations, each with a single proximally directed seta.
Gonopods, in situ, moderately e>^sed, partly retractile, directed caudad, parallel, and oriented along median axis of body, capable of moving independently, althouiÿi attached medially by thin membrane. Prefemur and gonocoxa extend over sternum between posterior legs of segment 7, causing legs to be displaced laterally.
Tibiotarsus directed anterioventrally, extending over anterior margin of segment 6 .
Goncpod aperture subelliptical with I/W ratio of 55-70%.
Gonocoxa large, slightly smaller than telopodite, with large, open, ventral concavity for retraction of gonopod, ventrolateral edge with two setae, anterior edge variously lobed. Prefemur very hirsute and 69
squared posteriorly. Cannula connected to telopodite under smaill
ledge. Femur curved and long. Seminal groove looped, extending to
endomerite vdiere it terminates in a large internal vesicle.
Tibiotarsus single, not divided. Caudal surface with various
processes, mesal (Mi_4 ) and ectal (Ei-^). Process located
proximal to endomerite. All processes directed caudad except E 2
viAiich is directed laterad. Process M 4 usually adjacent to small
distal setal series. Tip may be bent or strai^t, with small linear
series of flattened setae. Telopodite scythe-shaped or variously
bent: from subevenly curved to bent with one or two proximal or
distal angulations. Many variations occur. If a proximal bend is
present, the angle usually measures between 118° and 137°, vÆiile a
distal bend, if present, measures between 111° and 140°. The
telopodite tip is usually sharply hooked, or occasionally slightly
curved. The terminal setal tuft appears highly variable, primarily
due to breakage. Endomerite large, sloping away from telopodite.
Female very similar to male, except sterna flat without lobes,
legs not as robust as in males, tarsus usually shorter, femur
smaller, prefemur not curved or as massive, and trochanter not
expanded. Sphaerotrichomes absent. Syncoxostemum (Fig. 63) fused, massive, posteriorly flattened and curved sli<ÿitly
posteriolaterally. Sternum III (Fig. 6 6 ) slightly lobed ventrally.
cyphopods (Fig. 67) large, kidney-shaped, and usually withdrawn internally behind second pair of legs. Valves large,
flattened medially, lateral and medial surfaces with various numbers
of elongate setae, with 10-18 caudad macrosetae on dorsal edge. 70
Dorsal surface ej^anded into irregular plate. Operculum small,
closely associated with ventral surface of valves. Receptacle
absent.
DISTRIBUTION: Nearctic region (Map 3). Southeastern Canada west to Minnesota, south to northern Florida and eastern Texas.
Usually found in leaf duff of deciduous forests.
Key to males of the species of Pseudopolvdesmus
la. Process M 2 absent; process E 3 present; cyphopodal plate large and with long ventral setae; body with small posterior sternal lobes on 7th segment...... Groi:ç) Canadensis...... 2
lb. Process M 2 present; process E 3 absent; gonocoxae multi-lobed. Process M]^ & E^ connected by ventral ridge and with large posterior sternal lobes on 7th segment...... Group Serratus...... 6
2a. Process E^ present; denticles large; epiproct downtumed sliÿitly...... pinetorum
2b. Process E^ absent; denticles small; epiproct normal..3
3a. Processes E 2 and E 3 joined on single stalk (E2 +.3 ) ; process M 2 present; telopodite thickened longitudinally...... 4
3b. Processes E 2 and E 3 not joined; process M 2 absent; telopodite thickened longitudinally...... 5
4a. Process E 3 absent; found on Cumberland Plateau, .erasus
4b. Process E 3 present; found in southern ^palachians ...... tallulanus
5a. Process M 3 present; process M^ distal to base of endomerite; usually with red paranota; gonocoxae monoldbed...... canadensis 71
5b. Process M 3 absent; process M^ next to endomerite; brown paranota; gonocoxae multi-lobed...... collinus
6 a. Process M 4 present; large, greater than 20mm; cyphopod elongate; sometimes with red paranota; dorsal setae absent; habitat generalized..... serratus
6 b. Process M 4 absent; small less than 20mm; cy^opod oval; dorsal setae present; brown paranota; habitat lowlands...... 7
7a. Process £ 3 absent; M^ and £ 3 connected by posterior ridge...... paludicola
7b. Process £ 3 present; and £ 3 not connected...... 8
8 a. Process M 3 present; process M 4 absent; boc^ flattened, W/L ratio greater than 70%; sternum II of female modified...... minor
8 b. Process M 3 absent; process M 4 present; body much flattened, W/L ratio less than 80%; sternum II unmodified...... caddo
Group Canadensis
The Group Canadensis consists of five species,
Pseudopolvdesmus pinetorum. tallulanus. erasus. canadensis, and
collinus. Jpomorphies for the group are: not connected to £ 3 , posterior sternal lobe on 7th segment small, cyphopodal plate small,
long sternal setae, paranota with small denticles, and process £^ usually present. These are medium to large millipeds of the genus.
They have paranota sli^tly rounded laterally and posteriolateral
comer sli«ÿitly produced caudally, sitrple gonocoxae, cyphopod
elongate with laterally expanded plate, legs longer than in Group
Serratus, with longer trochanters and smaller coxae. The Appalachian 72
Mountains region has canadensis, collinus. erasus. and tallulanus, vSiile pinetorum occurs to the south and west.
Pseudopolvdesmus pinetorum (Bollman, 1888)
(Figs. 74, 80, 82, 8 6 , 90, 106, 111, 122-126?
Map 4; Tables 9-11)
Polvdesmus pinetorum Bollman, 1888, Ent. Amer. 4:3 (type locality: Arkansas, Pulaski Co., Little Rock; type: USNM). Polvdesmus americanus Carl, 1902, Rev. Suisse Zool., 10:611, fig. 37 (type locality: "Texas"; type: ΠN H ) . Polvdesmus natdhitoches Chamberlin, 1942, Bull. Univ. Utah, biol. ser., 6 (8 ) :10, figs. 34, 35 (type locality: Louisiana, Natxhitoches Par., 2mi. S. Saline; type: USNM). NEW SYNONYMY. Polvdesmus paroicus Chamberlin, 1942, Bull. Univ. Utiah, biol. ser., 6 (8 ): 11, figs. 37, 38 (type locality: Louisiana, Jackson Par., l.Smi. N. Clay; type: USNM). Polvdesmus modocus Chamberlin, 1943, Proc. Biol. Soc. Washington, 56:36, fig. 6 (type locality: Illinois, Randolph Co., between Ifodoc and Roots; type: FMNK). Polvdesmus hubrichti Chamberlin, 1943, Ent. News, 54:15, figs. 1, 2 (type locality: Missouri, St. Louis Co., St. Louis; type: ANSP). Polvdesmus medicus Chamberlin, 1943, Proc. Biol. Soc. Washington, 56:40, fig. 6 . Lapsus for modocus. Pseudopolvdesmus pinetorum. Causey, 1952, Chicago Acad. Sci. Nat. Hist. Misc. No. 106, p. 6 .
DIAGNOSIS: Medium to large (ll-28mm). Metiaterga and paranotza brown. Dorsal and paranotial setae absent. Dorsal sculpture medium.
Antennae short. Posterior sternum of 7th segment small. Gonocoxae monolobed. Goncpodal process M^ small, on base of endomerite, M 3 small to medium, M 4 small to medium, E^ small, if present on lateral edge, E 2 medium, E 3 small, E 4 small, cyphopods not es^ended 73
dorsally. cyphopod plate mediolateral lobe rounded, lateral edge with several small teeth.
DESCKEPTION: (Based on adult male and female from Missouri,
Barry Co., Roaring River State Park, VI-21-1983, C. P. Withrow,
(CPWC)). Adult male, 16.6mm long, maximum width (7th segment),
2.6mm, W/L 15.7%, and WL 43.2mm^. Segmental widths across paranota
as follows:
Collum 1 . 8 m m 9-13 2.4 ram
2 2 . 1 m m 14-15 2.3 mm
3-4 2 . 2 m m 16 2 . 2 m m
5 2.3 m m 17 2 . 0 m m
6 2.5 m m 18 1 . 6 m m
7 2 . 6 m m 19 1 . 0 ram
8 2.5 m m 2 0 0.4 m m
Color in life brown, paranota lighter.
Head typical for genus. Head width 1.06mm, interantennal distance 0.32, 30.2% of total head width. Antennae moderate
(2.49mm), 15.0% of total body length. An-tennomeric lengths (% of total antennal length); 1st antennomere, 0.24mm (9.8%); 2nd, 0.42mm
(16.8%); 3rd, 0.6.2mm (24.8%); 4th, 0.34mm (13.6%); 5th, 0.41mm
(16.6%); 6 th, 0.44mm (17.5%); 7th, 0.13mm (5.0%).
Dorsal sculpture moderately defined. Collum moderate, with acute posteriolateral angle and anteriomedial bulge. Paranota with 74 anterior, lateral, and posterior edges straight, anterior comer sliÿitly cephalad. Medial boss large, anterior bosses slightly separated (Fig. 74). Lateral denticles normal, located from anteriolateral comer (% of total length of 8 th segment) ; 1 st denticle, 0.04mm (6 .6 %); 2nd, 0.13mm (21.3%); 3rd, 0.25mm (41.0%);
4th, 0.46mm (75.4%). Posteriomedial teeth distinct on segments 15-
19. Ventrolateral surface of paranota r o u ^ and irregular.
Sterna cruciately inpressed, each quadrant moderately setose.
Stemum 2 edge sharply indented anteriomedially, posteriomedial margin also indented, with small medial cross-ridge. Stemum 3 posterior heavy margined, anterior edge moderately inflated, moderately r o u ^ without setae posteriomedially between penes, small, truncated anteriomedial swelling, posterior plate field moderate, and penes typical although margined laterally. Stemum 4 with 5-6 long setae and numerous short stubby setae. Stemum 5 similar to 4 but sternal lobes more defined. Stemum 6 typically shaped except posterior legs moderately displaced laterally, median depression striated, broad, and moderately deep. Posterior stemum of 7th segment with small pointed lobe. leg shape typical and short.
Length of riÿit posterior leg of 8 th segment 3.07mm, 18.5% of total body length. Length of podomeres (% of total leg length): coxae,
0.27mm (8.9%); trochanter, 0.47mm (15.3%); prefemur, 0.63mm (20.4%); femur, 0.37mm (12.1%); tibia, 0.31mm (10.2%); tarsus, 0.84mm
(27.4%); claw, 0.17mm (5.7%). Sphaerotrichomes typical. 75
E$)iproct strai^t (Fig. 80). Hypoproct and paraprocts normal.
Gonopod aperture subelliptical (0.84mm long, 1.17mm wide),
with I/W ratio of 71.8%. Gonocoxae large (0.81mm hi^, 1.03mm long),
posterioventral edge monolobed, blunt, extending outward twice its width, sides parallel, smooth, and rounded. Telopodite (Figs. 82,
8 6 , 90) moderate (1.10mm long). Prefemur normal. Femur thin with
rounded femoral ledge. Tibiotarsus typically shaped, small
(0.05mm), irregular, located 0.45mm (40.9%) from tip, on base of
endomerite; M 3 small (0.05mm), finned, 0.23mm (20.9%) from tip; M 4 medium (0.06mm), having a second smaller lobe for setal tuft, 0.14mm
(12.7%) from tip; very small (0.005mm), 0.40mm (36.4%) from tip;
E 2 medium (0.06mm), hooked, 0.27mm (24.8%) from tip; E 3 small
(0.01mm) and hooked, 0.17mm (16.3%) from tip; E 4 small (0.02mm), triangular, 0.08mm (7.3%) from tip. Tip slightly lobate. Telopodite with distal and proximal curves. Seminal groove typical. Telopodite thickness measured at femoral ledge perpendicular to body axis,
0.09mm, parallel to axis, 0.20mm. Tibiotarsus thickened proximal to
seminal groove. Endomerite blunt, 0.42mm (26.2%) from tip, extending
0.23mm on telopodite, projecting 0.13mm h i ^ and 0.21mm long.
Female 20.15mm long, maximum width 2.96mm, W/L 14.7%, and WL
59.6mm^. Segmental widths across paranota as follows:
Collum 2.2 mm 9-11 2.9 mm
2 2.5 mm 12-15 2.8 mm
3 2.6mm 16 2.6 mm
4 2.7 mm 17 2.5 mm 76
5 2.8 iron 18 2.1 im
6-7 2.9 mm 19 1.3 mm
8 3.0 mm 20 0.5 ram
Color and structure similar to male. Sterna and legs unmodified. Syncoxostemum strai^t laterally, quadrate and rough distally. Sternum III rounded distally, notched, and curved gently laterally, and rouÿi medially.
cyphopods (Figs. 106, 111) coirpact (0.53mm long, 0.24mm h i ^ ) .
Valves with 9 long macrosetae projecting from lateral half over intervalvular opening, and dorsal edge not flattened, cyphopod plate with distomedial lobe rounded, smooth, lateral edge with small notch containing 2-3 small teeth. Operculum small (0.11mm long, 0.18mm h i ^ ) and thin.
VARIATION: Based on 715 specimens, Pseudopolvdesmus pinetorum althou^ wide-ranging, showed slight size variation with little or no geographic significance. Based on 212 specimens (112 males, 100 females) measured, the means of the boc^ measurements for males and females, respectively, were: length 18.60mm, 18.70mm; width 3.1mm,
3.1mm; W/L 16.45%, 16.60%; IW 57.65mm^, 59.37mm^. The ranges and standard deviations are given in Tables 9, 10, and Figures 122-125.
Females are slightly longer than males. Females also have a hiiÿier
W/L ratio and dorsal area.
Based on ten specimens, the gonopodal processes varied little in their relative position on the tibiotarsus (Fig. 126). The 77 positions of the processes relative to telopodite length are:
56.15%, M 3 22.0%, M 4 13.6%, £ 3 27.55%, E 3 16.3%. For actual ranges see Table 11.
Gonopod processes of Pseudopolvdesmus pinetorum varied sli^tly in size and location. In several specimens the process was absent. In a number of specimens a process distal to E 3
(designated E^^.) was present. This condition being more prevalent in the northern and eastern parts of the range. Process E 3 tended to be smaller in the northeastern segment of the range, and varied from a single process with a lateral setal tuft to a double process with the tuft on the proximal prong. Process M^ was irregularly shaped.
Processes were generally shark-fin or hook-shaped.
DISIRIBUTION: The range of Pseudopolvdesmus pinetorum covers the lower and middle Mississippi River drainages from southern
Illinois and Iowa south throuiÿi eastern fânsas, Oklahoma, and Texas; eastward to central Alabama, and sporadically in central Tennessee and eastern Kentucky (Map 4). The distribution east of the
Mississippi River is not clear, due inadequate collecting. Records from central Kentucky and eastern Tennessee are very suspect, although those specimens from eastern Tennessee were examined and verified as pinetorum. I believe that the locality labels were somehow mixed vç). Pseudopolvdesmus pinetorum is the dominant species west of the Mississippi River and is most prevalent in the Ozark
Mountain region. The habitat is typical for the genus. 78
NOTES: Polvdesmus americanus Carl, holotype examined, was synonymized by Causey (1952a). Carl's description was atypical in that it was done well. All telopoditic processes were visible in his drawing.
In the original description, Polvdesmus natchitoches
Chamberlin, holotype examined, was illustrated in medial view with an enlargement of the telopodite tip, endomerite without setae, absent, and M 3 absent, and in distal view, M 3 was present as well as a small £ 3 +. The last structure is not present on tlae type!
Polvdesmus paroicus Chamberlin, holot^'pe examined, was synonymized by Causey (1952a). The gonopod figures in the description were again misdrawn. In the mesal view, M]_ was absent, with process £ 3 absent from the distal view..
Polydesmus modocus Chamberlin, holotype examined, was also synonymized with pinetorum by Causey (1952a). The Chamberlin (1943a) figure shows M^ missing. In his paper Chamberlin also drew two figures clearly representative of pinetorum but with no statement or identification in the text or key. While examining types from the
Chamberlin collection at the U. S. National Museum, a manuscript holotype identified as "Polvdesmus scholasticus" was found. It was collected from St. Louis, Missouri but a description was never published. This specimen may be the source of those figures.
Althou^i a type existzs, the name scholasticus has not been validly published, and my mention of it here is not to be constunied as validation. 79
Polydesmus hubrichti Chamberlin, holotype examined, was
synonymized by Causey (1952a). The gonopod drawings showed process
absent in medial view but present in distal view.
Pseudopolvdesmus tallulanus (Chamberlin, 1943) NEW OCMB.
(Figs. 75, 83, 87, 91, 107, 112, 122-126;
Map 5; Tables 9-11)
Dixidesmus tallulanus Chamberlin, 1943, Bull. Univ. Utah, biol. ser. 8(2): 19, fig. 34 (type locality: Georgia, Rabun Co., between Clayton and Tallulah Falls; type: USNM). Dixidesmus penicillus Chamberlin, 1943, Bull. Univ. Utah, biol. ser. 8(2) :19, fig. 35 (type locality: Georgia, Habersham Co., Clarkesville; type: USNM). NEW SYNONYMY. Dixidesmus humilidens Chamberlin, 1943, Bull. Univ. Utah, biol. ser. 8(2) :20, fig. 36 (type locality: Georgia, Hall Co., Gainesville; type: USNM). NEW SYNONYMY.
DIAGNOSIS: Medium to large (18-26mm). Metaterga brown with paranota red or brown. Dorsal sculpture li^t. Small dorsal and paranotal setae present. Denticles small. Antennae long. Gonocoxae monolobed, ventrodorsally large, smooth, and pointed. Telopodite thick longitudinally. Gonopodal process M^ medium, next to endomerite; M 3 medium; M 4 small with small setal pad; E^ size variable distal to endomerite; E 2 large, projecting posteriorly; E 3 medium; E 4 small, subterminal, cyphopod elongate, with sli^t lateral twist and small lateral projection.
DESCRIPTION: (Based on adult male and female from North
Carolina, Transylvania Co., 12.1mi. WSW. of Rosman, County Route 80
1152, 0.8ml. N. of CR. 1151, VlII-28-1975, R. M. Shelley, (NCMH) ).
M u l t male 25.7mm long, maximum width (7th segment), 4.7mm, and maximum thickness, 3.0mm. W/L 18.3% and WL 120.8 mm^. Segmental widths across paranota as follows:
Collum 2.9 mm 1 1 - 1 2 4.4 m m
2 3.5 m m 13 4.1 m m
3 3.8 mm 14 4.0 mm
4 4.0 m m 15 3.9 m m
5 4.2 mm 16 3.8 mm
6 4.6 mm 17 3.4 m m
7 4.7 mm 18 2 . 6 m m
8 4.6 m m 19 1 . 6 m m
9-10 4.5 m m 2 0 1 . 0 m m
Color in life, brown.
Head typical for genus. Head width 0.92mm, interantennal distance 0.23mm, 25.3% of total head width. Antennae long (4.44mm),
17.3% of total body length. Antennameric lengths (% of total antennae length): 1st antennomere, 0.22mm (5.0%); 2nd, 0.55mm
(12.4%); 3rd, 1.07mm (24.1%); 4th, 0.84mm (18.9%); 5th, 0.85mm
(19.1%); 6 th, 0.68mm (15.3%); 7th, 0.23mm (5.2%).
Dorsal sculpture slight. Lateral boss of paranotum elongate, anglirg anteriomedially, anteriomedial bosses well separated, medial boss large, equidistant from anterior and posterior edges (Fig. 75).
Collum moderate with rounded posteriolateral angle, medial anterior 81
bulge, and indented posteriomedially with single pair of denticles.
Paranota with anterior edge strai^t, posterior margin indented.
Denticles small, usually distinct, located from anteriolateral
comer (% of total length of paranotum, 8 th segment) : 1 st denticle,
0.10mm (7.2%); 2nd, 0.28mm (20.2%); 3rd, 0.67mm (47.6%); 4th, 0.95mm
(67.9%). Ventrolateral surface of paranota r ou^ and irregular.
Sterna cruciately impressed, each quadrant heavily setose.
Sternum 2 sharply indented anteriorly, smoothly indented
posteriorly, with wide r o u ^ border. Sternum 3 with small medial
swellings, not as rouÿi medially, penes normal, one posteriomedial
seta and two anteriomedial setae. Sterna 4 and 5 similar, with many
lateral setae. Sternum 6 anterior lobes widely separated, with short
and long setae; posterior sternal lobes sli^tly displaced
anteriorly, with numerous short setae, longitudinal groove inverted
V-shaped, deep, with distinct contour ridges. Remaining stema hirsute. Posterior sternum of 7th segment elongate, extending ventrally close to coxae. Leg shape typical and short. length of ri^t posterior leg of 8 th segment 5.10mm, 19.8% of total boc^ length. Length of leg segments (% of total leg length) : coxa, 0.44mm
(8 .6 %); trochanter, 0.96mm (18.8%); prefemur, 0.93mm (18.4%); femur,
0.50mm (9.8%); tibia, 0.55mm (10.6%); tarsus, 1.38mm (27.5%); claw,
0.32mm (6.3%), sli^tly curved. Sphaerotrichames typical.
Epiproct sli^tly downtumed. Hypoproct and paraprocts typical for genus.
Gonopod aperture subelliptical (1.27mm long, 1.89mm wide) with
I/W ratio of 67.3%. Gonocoxae large (0.92mm high, 1.57mm long). 82
triangular, slightly trilobed. Telopodite (Figs. 83, 87, 91) large
(1.68mm long). Prefemur irregular with small mound 1.35mm from distal tip. Femur thickened, femoral ledge round. Tibiotarsus t^ically shaped, medium (0.08mm), located 0.84mm (53.6%) from tip; M 3 medium (0.09mm), 0.36mm (22.9%) from tip; M 4 small (0.04mm),
0.21mm (13.4%) from tip; medium (0.08mm), 0.63mm (38.9%) from tip, distally on base of endomerite; E 2 large (0.14mm), 0.44mm
(28.0%) from tip; E 3 medium (0.08mm), 0.52mm (33.1%) from tip; E 4 small (0.03mm), 0.13mm (8.3%) from tip. Tip slightly Ibent.
Telopodite with distal bend 0.36mm from tip. Tîbiotarsal thickness measured at femoral ledge, perpendicular to body axis, 0.03mm, and parallel to axis, 0.03mm. Seminal groove normal. Endomerite typical,
0.72mm (46.1%) from tip, extending 0.28mm on telopodite, projecting
0.12mm high and 0.41mm long.
Female 23.3mm long,maximum width 4.5mm, W/L 19.3%, and WL
104.9mm^. Segmental widths across paranota as follows:
Collum 2.3 m m 13 4.5 mm
2 3.7 mm 14 4.3 mm
3 3.8 m m 15 4.2 ram
4 4.0 m m 16 4.0 ram
5 4.2 m m 17 3.5 ram
6 4.3 mm 18 2.7 mm
7 4.4 mm 19 1.7 ram
8-9 4.5 m m 2 0 0 . 8 mm
1 0 - 1 2 4.6 m m 83
Color and structure similar to male. Stema and legs
unmodified. Syncoxostemum wide medially, ventrolateral shelf small,
straiÿit laterally, r o u ^ anteriorly. Sternum III flattened,
projecting slightly ventrolaterally, and posteriorly smooth with
small medial ridge.
cyphopods (Figs. 107, 112) large (0.90mm long, 0.42mm high).
Valves not ejçanded, lateral setae moderate, cyphopodal plate
twisted laterally, projecting sli^tly laterally, sloping medially,
not curving. Operculum small with numerous long anterior curving
setae.
VARIATION: Based on approximately 250 specimens,
PseudoDolvdesmus tallulanus. showed slight size variation with
little or no geographical significance. Based on 55 (33 males, 22
females) specimens measured, the means of the boc^ measurements for males and females, respectively, were: length 23.27mm, 23.25mm; width 4.28mm, 3.95mm; W/L 18.39%, 15.99%; IW 99.60m^, 91.82mm^. The ranges and standard deviations can be seen in Tables 9, 10, and
Figures 122-125.
Based on ten specimens, the gonopodal processes varied little in their relative position on the telopodite (Fig. 126). The positions of the processes relative to telopodite length are:
52.6%, Mg 22.45%, Mg 10.9%, 42.35%, Eg 32.8%, Eg 22.9%, E4 8.65%.
For actual ranges see Table 11. The most notable variation observed is the length of process Eg ihich ranges from a small tooth to an elongate ezpine. 84
DISTRIBUTION: The range of PseudoDolvdesnnis tallulanus covers
the eastern slopes of the southern Appalachians from southwest North
Carolina and extreme eastern Tennessee into northwestern South
Carolina and northern Georgia (Map 5). The western limit of this
species borders its sister species Pseudopolvdesmus erasus. Based on
the localities recordel for both species, they appear to be
allopatric, althou^ this may be a collecting artifact.
NOTES: Dixidesmus oenicillus Chamberlin, holotype examinai.
Chamberlin's drawing of the gonopod shows sharper bends in the tibiotarsus and a smaller process E 3 than actually present in tallulanus and his holotype.
Dixidesmus humilidens Chamberlin, holotype examined.
Chamberlin's figure ^ows a smooth curvature of the tibiotarsus, also E 3 smaller than that of tallulanus. The species was said to be smaller than tallulanus although syrpatric.
Pseudopolvdesmus erasus (loomis, 1943) NEW OCMB.
(Figs. 84, 8 8 , 92, 108, 113, 122-126;
Map 5; Tables 9-11)
Polvdesmus erasus loomis, 1943, Bull. Mus. Comp. Zool., 92(7):406, fig. 17, and pi. 5, fig. 4 (type locality: Alabama, Madison Co., Huntsville; type: MCZC). Dixidesmus erasus. Causey, 1952, Chicago Acad. Sci. Nat. Hist. Misc. No. 106, p. 7. 85
DIAGNOSIS; Medium (15.8-24.0mm). Metaterga brown with reddish paranota. Dorsal sculpture weak, similar to tallulanus. Denticles small. Dorsal and paranotal setae small. Antennae moderately long.
Gonocoxae monolobed, sharp and thickened anteriorly. Gonopodal process medium, regular, close to base of endomerite; M 3 large, distal to E 3 ,* M 4 medium; large, on distal side of endomerite, E 2 large, basally directed; E 4 small, subterminal. Telopodite thick longitudally. Endomerite typical, cyphopods large, elongate, with many lateral setae. Valves with marginal macrosetae. cyphopodal plate with slight lateral twist, and small lateral projection.
DESCRIPTION: (Based on adult male and female from Tennessee,
Bradley Co., US Rt. 75, 25mi. S. of Cleveland, IV-14-1978, C. P.
Withrow, (CPWC) ). Adult male, 19.8mm long, maximum width (8 th segment), 3.2mm. W/L 16.2%, and WL 63.4 ram^. Segmental widths across paranota as follows:
Collum 2 . 2 mm 12-13 3.0 m m
2 2.7 ram 14-15 2.9 mm
3 2.9 mm 16 2.7 m m
4 3.0 mm 17 2.4 ram
5 3.1 mm 18 2 . 0 ram
6 - 1 0 3.2 mm 19 1.3 ram
1 1 3.1 mm 2 0 0 . 6 ram 86
Color in life, typically li«ÿit brown with reddish paranota.
Head typical for genus. Head width 1.72mm, interantennal distance 0.45mm, 26.1% of total head width. Antennae robust, moderately long (3.61mm), 18.2% of total body length. Antennomeric
lengths (% of total antennal length); 1st antennomere, 0.29mm
(8.0%); 2nd, 0.39mm (10.8%); 3rd, 0.82mm (22.7%); 4th, 0.67mm
(18.6%); 5th, 0.64mm (17.7%); 6 th, 0.55mm (15.2%); 7th, 0.25mm
(6.9%).
Dorsal sculpture weak, similar to tallulanus. Collum moderate, with acute posteriolateral angle, longer medially, and indented posteriomedially. Paranotal anterior edge straight, with posteriolateral angle acute. Dorsal and paranotal setae small.
Denticles small, located from anteriolateral comer (% of total paranotal length, 8 th segment): 1st denticle, 0.17mm (12.8%); 2nd,
0.39mm (28.9%); 3rd, 0.63mm (47.4%); and 4th, 0.99mm (75.0%).
Ventrolateral surface of paranota r o u ^ and irregular.
Stema similar to tallulanus. Leg shape normal and long, length of r i ^ t posterior leg of 8 th segment 4.17mm, 21.1% of total bo(^ length. length of leg segments (% of total leg length) : coxae,
0.37mm (8.9%); trochanter, 0.77mm (18.5%); prefemur, 0.78mm (18.7%); femur, 0.45mm (10.8%); tibia, 0.48mm (11.5%); tarsus, 1.12mm
(26.9%); claw, 0.20mm (4.8%).
Epiproct slightly down-tumed. Hypoproct and paraprocts normal.
Gonopod aperture subelliptical (0.98mm long, 1.26mm wide), with I/W ratio of 77.8%. Gonocoxae large (0.92mm high, 1.37mm long), 87
posterioventral edge monolobed, smooth, sharp, and thickened.
Telopodite (Figs. 84, 88, 92) large (1.94mm long). Prefemur
irregular with small posterioventral knob, 1.49mm (76.8%) from
distal tip. Femur thickened, femoral ledge round. Tibiotarsus
t^ically shaped, medium (0.08mm), located 0.96mm (49.9%) from
tip, next to endomerite; M 3 large (0.10mm), 0.54mm (28.3%) from tip;
M 4 medium (0.06mm), 0.37mm (19.2%) from tip; large (0.13mm),
0.94mm (48.3%) from tip, located distally on base of endomerite; E 2
large (0.14mm), 0.83mm (42.7%) from tip; E 4 small (0.05mm), 0.16mm
(8.4%) from tip. Tip curved dorsally. Telopodite with proximal bend
28.6% from tip. Seminal groove typical. Tibiotarsal thickness measured at femoral ledge, perpendicular to boc^ axis, 0.03mm, and
parallel to axis, 0.02mm. Endomerite sharp, 0.87mm (44.6%) from tip,
extending 0.35mm on telopodite, projecting 0.13mm h i ^ and 0.39mm
long.
Female 20.05mm long, maximum width, 3.25mm, W/L 16.2%, and WL
65,2mm^. Segmental widths across paranota as follows:
Collum 2.2 m m 14 2.9 ram
2 2.8 ram 15 2.8 mm
3 3.0 ram 16 2.7 ram
4 — 5 3.1 m m 17 2.5 ram
6-8 3.3 m m 18 2.0 ram
9 3.2 ram 19 1.4 ram
10-11 3.1 ram 20 0.6 ram
12-13 3.0 mm 88
Color and structure similar to male. Sterna and legs unmodified. Syncoxostemum similar to tallulanus. lateral angle gradually slopir^, medial lobe bulbous. Sternum III flattened, projecting slightly ventrolaterally, and posteriorly smooth, with small medial ridge.
cyphopods (Figs. 108, 113) elongate (1.24mm long, distally
0.77mm hi^, proximally 0.65mm high), lateral setae abundant. Dorsal plate twisted laterally, projecting sli^tly laterally, sloping medially, not curving. Operculum small with many long setae.
VARIATION: Based on approximately 200 specimens of
Pseudopolvdesmus erasus examined, little variation was observed.
Based on 8 8 (77 males, and 11 females) specimens measured, the means of the body measurements for males and females, respectively, were: length 20.38mm, 19.57mm; width 3.6 mm, 3.20mm; W/L 17.66%, 16.35%;
IW 73.37mm^, 62.62mm^. The ranges and standard deviations can be seen on Tables 9, 10, and Figures 122-125.
Normal variation in location and size of processes was present. Based on ten specimens, the gonopodal processes varied little in their relative position on the tibiotarsus (Fig. 126). The positions of the processes relative to telopodite length are:
55.45%, M 2 19.5%, M 3 11.45%, 40.5%, E 2 30.2%, E 4 8.0%. For actual ranges see Table 11.
On several specimens a small shoulder spine was located on E 2 , however this spine is not the same as the E 2 process seen in 89
Pseudopolvdesmus tallulanus. In several specimens the posterior margin of the collum had two large convex extensions.
DISTRIBUTION : The rar^e of Pseudopolvdesmus erasus covers the western slopes of the middle ^palachians from the southern tip of
Illinois, south to eastern Mississippi, southern Alabama, and western Georgia (Map 5). This species is allopatric with
Pseudopolvdesmus tallulanus. vhich occurs to the southeast at essentially higher elevations. Pseudopolvdesmus erasus appears to be limited to the Cumberland Plateau below 450 meters.
Pseudopolvdesmus canadensis (Neim^rt, 1844) NEW COMB.
(Figs. 55, 61, 70, 71, 72, 73, 76, 109, 114, 122-126;
Map 6 ; Tables 9-11)
Polvdesmus serratus of many authors, not Say, 1821. Polvdesmus canadensis Newport, 1844, Ann. Mag. Nat. Hist., 13:265 (type locality: CANADA; Ontario, Hudson's Bay, Albany River; type: BMNH). [hereby designated as holotype] Polvdesmus branneri Bollman, 1887, Proc. U. S. Nat. Mus., 11:620 (type locality: Tennessee, Jefferson Co., Jefferson City, Mossy Creek; type: USNM). NEW SYNONYMY. Polvdesmus nitidus Bollman, 1887, Ent. Amer. 3(1) :45 (type locality: Florida, Escambia Co., Pensacola; type: USNM). NEW SYNONYMY. Polvdesmus echinoaon Chamberlin, 1942, Bull. Univ. Utah, biol. ser., 6 (8 ): 10, fig. 33 (type locality: Pennsylvania, Luzerne Co., Shawanese (Harvey's Lake) ; type: USNM). NEW SYNONYMY. Polvdesmus conlatus Chamberlin, 1943, Bull. Univ. Utah, biol. ser., 8(2):18, fig. 5 (type locality: Tennessee, Sevier Co., Gatlihburg; type: USNM). NEW SYNONYMY. Dixidesmus svlvicolens Chamberlin, 1943, Bull. Univ. Utah, biol. ser., 8(2):20, figs. 37-38 (type locality: Georgia, Screven (not Briar) Co., 7mi. N. Sylvania; type: USNM). NEW SYNONYMY. Dixidesmus christianus Chamberlin, 1946, Proc. Biol. Soc. 90
Washijigton, 59:140, pl. XII, fig. 4 (type locality: Mississippi, Harrison Co., Pass Christian; type: USNM). NEW SYNONYMS. Dixidesmus catslcillus Chamberlin, 1947, Proc. Acad. Nat. Sci. Philadelphia, 99:24, fig. 2 (type locality: New York, Greene Co., Catskill; type: MSP). NEW SYNONYMY. Dixidesmus phanus Chamberlin, 1951, Great Basin Nat., 11(1-2):24, fig. 1 (type locality: Florida, Suwannee River; type: USNM). NEW SYNONYMY. Dixidesmus aausodicrorhachus Johnson, 1954, Chicago Acad. Sci. Nat. Hist. Misc., No. 137:1, figs. la-d (type locality; Michigan, Mackinaw Co., W. side of Grant Lake on Old US Rt. 2; type: USNM). NEW SYNONYMY. [Hereby designated as neoallotype of P. canadensis!. Dixidesmus branneri. Chamberlin and Hoffman, 1958, U. S. Nat. Mus. Bull. 212:65. Pseudopolvdesmus branneri. Hoffman, 1974, Proc. Biol. Soc. Washington 87(31):347.
DIAGNOSIS: Ifedium to large (ll-29mm). Metaterga vary from dark brown to black with red, pink, or brown paranota. Dorsal sculpture li^t. Dorsal and paranoial setae absent. Antennae long. Gonocoxae monolobed. GonopxDdal process M^ small, on edge, shark-fin shaped; M 2 medium, on edge across from E 2 + 3 ; M 3 medium, caudomedial; M 4 small with proximal setal patch; E^ long, thin, caudomedial; E 2 + 3 large, stalked, forked; E 2 prong large, extending proximad along side of tibiotarsus; E 3 prong large, extending away from tibiotarsus ; E 3 usually smaller than E 2 ; E 4 small, cyphopod projecting caudally, not flattened, cyphoped plate with straight distal edge, not sloping.
DESCRIPTION: (Based on adult male and female from Tennessee,
Campbell Co., on US Rt. 75, 19.7mi. S. of Jellico, VII-14-1982, C.
P. Withrow, (CPWC)). Adult male, 21.5mm long, maximum width (7th segment), 4.3mm, W/L 18.3%, and WL 84.9mm^. Segmental widths across paranota as follows: 91
Collum 2 . 6 m m 11-13 4.0 ram
2 3.2 m m 14 3.8 mm
3 3.4 m m 15 3.6 mm
4 3.5 ram 16 3.5 m m
5 3.9 m m 17 3.0 ram
6 4.1 m m 18 2.3 ram
7 4.3 m m 19 1.4 mm
8 - 1 0 4.2 ram 2 0 0.4 mm
Color in life, dark brown with red paranota.
Head typical for genus. Head width 2.08im, interantennal distance 0.58itim, 27.8% of total head width. Antennae long (4.06mm),
18.8% of total body length. Antennomeric lengths (% of total
antennal length): 1st antennomere, 0.35mm (8.9%); 2nd, 0.38mm
(9.7%); 3rd, 0.95mm (24.2%); 4th, 0.67mm (17.0%); 5th, 0.73mm
(18.6%); 6 th, 0.57mm (14.5%); 7th, 0.28mm (7.1%).
Dorsal sculpture li<ÿït. Collum moderate, subquadrate, slightly rounded along posterior edge. Anterior paranota rounded anteriorly and sli^tly cejhalad, becoming straight on posterior segments, sli^tdy sloping medially, lateral edge curved, posterior comer projecting caudalad only slightly. lateral boss of paranotum elonga-te, anteriomedial bosses well separated (Fig. 76). lateral denticles small, barely visible, distance from anteriolateral corner
(% of total length of 8 th segment): 1st denticle, 0.10mm (7.7%);
2nd, 0.28mm (21.5%); 3rd, 0.60mm (46.2%); 4th, 0.90mm (69.2%). 92
Posteriomedial teeth distinct on segments 17-19. Ventrolateral
surface of paranota rou^i and irregular.
Stema cruciately impressed, each quadrant heavily setose.
Sternum 2 sharply indented anteriorly, smoothly indented
posteriorly, with wide r o u ^ border. Sternum 3 with small medial
swellings, not as r o u ^ medially, penes normal, one posteriomedial
seta and two anteriomedial setae. Stema 4 and 5 similar, with many
long setae. Sternum 6 anterior lobes with short and long setae; posteriorly, goncpodal groove wide with many slender lateral setae, posterior sternal lobes sli^tly displaced anteriorly, lor^itudinal groove inverted V-shaped, deep, apices extending to coxae. Posterior
coxae of 7th segment projecting only s l i ^ t l y (Fig. 61). leg shape typical and long. Length of ri^t posterior leg of 8 th segment
4.16mm, 19.3% of total body length. length of podomeres (% of total leg length); coxae, 0.37mm (9.9%); trochanter, 0.80mm (19.2%); prefemur, 0.84mm (20.2%); femur, 0.45mm (10.8%); tibia, 0.52ram
(12.5%); tarsus, 0.95mm (22.8%); claw (Fig. 55), 0.23mm (5.5%).
Sphaerotrichomes typical.
Epiproct slightly downtumed. Hypoproct and paraprocts normal.
Gonopod aperture subelliptical (0.98mm long, 1.70mm wide) with
I/W ratio of 57.6%. Gonocoxae large (0.84mm high, l.lOmm long), posterioventral edge monolc±)ed, smooth, and subguadrate. Telopodite
(Fig. 70-73) large (2.50mm long). Prefemur normal. Femur thin with femoral ledge rounded. Tibiotarsus typically shaped, small
(0.02mm), blunt, on edge proximal to endomerite, 0.88mm (35.2%) from tip; M 2 medium (0.08mm), shaip, 0.44mm (17.7%) from tip; M 3 medium 93
(0.08mm), sharp, 0.36mm (13.2%) from tip; small (0.03mm), with
proximal setal patch, 0.15mm (6.0%) from tip; large (0.13mm),
blunt, 0.68mm (27.7%) from tip; E 2 + 3 bifurcate, stalk 0.61mm (24.4%)
from tip, prong E 2 large (0.28mm), hooked, extending proximally on
tibiotarsus; prong E 3 large (0.17mm), 0.87mm (34.8%) from tip, beside endomerite; E 4 small (0.03mm) and hooked, 0.09mm (3.6%) from tip. Tip sli^tly bent dorsally. Telopodite proximally curved, bent distally. Seminal groove t%)ical. Tibiotarsus thickness measured at
femoral ledge, perpendicular to body axis, 0.03mm, and parallel to
axis, 0.02mm. Endomerite sharp and thin, 0.78mm (31.2%) from tip,
extending 0.28mm on telopodite, projecting 0.15mm h i ^ and 0.24mm
long.
Female 21.7mm long, maximum width 3.63mm, W/L 16.7%, ard WL
78.7mm^. Segmental widths across paranota as follows:
Collum 2 . 6 m m 14-15 3.5 mm
2 3.0 m m 16 3.3 m m
3 3.2 m m 17 3.0 mm
4 3.3 mm 18 2.4mm
5 3.4 mm 19 1.5mm
6—7 3.5 m m 2 0 0.4 ram
8-13 3.6 ram
Color and structure similar to male. Stema and legs unmodified. Syncoxostemum flat posteriorly and slightly curved 94
posteriolaterally to protect cyphopods. Sternum III with raised
anterior margin.
Cyphopods (Figs. 109, 114) large and elongate (1.18mm long,
0.61mm h i ^ ) . Valves with numerous small macrosetae, posterior half with long setae laterally. Cyphopod plate large, edge straight, not
sloping.
VARIATION: Based on 1000 specimens, Pseudopolvdesmus canadensis appears to be the most variable species of the genus, thereby ej^laining the proliferation of specific names. The variability is probably correlated with the wide geographical range and hi(ÿi telopoditic coitplexity.
Based on 162 (100 males, 62 females) specimens measured, the mean boc^ measurements for males and females, respectively, were: length 22.79mm, 21.15mm; width 4.03mm, 3.74mm; W/L 17.76%, 17.60%;
IW 93.77mm^, BO.SOram^. The ranges and standard deviations can be seen on Tables 9, 10, and Figures 122-125. The length, width, W/L, and WL were higher in the males than females.
Based on ten ^>ecimens, the gonopodal processes varied little in their relative location on the tibiotarsus (Fig. 126). The average positions of the processes relative to telopodite length are: 52.8%, M 2 30.5%, M 3 13.4%, M 4 4.0%, 37.8%, E 2 + 3 23.7%, E 4
4.0%. For actual ranges see Table 11. Specimens from the southern part of the range tended to be smaller, with a more ccmpact telopodite. Process E 2 + 3 showed variation in the relative lengths and direction of the two components. Prong E2 tended to be longer in 95 the northern and southern parts of the range vhile both were equal in Michigan. The tibiotarsus was more robust and the red paranota seem to be most consistent in the i^^palachian Mountains. In some cases, small secondary spines were found associated with normal spines, althou^ no geographical continuity was observed. The tip
Tflhich is usually strai^t, may either be sharply hooked or slightly curved. Terminal setae, previously used as a diagnostic characteristic, primarily by Chamberlin and Causey, vary greatly, but most variation was due to breakage.
DISTRIBUTION : The range of Pseudopolvdesmus canadensis covers most of eastern North America from Hudson Bay, Canada, south into northern Alabama along the i^spalachian Mountains with scattered localities in northern Florida and southern Mississippi (Map 6). The species is most common in colder microclimates such as the higher elevations of the ^palachian Mountains. Discontinuous populations of Pseudopolvdesmus canadensis can be seen; 1) New England and southeast Canada, 2) J^jpalachian Mountains, 3) Michigan, and 4) the extreme southern populations. Little collecting has been recorded in
Pennsylvania so this gap is probably due to inadequate data. This is probably also true across southern Canada thereby linking the
Michigan and eastern populations. However there does appear to be a well-defined break between the Michigan and i^palachian populations.
Personal collecting in northern Ohio failed to turn iç» any canadensis specimens. Since canadensis appears to reside in cooler habitats, the southern populations from central Florida and southern 96
Mississippi are atypical. These may represent relic populations from the last period of glaciation. I esqject these populations are truly disjunct from the remaining range.
A specimen of canadensis has a Hawaii label and is either introduced or mislabelled. It would be interesting to know if the species has become established, because of the depaxçjerate native milliped fauna of the islands.
NOTES: Examination of Newport's single dried female holotype of Polvdesmus canadensis shcAB it to be distinct from
Pseudopolvdesmus serratus with vhich it had long been synonymized. I hereby designate the male type of crausodicrorhachus as the neo allotype of canadensis. due to it close proximity to the holotype locality and the need for a male type. The synonyms of canadensis are discussed below. As is the case with most Chamberlin species, all descriptions were based on 1 or 2 paragraphs of text.
Polvdesmus branneri Bollman, holotype examined. In the original description, Bollman states that the gonopods were different from serratus. although he did not esqplain how, and that they would be drawn in a later paper vhich was never published.
Polvdesmus nitidus Bollman, holotype examined, is hereby synonymized with canadensis.
Polvdesmus echinooon Chamberlin, holotype examined. The species was based on a specimen from Pennsylvania. In his gonopod drawing, M 2 was s l i ^ t l y more proximal than normal and the E 2 + 3 97 stalk has a small tertiary point between E 2 and E 3 . These conditions have been observed throu^out the range of canadensis.
Polvdesmus conlatus Chamberlin, holotype examined. The species had previously been synonymized with branneri by loomis and Hoffman,
1948. In the gonopod description, E4 was omitted.
Dixidesmus svlvicolens Chamberlin, holotype examined. The gonopods were misdrawn by placing a small secondary process adjacent to M 3 and omitting E4 .
Dixidesmus christianus Chamberlin, holotype examined. The species had been previously been synonymized (Loomis and Hoffman,
1948) with branneri. In the description, the gonopod figure was misdrawn with all the gonopodal processes slightly displaced distally.
Dixidesmus catskillus Chamberlin, holotype examined. The gonopod was misdrawn with M 2 missing and the tibiotarsus bent caudally.
Dixidesmus ohanus Chamberlin, holotype examined. The gonopod was misdrawn with M^ absent and either M 2 or M 3 absent.
I%x3n examination of the holotype, Dixidesmus cfausodicrorhachus
Johnson, was also determined to be a synonym of canadensis. the only difference being that E^ was Sorter than normal. In Johnson's description, E^ was short, E4 was placed next to E2 +3 , and E2 was short or at least equal to E 3 . The first two conditions were observed in the paratype, but not in the holotype. apparently the gonopods of the holotype were not removed and examined. The short length of E2 has been seen in other specimens throughout the range. 98
An effort was made to find and obtain part of Johnson's apparently large Michigan collection but neither he nor his collection could be found.
The species names, branneri. nitidus. echinoaon. svlvicolens. catskillus. phanus. and ouasodicrorhachus are here synonymized for the first time.
Packard (1886) described the "larval" form of canadensis althou^ it may be serratus due to similarities seen in juvenile forms.
Pseudopolvdesmus collinus Hoffman, 1974
(Figs. 85, 89, 93, 110, 115, 122-126;
Map 7; Tables 9-11)
Polvdesmus moniliaris Williams and Hefner, 1928, Bull. Ohio Biol. Surv. 4(3); 103, not Koch, 1847:135 (see notes). NEW SYNONYMY. Pseudopolvdesmus collinus Hoffman, 1974, Proc. Biol. Soc. Washington 87(31):346, figs. 1-2, (type locality: Virginia, Patrick Co., Pinnacles of Dan, circa 4mi. SW. Vesta; type: USNM).
DIAGNOSIS: Medium ( 12.8-25.3mm). Metaterga and paranota brown.
Dorsal and paranotal setae very small. Dorsal sculpture moderate.
Antennae long. Gonocoxae trilobed. Gonopodal process M^ small, on edge, close to endomerite; Mg medium; M 4 medium with small setal pad; very small to large, distal to endomerite; Eg medium; Eg medium; E4 small, cyphopod elongate, similar to canadensis.
DESCRIPTION: (Based on adult male and female from West
Virginia, Mason Co., Mud Run, 0.75mi. off State Route 2, IV-27-1977, 99
C. P. Withrow, (CEWC)). Adult male 20.7irati long, maximum width (7th segment), 3.6mm, W/L 17.6%, and WL 75.7ram^. Segmentai widths across paranota as follows:
Collum 2 . 1 mm 12-13 3.1 mm
2 2 . 6 mm 14 3.0 mm
3 2 . 8 mm 15 2.9 mm
4 2.9 mm 16 2.7 mm
5 3.1 mm 17 2.4 mm
6 3.3 ram 18 1 . 8 mm
7 3.6 mm 19 1 . 1 mm
8 3.5 mm 2 0 0 . 6 mm
9-11 3.2 mm
Color in life, brown.
Head typical for genus. Head width 2.2mm, interantennal distance 0.42mm, 19.1% of total head width. Antennae long (3.66mm),
17.7% of total body length. Antennomeric lengths (% of total antennal length): 1st antennomere, 0.19mm (5.2%); 2nd, 0.46mm
(12.6%); 3rd, 0.87mm (23.8%); 4th, 0.60mm (16.4%); 5th, 0.65mm
(17.8%); 6 th, 0.64mm (17.5%); 7th, 0.25mm (6 .8 %).
Dorsal sculpture weak, similar to canadensis. Collum moderate, gradually sloping posteriorly, posterior margin rounded. Paranota subquadrate with anterior edge perpendicular to body. Lateral denticles distinct, located from anteriolateral c o m e r (% of total length of paranotum, 8 th segment): 1 st denticle, 0 .1 0 mm (8 .8 %); 2 nd, 100
0.32mm (28.1%); 3rd, O.SBinm (50.9%); and 4th, 0.80mm (70.2%).
Posteriomedial teeth located on segments 15-19. Ventrolateral surface of paranota r o u ^ and irregular.
S t e m a cruciately impressed, each quadrant heavily setose.
Sternum 2 and 3 typical. Posterior sternum of 6 th segment deep, inverted U-shaped, not cephalad; coxae produced ventrally. Posterior sternal lobes of 7th segment only slightly elongate. leg shape normal and long. length of right posterior leg of 8 th segment
3.70mm, 17.8% of total body length. length of podomeres (% of total leg length); coxae 0.40mm (10.8%); trochanter 0.67mm (18.1%); prefemur 0.63mm (17.0%); femur 0.33mm (8.9%); tibia 0.42mm (11.4%); tarsus 1.05mm (28.4%); claw 0.20mm (5.4%).
Epiproct sli^tly downtumed. Paraprocts and hypoproct normal.
Gonopod aperture subelliptical (0.75mm long, 1.44mm wide) with
I/W ratio of 52.1%. Gonocoxae moderate, (0.62mm high, 1.17mm long), triangular, sli^tly trilobed. Telopodite (Figs. 85, 89, 93) large
(1.75mm long). Prefemur normal with small swelling situated 1.35mm from distal tip. Femur thin with straight femoral ledge. Tibiotarsus typically shaped, small (0.04mm), 0.99mm (56.6%) from tip; M 2 medium (0.06mm), 0.50mm (28.8%) from tip; M 4 medium (0.07mm), curved anteriorly similar to canadensis. 0.17mm (9.7%) from tip; medium
(0.06mm), 0.67mm (38.3%) from tip; E2 + 3 on combined stalk, 0.40mm
(22.9%) from tip; E2 medium (0.08mm), stubby; E3 medium (0.08mm), stubby; E4 small (0.02mm), 0.12mm (6.9%) from tip. Tip slightly bent. Seminal groove typical. Telopodite with distal bend.
Tibiotarsus thickness measured at femoral ledge, perpendicular to 101 boc^ axis, 0.27im and parallel to axis, 0.15mm. Seminal groove
rounded. Endomerite moderately long and rounded distally, 0.72mm
(41.1%) from tip, extending 0.26mm on telopodite, projecting 0.13mm h i ^ and 0.23mm long.
Female 22.1mm long, maximum width 3.7mm, W/L 16.8%, and WL
81.6mm^. Segmental widths across paranota as follows:
Collum 2 . 6 mm 14 3.5 mm
2 3.1 mm 15 3.4 mm
3 3.2 mm 16 3.1 mm
4 3.3 mm 17 2.7 mm
5—6 3.6 mm 18 2 . 2 mm
7-12 3.7 mm 19 1.3 mm
13 3.6 mm 2 0 0 . 6 mm
Color and structure similar to male, s t e m a and legs unmodified. Syncoxostemum wide medially, ventrolateral shelf small, laterally straight, rouÿi anteriorly. Sternum III smooth but lightly punctate, with small round median lobe.
cyphopods (Figs. 110, 115) elongate (0.90mm long, 0.42mm h i ^ ) . Valves not as extended as canadensis, lateral setae moderate.
Operculum small with large anterior curving setae. Cyphopod plate similar to canadensis.
VARIATION: Based on 40 (22 males, 18 females) specimens measured, the means of the boc^ measurements for males and females, 102 respectively, were: length 19.73mm, 19.43mm; width 3.92mm, 3.60mm;
W/L 16.95%, 17.15%; and IW 78.45mm^, 6 8 .75mm^. The ranges and standard deviations can be seen on Tables 9, 10, and Figures 122-
125.
Based on ten specimens, the gonopodal processes varied little in their relative position on the tibiotarsus (Fig. 126). The average positions of the processes relative to telopodite length are: 57.9%, M 2 26.6%, M 4 5.6%, 40.4%, E2 + 3 22.0%, E 4 6.3%. For actual ranges see Table 11. Process Ej variation in size and shape was most notable, ranging from a small swelling to a long slender process similar to canadensis.
Several specimens of collinus possess dorsal setae. The setae were very short conpared to those species that normally have dorsal setae.
DISTRIBUTION: The range of Pseudopolvdesmus collinus covers eastcentral North America, from the Atlantic coast of Maryland and northern South Carolina to western Ohio and eastern Tennessee (Map
7). The species is uncommon and sporadically found. The habitat is variable. Specimens have been collected from Ohio River mud-clay floo(%)lains, above 3000 ft in the Appalachians, and the sanc^ soils along the Atlantic coast.
NOTES: Chamberlin and Hoffman (1958) state that even though the name, moniliaris has been used by several authors in reporting northeastern polydesmids, the exact identity, because of the absence 103 of specimens, has never been conclusively settled. They also state that the specimens were probably the introduced species Polvdesmus inconstans or some other synanthrqpic species. This, however, does not appear to be the case with Williams and Hefner's usage of the name. Even though the specimens were missing from the Hefner collection, based on their description and illustration I believe their P. moniliaris is collinus. This argument is strengthened by a collection of collinus from westcentral Ohio, very close to the area in vhich Williams and Hefiner worked. There are still other records of moniliaris in the literature vhich remain unidentified.
The Grotç) Serratus consists of four species, serratus. paludicola. caddo. and minor. The characteristics of the grovç) are; posterior sternum of 7th segment with long sternal lobes, absent, gonocoxae multiple, and cyphopod sliiÿitly elongate, with plate ccfipressed laterally. The range of the grou^ is middle eastern North
America, extending into the lower Mississippi River Valley and eastern Texas.
Pseudopolvdesmus serratus (Say, 1821)
(Figs. 12, 44, 46, 48, 57, 59, 63, 6 6 , 6 8 , 65, 78,
94, 98, 102, 122-126; Map 8 ; Tables 9-11)
Polvdesmus serratus Say, 1821, J. Acad. Nat. Sci. Philadelphia, 104
2:106 (type locality: ’’Eastern ^ o r e of Virginia"; type: vinknawn, presumed lost; Neotype designated below). Polvdesmus pensvlvanicus [sic] Kbcii, 1847, jji Krit. Rev. Insect. Deutsdilands, 3:133.— 1863, Die Myriapoden, 1:18, pi. 69, fig. 142 (type locality: Pennsylvania; type: unknown, presumed lost; Neotype designated below). NEW SYNONYMY. Polvdesmus glaucescens Koch, 1847, in Krit. Rev. Insect. Deutsdilands, 3:133.— 1863, Die l^riapoden, 1:59, pi. 26, fig. 51 (type locality: North America; type: unknown, presumed lost; Neotype designated below). NEW SYNONYMY. Polvdesmus scopus Chamberlin, 1942, Canadian Ent., 74:16, fig. 1 (type locality: Iowa, Boone Co., 6 miles south of Boone; type: USNM). NEW SYNONYMY. Polvdesmus planicolens Chamberlin, 1942, Canadian Ent., 74:16, fig. 2 (type locality: Iowa, Story Co., Ames; type: USNM). NEW SYNONYMY. Pseudopolvdesmus serratzis, Chamberlin and Hoffman, 1958, U. S. Nat. Mus. Bull. No. 212:71.
DIAGNOSIS: Medium tuD large (13-32mm). Metatzerga brown, paranota sometimes red. Dorsal sculpture moderate. Dorsal and paranotal setae absent. Few sternal setae. Antennae moderate and filiform. Gonocoxae bilobed. Gonopodal process M^ small to medium, at base of endomerite, M 2 small, M 4 small, E 2 medium, connected to
M 2 by inner ridge, E4 medium, cyphopod elongate, raised marginally with expanded anterior surface flat, cyphopod plate small, sloping sli^tiLy medially.
DESCRIPTION: (Based on adult male and female from West
Virginia, Cabell Co., SR. 10, Imi. S. of jet. Alternate Stete Route
10, V-20-1983, C. P. Wi-fchrow, (CPWC)). Adult male, 23.3mm long, maximum width (8 th segment), 3.9mm, W/L 16.7%, and WL 90.40mm^.
Segmentel widths across paranota as follows: 105
Collum 2.5 mm 13-14 3.7 mm
2 3.0 mm 15 3.6 mm
3 3.3 mm 16 3.4 mm
4 3.5 mm 17 3.0 mm
5 3.6 mm 18 2.4 mm
6 3.8 ram 19 1.5 mm
7-8 3.9 mm 2 0 0.4 mm
9-12 3.8 mm
Color in life, brown.
Head typical for genus. Head width 2.5inm, interantennal distance 0.54inm, 21.6% of total head width. Antennae moderately robust (Fig. 57) long (4.89mm), extending to posterior edge of 4th segment, 18.9% of total bo<^ length. Antennomeric lengths (% of total antenneil lergth) : 1st antennomere 0.22mm (4.5%); 2nd 0.77mm
(15.8%); 3rd 1.19mm (24.4%); 4th 0.85mm (17.5%) ; 5th 0.94mm (19.3%);
6 th 0.64mm (13.1%); 7th 0.26mm (5.3%).
Dorsal sculpture moderate. lateral and medial bosses of paranotum small, medial boss almost round (Fig. 12). Anterior bosses of dorsum well separated. Collum large with acute posteriolateral angle, anteriomedial bulge. Anterior terga projecting laterally
(Fig. 44). Dorsal and paranotal setae absent. Paranota with straight anterior edge, posterior c o m e r directed caudally, lateral edge curved. Lateral denticles distinct, located from anteriolateral comer (% of total length of paranotum, 8 th segment) : 1 st denticle,
0.08mm (6.7%); 2nd, 0.08mm (13.4%); 3rd, 0.48mm (38.3%); and 4th, 106
0.83im (66.0%). Postericmedial teeth distinct on segments 14-19.
Ventrolateral surface of paranota r o u ^ and irregular. Laterally diploposegments as in figure 46.
Sterna cruciately inpressed, each quadrant moderately setose
(Fig. 59). Sternum 2 very irregular, short, anterior and posterior plate fields flat, small, and regular, with medial ridge irregularly asperate. Sternum 3 posteriomedially between large penes, sli^tly depressed ventrally, anterior edge pointed slightly with lateral and posterior margins raised, surface depressed, smooth, with about 1 2 long, thin setae; interpenial distance small, with a pair of longitudinal parallel ridges on sternum. Sterna 4 and 5 large, with many long, thin setae, however gap between sternum lobes shallow and steep. Lobes of sternum 6 blunt distally, each with one medial and one lateral pair of setae and with two pairs of posterior setae between them. Sternal setae ^ o r t and stubby with a small patch of similar setae posteriolaterally. Medial longitudinal depression on posterior of sternum 6 narrow and deep. Coxae covered with short, stubby setae and single, large, thin, ventrad seta. Posterior sternum 7 with long, thin, coxal lobes. Legs long (Fig. 69). Length of r i ^ t posterior leg of 8 th segment 3.87mm, 16.1% of total body length. length of podomeres (% of total leg length) ; coxa 0.41mm
(10.7%) ; trochanter with large dorsal protuberance vhich is slightly wider distally, 0.64mm (16.5%); prefemur 0.79mm (20.5%); femur
0.37mm (9.5%); tibia 0.41mm (10.7%); tarsus 1.04mm (26.9%); claw
0.20mm (5.2%). Sphaerotrichomes normal. Cross section as in figure
48. 107
Epiproct strai^t (Fig. 81). Hypcproc± trilobed, medial lobe small with single lateral pair of setae. Paraprocts typical.
Gonopod aperture subelliptical (1.6mm long, 1.05mm wide) with
I/W ratio of 65.6%. Gonocoxae large (0.80mm hi ^ , 1.13mm long), posterioventral edge bildbed, ventral lobe blunt, extending outward twice its width, sides parallel, smooth, rounded, and directed ventrally; dorsal lobe irregular, flattened with edges. Telopodite
(Figs. 94, 98, 102) large (1.56mm h i ^ ) . Prefemur normal. Femur thin with strai^t femoral ledge. Tibiotarsus typically shaped, process
small to medium (0.05mm), 0.60mm (57.0%) from tip, close to base of endomerite; Mg small (0.04mm), regular, 0.42mm (40.3%) from tip;
M 4 small (0.04mm), 0.18mm (17.0%) from tip; Eg medium (0.07mm), connected to Mg by small ridge, 0.38mm (36.4%) from tip; E4 medium
(0.06mm), 0.18mm (17.7%) from tip. Tip hooked dorsally. Seminal groove typical. Tibiotarsus with distal and proximal curves.
Tibiotarsal thickness measured at femoral ledge, perpendicular to body axis, 0.19mm, and parallel to axis, 0.21mm. Endomerite thin and sharp, 0.63mm (46.9%) from tip, extending 0.24mm on telopodite, projecting 0.11mm high and 0.15mm long, with acute tip.
Female 22.9mm long, maximum width 3.9mm, W/L 16.9%, and WL
8 8 .9mm^. Segmental widths across paranota as follows:
Collum 2.8 ram 11-13 3.9 mm
2 3.1 mm 14-15 3.8 mm
3 3.4 mm 16 3.5 mm
4 3.5 mm 17 3.3 mm 108
5 3.6inm 18 2 . 5 m m
6 3.7 ram 19 1.6 ram
7-10 3.8 ram 20 0.5 ram
Color and structure similar to male. Sterna and legs unmodified. Syncoxostemum quadrate distally, raesal surface very rou^, and posterior surface smooth with small grooves (Fig. 63).
Sternum III sli^tly enlarged with lateral projection moderate, ventral surface very r o u ^ and irregular (Fig. 6 6 ).
cyfiiqpods (Fig. 67) elongate (1.2ram long, 0.87ram high), valves with large number of short, raacrosetae along valvular lips. Lateral surface with numerous setae on ventral 1/3 grading to none dorsal, anterior edge rounded and flattened laterally with edge sli^tly thickened. Operculum small with several long setae. Dorsal plate
(Fig. 120) distinct and slightly sloped medially.
VARIATION: Based on approximately 4500 specimens,
Pseudopolvdesraus serratus. although wide-ranging, has s l i ^ t size variation with little or no geographic significance. Based on 500
(250 males, 250 females) specimens measured, the means of body measurements for males and females, respectively, were: length
23.86mm, 21.50mm; width 4.14mm, 3.72mm; W/L 17.31%, 17.90%; and LW
99.42mm^, 86.99mm^. The ranges and standard deviations can be seen on Tables 9, 10, and Figures 122-125. Males are only sli^tly longer and wider than females (23.9mm by 4.1mm to 21.5mm by 3.7mm, respectively). The females had a slightly hi^er W/L ratio. Because 109 males were longer and wider than females, dorsal area was also greater, 99.4mm^ to ST.Oram^. specimens from the i^ppalachians also appear sli^tly lai^ger. When serratus and other species, usually canadensis, were synpatric, serratus was consistently the larger.
Denticles tended to be smaller on extreme northern populations
(Ontario).
Coloration was usually brown, however paranota may be reddish in some specimens.
The hypoproct was usually bilobed althouiÿi a trilobed condition had been observed on several specimens.
The gonopods of Pseudopolvdesmus serratus showed extensive non-geographic variation in the size, shape, and location of the processes, the latter varying from spined, triangular, to finned.
The relative location of processes of the tibiotarsus varied little
(Fig. 126). The positions of the processes relative to telopodite length are: 56.1%; M 2 37.8%; M 4 13.0%; E3 37.1%; and E4 8.1%. For actual ranges and standard deviation see Table 11. The inner ridge between M 2 and E3 may be toothed.
The cyphopods varied very little, although the plates of southern populations (ex. Louisiana) tend to be more rounded.
DISTRIBUTION: The range of Pseudopolvdesmus serratus covers eastern North America from southern Quebec, Canada and Iowa, south to Georgia, the Gulf Coast and eastern Texas (Map 8 ). P. serratus is the most common species east of the Mississippi River and is only sporadically found farther west. Generally, it occurs in deciduous 110
hazdwoods in non-urbamzed environments. Even in many di.sturbed
areas, Pseudopolvdesmus serratus is common althou^ Polvdesmus
inconstans and Oxidus gracilis often appear to displace it. Mass
migrations have been recorded from Ohio (Ramsey, 1966).
Due to incdtplete records, the exact western extension of the
species in Canada is unknown.
NOIES: Polvdesmus canadensis, based on a female, has been
considered a synonym of serratus by all authors since Attems (1898),
but is instead the senior synonym of branneri.
Because the holotypes for Polvdesmus serratus. P.
pensvlvanicus. and P. alaucescens are unknown and presumed lost,
neotypes are here designated. Polvdesmus serratus: male neotype
designated from Virginia, Dinwiddle Co., Pocahontas S. P., (FSAC).
Polvdesmus pensvlvanicus : male neotype designated from Pennsylvania,
Bradford Co., 4.5 mi. S. of Mbnroeton on US Rt. 220, neotype and two male paratypes, VI-8-1958, R. Hiÿiton (RIHC). Polvdesmus glaucescens: male neotype designated from Ohio, Fairfield Co.,
Bamebey Center, IV-27-1984, C. P. Withrow, (USNM).
It appears that character displacement may be occurring. This deals with the case of red and brown tergite color between
Pseudopolvdesmus serratus and canadensis. Under normal conditions both have brown paranota, however vhen both species are synpatric, canadensis usually has red paranota and serratus has brown. However, many specimens of serratus collected from Ohio, vhere no canadensis occurs, have definite red paranota. I l l
Both Polvdesmus scoous Chamberlin, holotype examined, and P.
planicolens Chamberlin, holotype examined, were based on five-line
descriptions with poorly drawn gonopod figures. In describing P.
scopus. Chamberlin presented an inaccurately drawn figure of the
gonopods, vitiile giving no verbal description of them. Examination of the type shows that the gonopod figure was misleading in the
following ways: processes M^, M 2 , or E 3 , and E 4 were not shown but are present and the elongation of the femur of the telopodite is exaggerated. From the same article, similar problems occur with the
figure of P. planicolens: the inner ridge between M 2 and E 3 is spined, E4 is present but not shown, and the shape is exaggerated.
All other characters match serratus and fit within the limits of variation. Chamberlin described the two species from single males collected within 12 miles of each other. A trip was made to both type localities but no additional specimens were found.
Pseudopolvdesmus paludicola Hoffman, 1950
(Figs. 95, 99, 103, 125;
Map 9; Table 11)
Pseudopolvdesmus paludicolus Hoffman, 1950, Virginia J. Soi., 1:222, fig. 4 (type locality; Virginia, Princess Anne Co., Sand Bridge, 5mi. S. of Virginia Beach; l^pe: USNM).
DIAGNOSIS: Small (ll-13ram). Metaterga and paranota brown.
Dorsal sculpture well defined. Dorsal and paranotal setae present, lateral denticles very small or absent, caudolateral c o m e r produced 112 caudally. Antennae short. Gonocoxae monolobed. Gonopodal process medivrai, hooked, and at base of endomerite; M 2 medium and reduced to a low swelling; M 3 small and triangular; inner ridge with median elongated process extending proximally connecting E 4 and M 2 ; E 4 medium. No females have been recorded.
DESCRIPTION: (Based on adult male from Virginia, Princess Anne
CO., Norfolk, east side of Norfolk Reservior, V-18-1963, P. Hall,
(RLHC) ). Adult male 12.4mm long, 1.83mm maximum width (8th segment),
W/L ratio 14.8%, and WL 22.7mm^. Segmental widths across paranota as follows:
Collum 1.2 mm 15 1.6 mm
2 1.3 mm 16 1.5 mm
3-4 1.5 mm 17 1.4 mm
5-6 1,7 mm 18 1 . 1 mm
7-8 1.8 mm 19 0 . 8 mm
9-14 1.7 mm 20 0.3 mm
Color in life, brown, as described by Hoffman (1950), preserved specimens viiite.
Head typical for genus. Head width (1.0mm), interantennal distance 0.33mm, 33.3% of total head width. Antennae short (1.29mm),
10.4% of total body length. Antennomeric lengths (% of total antennal length): 1st antennomere, 0.11mm (13.7%); 2nd, 0.19mm 113
(15.1%); 3rd, 0.29rtim (22.7%); 4th, 0.21mm (16.2%); 5th, 0.22mm
(17.2%); 6 th, 0.19mm (15.1%); 7th, 0.07mm (5.2%).
Dorsal sculpture well defined, similar to serratus. collum small , without denticles, wider along posterior margin, straight anteriorly but rounded laterally. Lateral denticles very small or absent. Dorsal setae present. Paranota rounded anteriorly and pointed posteriolaterally with lateral edge curved. lateral surface ventral to paranota smooth, without denticles.
Sterna cruciately impressed. Each quadrant moderately setaceous. Sternum 3 with penes normal, with anteriomedial and postericmedial pairs of setae. Sterna 4-5 narrow, unmodified, with numerous long setae. Sternum 6 with large anterior lobes, posterior sternal lobes displaced slightly anteriolaterally ; medial V-shaped indentation wide and shallow. Sternum 7 with posterior sternal lobes long and thin. Sterna 8-20 typical with numerous short setae. legs short but typical. length of right posterior leg of 8 th segment,
(1.14mm), 9.1% of total body length. length of podomeres (% of total leg length) : coxae 0.18mm (1 2 .8 %) ; trochanter with dorsal extension larger proximally 0.28mm (19.6%); prefemur 0.22mm (15.7%); femur
0.14mm (9.8%); tibia 0.11mm (7.8%); tarsus 0.40mm (28.5%); claw
0.08mm (5.9%). Sphaerotrichomes on prefemur, femur, and tarsus.
Hypoproct slightly deflexed. Epiproct and paraprocts t^ical for genus.
Gonopod aperture subelliptical (0.71mm long, 1.04mm wide) with
I/W ratio of 69.2%. Gonocoxae large (0.19mm high, 0.47mm long), loig, and posterioventrally bilobed. Telopodite (Figs. 95, 99, 103) 114
small (0.98mm h i ^ ) . Preferaur small and hirsute. Femur thin with
femoral ledge straight. Tibiotarsus typical with process medium
(0.07mm), pointed, located 0.51mm (52.8%) from tip, at base of
endomerite; Mg medium (0.07mm), sharp, connected distally to small,
medial, secondary spine, 0.28mm (28.9%) from tip; M 3 small (0.02mm),
with barely visible medial ridge to secondary spine of E4 , 0.20mm
(20.1%) from tip; E 4 medium (0.08mm), 0.5mm (5.0%) from tip. Tip
thick with slight dorsal bend. Telopodite with slight curve. Seminal
groove typical. Tibiotarsus thick at femoral ledge, measured perpendicular to body axis, 0.08mm, parallel to axis, 0.06mm.
Endomerite thin and sharp, 0.57mm (58.6%) from tip, extending 0.20mm
on telopodite, projecting 0.03mm h i ^ and 0.13mm long.
No females have been positively identified.
VARIATION: Almost nothing can be said about variation because
only two males are known. The positions of the processes relative to telopodite length (Fig. 126) are: M^ 55.75%, Mg 30.55%, M 3 21.15%,
E4 5.55%. For actual ranges of the percentages see Table 11.
DISTRIBUTION: The range of Pseudopolvdesmus paludicola is
limited to two localities in southeastern Virginia, south of
Chesapeake Bay (Map 9). Shelley (1978) stated that paludicola can be ejpected in the Piedmont of the Carolinas. This was based on a personal statement by Hoffitian that a ^jecimen was collected from
Orangeburg, Orangeburg Co., South Carolina. However, this specimen could not be located in any collection. I expect paludicola to be 115
present throu^out the Mid-Atlantic Coastal Plain and possibly to
extend into the Piedmont.
NOTES: According to Hoffman (1950b) the holotype was found
under a board on the inner dunes, several hundred feet from the
ocean (generally an atypical locality for millipeds). The recorded
vegetation was g r ^ s and sedge, a few pines, and Mvrica thickets.
The specific name is Latin for swaitp-inhcjbitant; althou^ not
found in a swaitp it probability refers to the close proximity to the
Dismal Swattp. The name does not take the masculine gender suffix of
the genus. A female specimen from Greene Co., North Carolina (NMNH) was tentatively identified by Shelley to be paludicola. but was not
included here until males can be collected from the site.
Pseudopolvdesmus caddo Chamberlin, 1949
(Figs. 50, 77, 96, 100, 104, 116, 118,
122-126; Map 9; Tables 9-11)
Pseudopolvdesmus caddo Chamberlin, 1949, J. Washington Acad. Sci., 39:97, fig. 11 (type locality: Louisiana, Caddo Par., 5mi. NW. Shreveport; type: USNM). Pseudopolvdesmus bidens Loomis, 1959, J. Washington Acad. Sci., 49(5): 163, fig. 8 (type locality: Louisiana, leblanc Par., Kinder, US Rt. 190; type: USNM). NEW SYNONYMY.
DIAGNOSIS: Small (9-14ram). Metaterga and paranota light brown.
Dorsum sli^tly arched and well defined. Small dorsal and paranotal
setae present. Collum subquadrate. Paranota extending slightly anteriorly and squared laterally, flatter than anyother species. 116
Antennae short. Gonocoxal margin trilobed. Gonopodal process
small, at base of endomerite, M 2 small to medium, across from E2 , M 4
small, E2 small, elongate, usually with small distal tooth, E4
small. Cyphopod corrpact, edge with setae evenly spaced along length
of valve, lateral setae long, cyphopod plate elongate and loping medially.
DESCRIPTION: (Based on adult male and female from Louisiana,
Iberville Par., Maringouin Levee, 13.3mi. N. of Bayou Sorrel, X-30-
1965, R. E. Tandy, (ESAC)). Adult male, 12.6mm long, 2.1mm maximum width (8 th segment), maximum thickness 1.4mm, W/L 16.0%, T'JL 27.0mm^,
and thickness/width ratio of 70.0%. Segmental widths across paranota as follows:
Collum 1.5 mm 11-15 2 . 0 mm
2 1.7 mm 16 1.9 mm
3 1 . 8 mm 17 1 . 8 mm
4 1.9 mm 18 1.4 ram
5—6 2 . 0 mm 19 0.9 mm
7-10 2 . 1 mm 2 0 0.3 ram
Color in life, brown.
Head typical for genus except genae bulge only slightly. Head width 0.83mm, interantennal distance 0.25mm, 30.1% of total head width. Antennae short 1.87mm, 14.8% of total body length.
Antennomeric lengths (% of total antennal length) : 1st antennomere, 117
0.20mm (10.2%); 2nd, 0.31mm (15.8%); 3rd, 0.39mm (19.5%); 4th,
0.24mm (11.9%); 5th, 0.37mm (17.0%); 6 th, 0.35mm (17.9%); 7th,
0.15mm (7.7%).
Dorsal sculpture distinct. Anteriomedian bosses not well
separated, lateral boss extending entire length of each paranotum
(Fig. 77). Tergites directed sli^tly ventrolaterally. Collum small, bulging slightly posteriomedially, rounded anteriolaterally, and with 3 transverse rows of setae. Dorsal and paranotal setae small and filiform (Fig. 50). Paranota quadrate, extending slightly
cephalad, anterior edge straight. Lateral denticles distinct,
located from anteriolateral comer (% of total length of 8 th paranotum); 1st denticle, 0.17mm (22.7%); 2nd, 0.23mm (31.8%); 3rd,
0.40mm (54.5%); 4th, 0.60mm (81.8%). Postericmedial teeth not visible. Ventrolateral surface of paranota smooth.
Sterna cruciately impressed, each quadrant moderately setose.
Stema 2-5 typical for groi:p. Lobes of sternum 6 large, blunt, displaced anteriorly, with short sternal setae; deep, inverted, U- shaped longitudinal groove, with contour ridges. Posterior sternum of 7th segment long and thin, coxae with short, stubby setae, and each with a single, long, filiform macroseta. Legs short, prefemur slightly curved ventrally, femur and tibia with small, distal, dorsal process. leg shape typical but short. Length of right posterior leg of 8 th segment, 1.80mm, 14.3% of total body length.
Length of podomeres (% of total leg length) : coxae 0.28mm (15.7%) ; trochanter 0.38mm (21.0%); preferaur 0.32mm (17.9%); femur 0.20mm 118
(10.8%); tibia 0.20mm (11.2%); tarsus 0.47mm (26.0%); claw 0.14mm
(7.6%). ^haerotrichcfmes normal.
Hypoproct straiç^t. Ipiproct and paraprocts normal.
Gonopodal aperture subelliptical (0.67mm long, 0.99mm wide) with I/W ratio of 67.7%. Gonocoxae small (0.50mm high, 1.00mm long),
triangular. Telopodite (Figs. 96, 100, 104) small (0.94mm hi^).
Prefemur normal. Femur thin with femoral ledge slight and straight.
Tibiotarsus typical, process small (0.03mm), triangular, at base
of endomerite, 0.52mm (55.3%) from tip; M 2 small (0.03mm), hooked,
0.26mm (27.7%) from tip; M 4 small (0.03mm), triangular, 0.08mm
(8.5%) from tip; E2 small (0.02mm), long and thin, extending from
0.34-0.46mm (36.1-48.9%) from tip, E 4 small (0.02mm), triangular,
0.8.0mm from tip. Tip with small, dorsally directed tooth.
Telopodite evenly and gently curved. Seminal groove normal.
Telopodite width measured at femoral ledge, perpendicular to body axis, 0.14mm, and parallel to axis, 0.08mm. Endomerite sharp, base
0.50mm (0.53.2%) from tip, extending 0.20mm on telopodite, projecting 0.03mm high and 0.14mm long.
Female 13.2mm long, maximum width 1.8mm, W/L 16.3%, and WL
22.7mm^. Segmental widths across paranota as follows;
Collum 1.3 mm 17 1 . 6 mm
2 1 . 6 mm 18 1.3 mm
3 1.7 mm 19 0.9 mm
4-15 1 . 8 mm 2 0 0 . 2 mm
16 1.7 mm 119
Color and structure similar to male, with s t e m a and legs
unmodified. Syncoxostemum quadrate distally, straight laterally,
and very rou^i mesally and distally. Sternum III posterior edge
ridged.
cyphopods (Figs. 116, 118) small (0.61mm long, 0.42mm h i ^ ) ,
conpact. Valves with 14-15 overlapping macrosetae, dorsal edge
thickened, deeply marginated dorsally, and lateral setae extending
dorsally. Operculum normal, cyphopod plate elongate, sloping
medially.
VARIATION: Based on 28 (17M, IIF) specimens, the means of the
body measurements for males and females, respectively were: length
10.21mm, 9.75mm; width 1.42mm, 1.37mm; W/L 13.04%, 12.87%; and IW
15.22mm^, 14.37mm^. Ranges and standard deviations can be seen in
Tables 9, 10, and Figures 122-125. Females were only slightly
smaller than males. Thickness ranged from 0.84 to 1.02mm and the
thickness/width ratio ranged between 72.7 and 87,2%.
Based on ten specimens, the gonopodal processes varied little
in their relative position on the telopodite (Fig. 126). The positions of the processes relative to telopodite length are:
53.6%; Mg 29.6%; Mg 9.7%; and Eg 38.5%. For actual percentages see
Table 11.
DISTRIBUTION: The known range of Pseudopolvdesmus caddo is
along the Gulf Coast, from eastern Texas, through Louisiana.
Pseudopolvdesmus caddo appears to be associated with wet low areas 120 along the Lower Mississippi and Red Rivers, and along the Texas
Coastal Plain. Because of habitat similarities caddo probably extends into western Mississippi (Map 9).
NOTES: The holotype of Pseudopolvdesmus bidens Loomis, examined, was misdrawn. Loomis only showed M 2 and M 4 on the telopodite. Like Chamberlin, Loomis gave no written description of the gonopods.
Pseudopolvdesmus minor (Bollman, 1888)
(Figs. 62, 79, 97, 101, 105, 117, 119, 122-126;
Map 9; Tables 9-11)
Polvdesmus minor Bollman, 1888, Ent. Amer. 4:2 (type locality: Arkansas, Pulaski Co., Little Rock; type: USNM). Polvdesmus neoterus Chamberlin, 1942, Bull. Univ. Utah, biol. ser. 6 (8 ): 10, fig. 30 (type locality: Louisiana, Orleans Par., New Orleans; type: USNM). NEW SYNONYMY. Polvdesmus euthetus Chamberlin, 1942, Bull. Univ. Utah, biol. ser., 6 (8 ): 11, fig. 36 (type locality: Missouri, St. Louis Co., Buder Park, Imi. SE. of Valley Park; type: USNM). NEW SYNONYMY. Pseudopolvdesmus minor. Chamberlin and Hoffman, 1958, U. S. Nat. Mus. Bull. 212:70.
DIAGNOSIS: Small (9-14mm). Metiaterga and paranota light brown.
Dorsum slightly arched and well sculptured. Small dorsal and paranotal setae present. Antennae short. Gonocoxae bilobed, not as flat as Pseudopolvdesmus caddo. Gonopodal process M^ medium, separate from base of endomerite; M 2 medium, distal txo endomerite;
M 3 small, medial; E2 small, elongate, usually with distal process; 121
E 4 small, subterminal. Endomerite small and sharp. S t e m a 2 and 3 of
female with irregular ventral swellings. Cyphopods small, compact,
few small lateral setae present, flattened dorsally, with a dorsal
clump of macrosetae overlapping the valves, with small medially
sloping dorsal plate.
DESCRIPTION; (Based on adult male and female from Arkansas,
Desha Co., Mb(3iee, 1-7-1954, N. B. Causey, (ESAC)). Adult male
10.1mm long, maximum width (8 th segment) 1.5mm, thickness 1.2mm, W/L
16.8%, WL 14.9mm^, and thickness/width 80.0%. Segmental widths
across paranota as follows:
Collum 0 . 8 mm 15 1.3 ram
2 1 . 2 mm 16 1 . 2 mm
3-4 1.3 mm 17 1 . 0 mm
5-7 1.4 mm 18 0.9 ram
8-9 1.5 mm 19 0.7 mm
10-14 1.4 mm 2 0 0 . 2 mm
Color in life, licÿat brown.
Head typical for genus. Mandible with s l i ^ t lateral bulge.
Head wide 0.83mm, interantennal distance 0.25mm, 30.1% of total head width. Antennae short (1.22mm), 12.1% of total body length.
Antennomeric lengths (% of total antennal length): 1st antennomere,
0.13mm (10.6%); 2nd, 0.17mm (14.3%); 3rd, 0.26mm (21.2%); 4th, 122
0.16im (13.2%); 5th, 0.20mm (16.4%); 6 th, 0.20mm (16.4%); 7th,
0.10mm (7.9%).
Dorsal sculpture well defined. Collum moderate with single
lateral denticle, three transverse roi-js of dorsal setae, and
postericmedial edge indented. Tergites are directed ventrolaterally
sli^tly. Paranota quadrate, not extending cephalad, anterior edge
strai^t, posterior c o m e r pointed, lateral edge straiÿit.
Anteriomedial bosses not well separated, lateral boss extending
entire length of paranota. lateral denticles distinct, located from
anteriolateral comer (% of total length of 8 th paranotum) 1 st
denticle, 0.07mm (13.6%); 2nd, 0.15mm (30.5%); 3rd, 0.26mm (52.5%);
4th, 0.40mm (81.4%). Denticles usually with one or sometimes two
short, blunt, setae. Postericmedial teeth not distinct, only visible
on segments 18-19. Surface ventrolateral to paranota rough and
irregular.
S t e m a cruciately impressed, each quadrant heavily setose.
S t e m u m 2 with normal penes, with anteriomedial and postericmedial
pair of setae. S t e m u m 3 with 2 anteriomedial setae on small
swelling, and one postericmedial seta. S t e m a 4-6 typical, with
numerous long setae. Sternal lobes on 6 th segment largest. Posterior
stemal lobes of 6 th segment slightly displaced anteriorly, medial
indentation of stemum deep from coxa to coxa. Posterior stemal
lobes of 7th segment long and thin (Fig. 62). Contour ridges of groove longer than minor. Leg shape typical although short. length
of right posterior leg of 8 th segment, 1.43mm, 14.3% of total body
length. Length of podomeres (% of total length): coxae 0.21mm 123
(12.9%); trochanter 0.34mm (21.1%); preferaur 0.26ram (16.0%); femur
0.13ram (8.2%); tibia 0.19ram (11.4%); tarsus 0.40ram (24.3%); claw
O.lOrara (6.2%). Sphaerotrichomes normal.
Hypoproct strai^t. Epiproct and paraprocts normal.
Gonopod aperture subelliptical (0.62mm long, 0.80mm width), with I/W ratio of 70.7%. Gonocoxae small (0.47ram high, 0.50mm long),
similar to serratus. with large ventral and small dorsal lobes, ventral setae absent. Telopodite (Figs. 97, 101, 105) short
(0.96mm). Preferaur rounded and elongate. Femur thin with straight
femoral ledge. Tibiotarsus i^ically shaped, process medium
(0.06mm), located 0.54mm (56.3%) from tip, away from base of endomerite; M 2 medium (0.05mm), finned, and 0.38mm (39.6%) from tip;
M 3 small (0.03mm), located 0.17mm (17.7%) from tip; £ 3 small
(0.02mm), elongate, 0.34-0.28mm (35.5-29.2%) from tip; E4 small
(0.02mm), 0.07mm (7.3%) from tip. Tip hooked dorsally. Telopodite with distal and proximal curves. Seminal groove typical. Tibiotarsus thickness, measured at femoral ledge, perpendicular to body axis,
0.08mm and parallel to axis, O.llram. Endomerite thin, sharp, 0.43mm
(44.8%) from distal tip, extending 0.21mm on telopodite, projecting
0.02mm h i ^ and 0.14mm long.
Female 13.4mm long, maximum width 2.2mm, maximum thickness
0.18mm, W/L 16.4%, WL 29,5mm^, and thickness/width 83.5%. Segmental widths across paranota as follows:
Collum 1.5 mm 12-13 2.0 mm
2 1.7 mm 14 1.9 ram 124
3 1 . 8 mm 15 1 . 8 mm
4 1.9 mm 16 1.7 mm
5 2 . 0 ram 17 1.5 mm
6 2 . 1 mm 18 1 . 2 mm
7-9 2 . 2 mm 19 1 . 0 mm
1 0 - 1 1 2 . 1 mm 2 0 0.4 mm
Color and structure similar to male, however legs and s t e m a unmodified except for stema 2 and 3 with small irregular swellings
(Fig. 79). Syncoxostemum rounded distally, lateral edge proitMting caudally with posterior margin turned medially, and very irregular mesally. Stemum III slightly enlarged with lateral projection moderate, and ventral surface rough.
cyjtiopods (Figs. 117, 119) small (0.56mm long, 0.40mm high), large dorsally, and similar to caddo. Valves with many short macrosetae along margin. Dorsal plate ccmpressed laterally, not elongated, but sloping medially, cyphopods with several setae on posterior 1 / 2 grading to none dorsally, dorsal edge rounded, flattened laterally and sli^tly marginate.
VARIATION: Based on 31 (17M, 14F) specimens, the means of the body measurements for males and females, respectively, were:. length,
10.34mm, 10.78mm; width, 1.34mm, 1.34mm; W/L 12.95%, 12.45%; and IW,
13.92mm^, 14.59mm^. The ranges and standard deviations can be seen in Tables 9, 10, and Figures 122-125. No significant difference was 125
observed between males and females. Thickness ranged between 0.97
and 1.25mm. The thickness/width ratio ranged from 83.2 to 112.2%.
Based on ten specimens, the gonopodal processes varied little
in their relative position on the telopodite (Fig. 126). The positions of the processes relative to telopodite length are:
57.2%, M 2 37.8%, M 3 19.8%, E2 32.3%, and E 4 7.8%. These percentages were derived as described in the methods section. For actual ranges of the percentages see Table 11.
DISTRIBUTION: The range of Pseudopolvdesmus minor is centered in the middle Mississippi River drainage. Specimens were collected from northern Illinois, Missouri, and Arkansas, east through
Ifentucky and Tennessee (Map 9). The New Orleans locality was probably due to flood transport or mislabeling. All localities for minor appear to be associated with flood plains of major rivers, including the Illinois, Wasbash, Lower Missouri, and Lower Arkansas
River systems.
NOTES: The holotype of Polvdesmus euthetus Chamberlin, examined, was misdrawn in the original description. Chamberlin drew the M^ process too close to the endomerite.
The holotype of Polvdesmus neoterus Chamberlin, exanüned, was typical except that the apical tip of the gonopod was bent laterally, and the apical row of setae was absent.
In both descriptions, as is the case with many of Chamberlin's 126 works, the gonopods were not described verbally, and only poorly drawn figures were given.
Subfamily Epanerchodinae, NEW SUBFAMILY
(Figs. 3, 15; Map 1)
Epanerchodus Attems, 1901. Mitt. Mas. Hamburg 18:102. (subgenus of Polvdesmus). Epanerchodus Attems, 1914. Arch. Naturg. Abt. A 80(4) :157.
DIAOTOSIS: Small to large (6-29ram), one species smaller than
13mm. Body with 20 segments. I/W ratio 8.8-19.5%. IW between 4.3 and
59. 9mm^. Tergal width to he i ^ t 1.3. Body width to height 1.00.
Brown or depigmented. Head subglobose or slightly quadrate with microsetae. Mandible rounded laterally. Gnathochilarium quadrate.
Antennae long and either filiform or slightly clavate. length relationships of antenncmeres 7<1<2<6<5<4<3. Antennomeres 2-4 cylindrical, 5 & 6 slightly clavate, and 7 subconical, with typical dorsal knob and also small dorsoproximal knob. Interantennal distance large. Metazonite with small pits. Terga (Fig. 3) horizontal with polygonal bosses, not tuberculate. Setae may be associated with bosses. Anterior terga directed anteriad or laterad. collum semiovate or ovate and usually large, wider than head.
Paranota directed ventrally or horizontally, may be large or small.
Lateral edge usually dentate with dorsal filiform or clavate setae.
Peritremata thick. Poriferous arrangement normal. Ozopores small, marginal, opening dorsolaterally, and not elevated. Stema 127 unmodified. Vasa deferentia large. Hypoproct transversely oval and non-hirsute. Paraprocts typical with thin margins. Epiproct non- hirsute and usually sli^tly trilobed. Legs slender, long, 23% of body length and unmodified. Length relationship of podomeres
1<7=4<5<2=3=6. Claw long, thin, and tapered. Trochanter and preferaur with sphaerotrichomes.
Gonopods (Fig. 15) moderate, bent at right angle and not extending over anterior edge of 7th segment, aperture large, transversely oval, extending laterad beyond ends of coxal sockets of
7th pair of legs, lower edge of posterior side sliiÿîtly concave and rpper edge markedly extending forward to form a broad shelf. Stemum
7 broadly depressed between 8 th coxae. Seminal groove typical, extending subterminally on the tibiotarsus, usually with seminal vesicle, and terminating near the endomerite T^hich opens on surface, near an enlargement (=clivus of Attems, 1901) or proximal to it.
Cannula distinctly attached to preferaur under large ledge. Gonocoxae triangular, small, and hirsute. EYeferaur large, irregular, hirsute, and may have patches of elongate setae. Femur short, unbranched, with small proximal cingulum, and usually with large caudad process.
Clivus variable, from a large inflated sac to a small, hirsute mound. Tibiotarsus attached ventrally to gonocoxae, highly variable, bent 90° to the femur, and with caudally projecting processes.
Syncoxostemum thickened, not projecting. Stemum III may be modified. Cyphopods oval, without anterior plate. 128
CONTENT : This subfamily presently contains one large heteroraorphic genus, Epanerchodus. with 8 subgenera containing approximately 98 species. This genus can probably be divided into several genera.
The species were placed in the genus Polvdesmus prior to 1901, vdien Attems erected a new subgenus. In 1914, he raised it to generic level. In the early 1940's Verhoeff in a series of publications erected 6 subgenera. Two additional subgenera were later added by
Chamberlin and Wang (1953) and Miyosi (1954). Little work except for species description has been done on this taxon.
DISTRIBUTION: Palearctic (Map 1). Restricted to East Asia, primarily Japan, with several species in Siberia and Korea. Many species are known from lava caves around Mt. Fuji, these caves are known to be less than 1 0 , 0 0 0 years old, yet many contain unique species. CLADISTIC ANALYSIS
In the Phylogenetic and Classification Methods section of this study, some of ity cladistic ideas and techniques were outlined. What follows is an analysis of taxonomic relationships based on those ideas. The relationships are based on the sequences and patterns of branching, vhich in turn are based on shared apctmorphic character states. This analysis includes listing of monophylies, characters, character states, polarities, and analysis of phylogenetic trees.
CHARACTER STATES
This section presents the characters, their relative states, and the polarities that were used in formulating the phylogenetic hypotheses. This is basic for further study at family and superfamily levels. The arrangement of characters and character states is based on the clade level to #iich that character belongs.
129 130
The Si:ç)erfamily Polydesmoidea is a large, almost entirely
Holarctic taxon; dealing with it except in a generalized sense is beyond this stuc^. The characteristics of each of the five families, except for the Polydesmidae and its sister groiç), Doratodesmidae, have not been studied. Because this stuc^ deals with the North
American polydesmid fauna, vhich contains only two subfamilies, exotic taxa are not covered in depth.
Monophvlv of the Familv Doratodesmidae
There has been no in depth stuc^ of this small southeast Asian family. It is not the author's intention to rectify this problem in this stucfy. However, there was a need to obtain sorte understanding of this taxon beyond Hoffman's (1977) family description, in vhich he also suggested that Doratodesmidae is the closest relative to the
Polydesmidae. In Hoffman's 1979 classification, 9 genera, mostly monotypic were listed. The systematics of the family has two major problems: one is the lack of any previous revision and the other is the lack of specimens due to incomplete collecting or scarcity of iixlividuals.
Based on the few specimens I have examined, I agree with
Hoffman (1977) that Doratodesmidae is the sister group of the
Polydesmidae and that at least two subfamilies can be recognized.
The following list of apamorphies is probably not ccmplete. It is based on the several specimens examined (Appendix D) and the literature. The 5 autapomorphies for the Family Doratodesmidae are: 131
(Character 1: Paranota of 2nd Segment Flabellately Enlarged
In the plesiamorphic state, the paranota of an individual are essentially all the same relative size. The flabellate enlargement of the second paranotum is present in the doratodesmids, therefore autapomorphic (Fig. 1). This enlargement gives added protection to the head and legs vhen the milliped is curled. There is also a corresponding reduction in the collum.
Clmracter 2: Ability to Involute
Most millipeds lack the ability to curl into a ti^t ball and is considered the plesicmorphic condition. Those in several unrelated taxa, however do curl vp and are conpletely protected by dorsal arxî paranotal (character 1) modifications. In the Order
Polydesmida, this ability is shared by several families,
Doratodesmidae, Sphaeriodesmidae, cyrtodesmidae, and Oniscodesmidae.
This ability is not present in any other taxon of the Siçierfamily
Polydesmoidea. This condition is considered by me to be autapomorphic for the doratodesmids within the suç)erfamily and to have arisen independently in all four families. In each family different tergal modifications are ertployed.
Character 3: Gonocoxae with Ventral Macrosetae
The plesiamorphic condition is the absence of ventral macrosetae in the excavation. The presence of a patch of numerous macrosetae in the ventral excavation is autapomorphic in doratodesmids (Fig. 13). Although most polydesmids have a pair of 132 elongate setae on the ventrolateral edge, all lack setae within the ventral excavation.
Character 4: Sternal Width Narrow
This autapomorphy appears to be related to character 2. The doratodesmids have very narrow or contiguous sterna permitting tighter involution. In the polydesmids the sterna are generally very wide.
Character 5; Boc^ Length/Width Ratio High
In the doratodesmids the paranota are very wide, extending relatively far from the bo<^. For this reason the width is relatively large compared to total boc^ length. In the autapomorphic state, seen in the doratodesmids, this ratio ranges from 18 to 30%.
In the plesiamorphic state, seen in the remaining polydesmoids, this ratio ranges from 8 to 2 2 %.
MonoPhvlv of the Familv Polvdesmidae
The monophyly of the Family Polydesmidae is not definitively established; this problem can only be alleviated by a total ordinal revision. Althou^ some progress has been made in the last decade
(e. g., Hoffman, 1979), problems remain, especially how the trichopolydesmids, macrostemodesmids, and fuhrmannodesmids fit into the overall picture. In the past 20 years, all three taxa have been placed in the family at one time or another; and at the other 133 extxeane, Hoffitian (1979) placed each in a separate family within 2 other siçjerfamilies! Most of the problems are due to inadequate descriptions of taxa by past workers. In most cases the original and only generic description covers at most one or tvro paragraphs.
Hoffinan, i^hose works and studies are based primarily on gonopods, states that the apomorphic characters for Polydesmidae are development of a terminal chamber of the seminal groove, and the cordate or subcordate shape of the gonopod aperture, with the posterior margin projecting inward forming a transverse shelf. Only recently (e. g. Enÿioeff, 1981) has the studi'’ of milliped systematics progressed beyond the almost pragmatic use of gonopods to the utilization of other structures.
Presently I agree with Hoffman (1979) on the monophyly of the family Polydesmidae. An indepth study of the Polydesmoidea may uncover more apomorphic characters for the Polydesmidae. Characters vdiich are apomorphic in the family also include 18 and 64 viiich are plesiamorphic in several taxa and will be discussed later. The family has the following 5 apomorphies (characters 6-10) :
Character 6 : Interantennal Distance Wide
This autapomorphy is characteristic of the family
Polydesmidae. The antennae are well separated, by approximately 25% of the total head width. In the doratodesmids and several other taxa in the superfamily the antennal bases are or almost are contiguous.
Character 7; Antennae Long 134
This autapomorphy is characterized by the elongation of the antennae relative to boc^ length. As in character 4, several families as well as the doratodesmids have short, very clavate antennae. There is shortening of the relative length in some of the smaller-sized taxa (ex. mastigonodesmines and Pseudooolvdesmus minor) but this appears to be secondarily derived.
Character 8 : Seminal Groove Subterminal
The opening of the seminal groove, located subterminally on the tibiotarsus, is apomorphic for the Polydesmidae. In the polydesmines, it is about midway ig the tibiotarsus near the endomerite. In the epanerchodines, it is near the clivus. The plesiamorphic condition, vhere the opening is terminal, is seen in many taxa of the superfamily. In several genera of doratodesmids the plesiamorphic condition is observed, however, in these cases I believed that the subterminal condition is independently derived.
Character 9: cyphopod Elongate
The elongation of the cyphopod is apomorphic for the polydesmids. The plesicmorphic condition of an ovate cyphopod is seen in the doratodesmids and several polydesmines and epanerchodines but the state in the latter two taxa is believed to be secondarily derived due to small body size.
Character 10: Metazonal Tubercles 135
The absence of metazonal tubercles is autapomorphic in the polydesmids. The metazonal surface is usually covered by small regular pits.
The following apomorphies are present for the subfamilies
Mastigonodesminae and Scytonotinae. The placement of these two subfamilies into a major separate clade was not ej^jected, due to distance between the two subfamilies. I es^ject this clade to represent relict taxa. This clade has the following 6 apomorphies
(characters 11-16);
Character 11; Mandible Round
The apomorphic state, vhere the mandible is rounded laterally contrasts to the plesiomorphic state, vhere it is quadrate. This character is easily observed in dorsal view. The plesiomorphic condition is seen in Doratodesmidae, the subfamily
Mastigonodesminae, and the genus Bidentogon in the subfamily
Scytonotinae. This condition in these latter cases was probably derived independently, but the strength of rounded mandible as an autapomorphy is diminished.
Character 12: Ozopore Marginal
The plesiomorphic condition is the location of the ozopore medial to the margin of the paranotum. The autapomorphic condition, seen in the Polydesmidae, has the ozopore located on the lateral or posterior itargin, in contact with the edge (Fig. 12). 136
Character 13: Ozopore Posteriorly Located
The posterior placement of the ozopore is apomorphic for several taxa, the subfamily Mastigonodesminae and in the subfamily
Scytonotinae, the genera Soeodesmus. Coronodesmus. and Idahodesmus.
The plesiomorphic condition is the placement of the ozopore laterally on the paranotum.
Character 14: Terga Trapezoidal
The plesiamorphic character is the normal rectangular shape with lateral paranota. However in two taxa, Mastigonodesminae (Fig.
2) and the genus Speodesmus (Fig. 4), the apomorphic state in viiich the paranota are equilaterally trapezoidal is present. The anterior edge is narrower transversely than the posterior edge.
Character 15: No Individuals Over 2 Oram
For the purpose of this study size was divided into three ranges; small, those individuals under lOram; medium, between 10-2Oram and large, over 2Oram. The absence of large individuals is considered apomorphic. Large boc^ size appears to be the plesiomorphic condition; however, a number of taxa have evolved various degrees of smallness. This is probably due to habitat constraints.
Character 16: Tibiotarsus Attachment Lateral
The apomorphic characteristic is the lateral origin of the tibiotarsus on the gonocoxae. It appears that this attachment is 137 less secure than the plesiomorphic or ventral condition. The apomorphic condition is present in the Scytonotinae and some elements of the Polydesminae.
Monophvlv of the Subfamily Mastigonodesminae
In most classifications, this taxon is placed at the familial level (e. g., Strasser, 1974). The latest examination, by Hoffman
(1979), placed it in the Family Polydesmidae. He speculated that this conplex of 3 genera deserved subfamilial designation. Specimens and literature examined suggest that Hofûnan was correct. The following 5 apomorphies (characters 17-21) are present for the subfamily Mastigonodesminae:
Character 17: Unpigmented
The apomorphic condition is the loss of boc^ pigmentation.
This state is seen in the subfamily Mastigonodesminae, Speodesmus.
Speorthus. Idahodesmus. and some species in the Epanerchodinae. This characteristic is common in troglcx^rtic ^)ecies and in various small-sized non-troglodytic taxa.
Character 18: leg Modifications Present
This apomorphic state includes various modifications of either the third pair of legs or of multiple legs in the males, #d.ch have probably been derived independently. The first type is present in the subfamily Mastigonodesminae, Bidentogon. and Coronodesmus where there is an enlargement of the tibial podomere (Fig. 53). The second 138
case, several species of Scvtonotxis. involves various knobs and projections on the tibial podomeres of leg pairs 5-11 (Fig. 54).
Character 19: legs with Elongate Setae
The apomorphic condition is the presence of a number of long, dorsodistal, sensory setae, on at least the tibial and tarsal podomeres. These are not present in the plesiomorphic condition. The
Mastigonodesminae and the genera Speorthus and Idahodesmus exhibit the apomorphic condition. These three taxa include the smallest
individuals studied and may be related to size. In Speodesmus the legs possess a large number of moderately long setae ^mhich may be a troglodytic modification.
Character 20: Paraprocts Hirsute
The presence of hirsute paraprocts is apomorphic. The plesiamorphic condition is the presence of only two pairs of setae.
The apomorphic condition vhich varies from 7 to 18 pairs, is present in four taxa, Mastigonodesminae, and the genera Bidentogon.
Speorthus. and Idahodesmus.
Character 21: Sternum III Modified
The modification of the third sternum in males is apomorphic in the Mastigonodesminae and the genus Idahodesmus (Fig. 39).
Althou^ similar in both taxa, these bifurcated Y-shaped projections may have arisen independently. 139
Monophvlv of the Subfamily Scytonotinae
The monophyly of the subfamily is well documented. The following 11 apomorphies (characters 22-32) are present in this subfamily:
Character 22: Syncoxostemal Hooks Present
The presence of lateral syncoxostemal hooks is autapomorphic for the Scytonotinae (Fig. 64). These small ventrocaudal hooks have no known function. The plesiamorphic condition is the absence of these hooks.
Character 23: Claws Long
The plesiamorphic condition is the presence of ^ o r t claws, less than 4% of total leg length. Taxa with the autapomorphic condition have longer claws, usually over 5.5% of leg length. The apomorphic condition is seen in the scytonotines, the polydesmines, and a few epanerchodines.
Character 24: Biogeography
Biogeographical information is usually used to substantiate the generated tree. Here a possible series was set up with Southeast
Asia the most plesiamorphic as represented by the Family
Doratodesmidae; the Holarctic region as intermediate, and the
Nearctic as apomorphic. The subfamilies Mastigonodesminae,
Polydesminae, and Epanerchodinae are intermediate, vhile the apomorphic cordition is ejdiibited by the scytonotines. 140
Character 25: Only 19 Segments Present
The plesiomorphic condition is vdiere 20-segmented or 19- and
20-segmented species are present. The apomorphic state of only 19- segnfânted individuals is present in two subfamilies, the
Mastigonodesminae and the Scytonotinae. Some individuals in the
Polydesminae have 19 segments and are believed to be independently derived. Character 65 also deals with segment number.
Character 26: Gonqpodal Prefemur Long
The apomorphic condition is an elongated gonopod (Fig. 24).
This state is seen in the Subfamily Scytonotinae with the exception of Scvtonotus and Speodesmus. the two most primitive taxa. The plesiomorphic condition is a relatively short gonopod with some degree of curvature.
Character 27 : Gnathochilarium Rectangular
A rectangular gnathochilarium is autapomorphic for the subfamily Scytonotinae. Taxa with the plesiomorphic condition exhibit a quadrate gnathochilarium.
Character 28: Syncoxostemal Spine Present
Another autapomorphy for the subfamily Scytonotinae, is the presence of lateral ^ines on the syncoxostemal plate lateral to the hooks (Character 22) (Fig. 64). 141
Character 29: Claw Constricted
The autapomorphic condition, present only in the Scytonotinae,
is a distally constricted claw (Fig. 54). The plesiomorphic state has a gradually tapered claw (Fig. 55).
Character 30: Leg Sphaerotrichomes
In the Scytonotinae, the autapomorphic condition involves sphaerotrichomes present only on the tibia and tarsus. The plesiomorphic state has other podomeres with sphaerotrichomes.
Character 31: Collum Enlarged
The apomorphic state, viiere the collum is enlarged and equal to or wider than the head is present only in the Scytonotinae. The plesiomorphic state involves a small collum much narrower than the head.
Character 32: Head with Microsetae
The head in the Scytonotinae is covered with numerous short microsetae. This autapomorphy gives the head an almost fuzzy appearance. The plesiamorphic state has a few, long setae present.
The following 24 characters (characters 33-56) deal with the genera in the Subfamily Scytonotinae: 142
Character 33: legs Robust
The apomorphic state of relatively robust male legs is characteristic of the subfamilies Polydesminae and Epanerchodinae as well as the genus Speodesmus in the scytonotines.
Character 34: Gonopod Prefemur Not Hirsute
The gonopod prefemur is not hirsute in the genus Speodesmus.
The plesiomorphic state is #iere the prefemur is hirsute.
Character 35: Anterior Edge of Gonopod With Serrations
This apomorphy in vbich the anterior edge of the gonopod has a small field of serrations is present in the genera Speodesmus (Fig.
16) and Coronodesmus (Fig. 30).
Character 36: Gonopodal Femur with Setal Row
This apomorphy, present in the genera Speodesmus and
Bidentogon. is a linear row of long setae on the posterior surface of the femoral region of the gonopod. These setae are not present on any other polydesmids.
Character 37: Tibiotarsus with Caudal Projection
Speodesmus is the only taxon with a proximally directed ^in e on the anterior edge of the tibiotarsus distal to the serrations
(Fig. 16). The plesiomorphic condition is spineless. 143
Character 38: Hypqproct Hirsute
The hirsute hypqproct is apomorphic for Speodesmus and
Coronodesmus. This condition usually involves over 6 pairs of setae, as coitpared to the plesiomorphic condition with 2 pairs.
Character 39: Epiproct Hirsute
The hirsute epiproct is apomorphic for most of the subfamily
Scytonotinae with the exception of some species in the genus
Speodesmus. The plesiamorphic condition is a single pair of setae.
Character 40: Terga Directed Ventrad
This apomorphic state, limited to the subfamily Scytonotinae, is caused by the paranota being ventrally directed. This is opposed to the laterally projecting paranota of most polydesmoids vhere the milliped has a definite flat-backed appearance (Fig. 48). This condition is most extreme in males of the genus Scvtonotus vhere the boc^ is almost circular in x-section (Fig. 49).
Character 41: Dorsal Setae Hooked
This autapomorphic state is present only in the genus
Scvtonotus. Dorsal setae of adults are filiform and hooked terminally. This condition is not exhibited by juveniles, vhich have irregular clavate setae. The difference between adult and juvenile setae in other taxa is unknown. 144
Character 42: Peritremata Large
The apomorphic condition involves large, wide peritremata.
This character is found in Scvtonotus and Idahodesmus of the
Scytonotinae, and in the subfamilies Epanerchodinae and
Polydesminae. In the plesiamorphic condition, the peritremata are
either absent or very small.
Character 43: Gonopodal Femur Branched
The autapomorphic state of a branched femur is present only in
the genus Scvtonotus (Fig. 21). Both branches are movable. The
primitive state has an unbranched femur although it may have various
lobes and processes.
Character 44: Anterior Edge of Collum Flared
This autapomorphy for the genus Scvtonotus is not present in
any other taxa studied. It allows the collum to over-ride the head
giving additional protection to it while the milliped is flat and
extended.
Character 45: Ozopores Elevated
This apomorphy is present in the more derived
Scytonotinae. The ozopores are elevated on large bulbous mounds
(Fig. 9). This state is present in the genera Utadesmus. Bidentogon.
Coronodesmus. Speorthus. Idahodesmus. and several species of the genus Scvtonotus. 145
Character 46: Dorsal Tubercle Formula 10, 10, 8
The dorsal tubercle formula, 10, 10, 8 , representing the
number of tubercles in each transverse row on the dorsum, is
apomorphic in three genera, Bidentogon (Fig. 7), Coronodesmus (Fig.
8 ), and Idahodesmus (Fig. 10). The plesiomorphic condition is of
three rows with 8 tubercles.
Character 47: Gonopod Elongate
The apomorphy, present in genera Utadesmus (Fig. 24) and
Bidentogon (Fig. 27), is exhibited as an elongated gonopod. The plesiomorphic state is a ^ o r t arcuate gonopod.
Character 48: Endomerite Flattened
In the genus Utadesmus this autapomorphy is the transverse flattening of the endomerite. In the plesiomorphic condition the endomerite is usually slightly flattened longitudinally as in
Pseudopolydesmus.
Character 49: Dorsal Setae Clavate or Peg-Shaped
The apamorgdiic state present in several genera, Utadesmus.
Speorthus. and some Speodesmus. is that the setae of adults are short and either peg-shaped (Fig. 52) or slightly clavate (Fig. 51).
The exact relationship between the peg or clavate setae is unknown.
The plesiamorphic condition is the presence of typical filiform setae (Fig. 50). 146
Character 50: Paranota with Cephalad-Directed Denticles
The apomorphic state is distinguished by a large denticle on
the anterior edge of each paranotum (Fig. 9). This state is present
in Bidentogon. Coronodesmus. Speorthus. and Idahodesmus.
Character 51: Tibiotarsus Branched
The apomorphic state of major branching of the tibiotarsus is present in Bidentogon. Coronodesmus. Speorthus. and Idahodesmus.
Al thou^ some branching occurs in other taxa, such as Polvdesmus and
Epanerchodus. these are believed to be derived independently.
Character 52: Anterior Row of Dorsal Setae Directed Cephalad
The apomorphic state, present in Bidentogon (Fig. 7) and
Coronodesmus (Fig. 8 ), is vhere the setae associated with the anterior of three dorsal rows of tubercles are directed cephalad with the posterior two rows of setae directed posteriorly. The plesiomorphic condition has all three rows of setae directed posteriorly.
Character 53: Sterna Modified
The apomor%hic state, exhibited by Coronodesmus.
Epanerchodinae, and some Polydesminae, is the modification of the sterna with spines, and plates. It is atypical in Coronodesmus vhere sternal spines are located only on the tenth sternum. The plesiomorphic state is a flat and unmodified sternum. 147
Character 54; Dorsum with 4 or 5 Rows of Tubercles
The apomorphic state of 4 or 5 transverse rows of dorsal tubercles is present in Scvtonotus (Fig. 5), Speorthus (Fig. 9), and
Idahodesmus (Fig. 10). The remaining scytonotines have the plesiamorphic condition of only 3 rows of tubercles. The rows are less regular in 4- or 5-row taxa than in 3-row taxa.
Character 55: Leg Bairs Separate on Sternum
The apomorphic state, observed in the Sc^dionotinae, is eidiibited by the slight separation of the anterior pair of legs from the posterior pair on the midbody sternum (Fig. 60). The plesiamorphic state has the leg pairs contiguous (Fig. 59).
Character 56: Medium-sized Individuals Absent
This character is used in conjunction with character 15 to obtain some usable information dealing with size. The apomorphic condition is the absence of medium-sized individuals (between 1 0 -
20mm), all being small (less than 10mm). Speorthus and Idahodesmus are apomorphic.
The placement of the subfamilies Epanerchodinae and
Polydesminae in an independent clade is due to similarities observed by previous workers and their overall general appearance. The following 3 apomorphies (characters 57-59) support this clade: 148
Character 57: Anterior Terga Directed Lateral
This apomorphic condition, present in the epanerchodines and
polydesmines, is the presence of laterally-directed paranota on
segments 2-4 (Fig. 44). The plesiomorphic condition has paranota 2-4
curving or bending anteriorly (Fig. 45).
Character 58: Seminal Vesicle Present
The presence of the seminal vesicle at the terminal end of the
seminal groove has always been a good apomorphy for the
epanerchodines and polydesmines as stated by Attems (1914). The plesiomorphic condition is the lack of this terminal seminal vesicle.
Character 59: Dorsum with Smooth Bosses
The apomorphic state, dorsum covered with flat, non- tuberculate bosses, is present in the epanerchodines and polydesmines. In the plesiomorphic condition, the dorsum is covered to various degrees with tubercles.
Character 60; Endomerite large
The apomorphic state, an enlarged endomerite, is present in the epanerchodines and seme polydesmines. The plesiomorphic condition is either the absence or the presence of a small endomerite. 149
Character 61: Cyphopod Modified Ventrally
The apomorphic condition is the modification of the ventral surface of the cyphopod. This condition is best ej^ressed in the genus Pseudooolvdesmus vhere the surface is enlarged into a large irregular flat surface (Fig. 67).
Monophvlv of the Subfamily Epanerchodinae
As stated in the descriptions, Attems (1901) placed this taxon in Polvdesmus as a subgenus. He corrected this in 1914. This taxon of approximately 1 0 0 species has not been revised althou^ a need is present. An indepth study of this taxon is beyond the scope of this study. From the limited number of specimens examined (Appendix D) and the literature, there are 4 autapomorphies (characters 62-65) for the subfamily Epanerchodinae:
Character 62: Clivus Present
The presence of the clivus on the gonopod is an autapomorphy for the epanerchodines (Fig. 15). This usually bulbous structure is located laterally about 1/3 distance rp the tibiotarsus. In many cases it is closely associated with the endomerite.
Character 63: iperture Triangular
The apomorphic state, present in the epanerchodines, is the triangular shape of the gonopodal aperture on the posterior sternum 150
of the 7th segment. The plesiamorphic state is a transversely oval
aperture.
Character 64: Secondary Lobe on 7th Antennomere
In the epanerchodines another apomorphy is the presence of a
small secondary lobe located proximal to the primary lobe. The
plesiomorphic state has only the small primary lobe present.
Character 65: Boc^ with 19 or 20 Segments
In conjunction with character 25, presence of 19 or 20 body
segments is apomorphic %hile the plesiamorphic state is only 20. The
apomoiphic condition is seen only in the subfamily Epanerchodinae.
The phylogenetic tree generated by the PAUP program for the
family Polydesmidae and the genera of the subfamily Scytonotinae is
based on the preceding 65 character states (Tables 2a & b). The
individual states for each taxon can be seen in Tables 3a & b. The
minimum tree length is 92 character steps. The difference between
the absolute minimum length of 6 6 and the determined minimum of 92
is due to reversals and parallelisms.
The cladogram (Fig. 43) shows that there are two major clades
in the family Polydesmidae; one including the subfamilies
Mastigonodesminae and Scytonotinae, and the second including the
subfamilies Polydesminae and "Epanerchodinae. In the subfamily
Scytonotinae, vhich includes only North American representatives. 151 the first genus to separate is Speodesmus. followed by Scvtonotus and then utadesmus. The remaining four genera form two clades, one consisting of the genera Bidentogon and Coronodesmus and the second
Speorthus and Idahodesmus.
The polydesmine-epanerchodinae clade is well established. The
Polydesminae includes two large genera, Pseudooolvdesmus and
Polvdesmus. the first Nearctic, the second Palearctic. There are also several small, mostly monotypic genera from central Asia of uncertain positions although based on the literature, they have close affinity to Polvdesmus.
The consistency index (Cl) is a h i ^ 0.717. If the tree contained no reversals or parallelisms, this index would be 1 .0 0 0 .
Variation is responsible for this value.
The f-value and normalized f-value were 3.124 and 0.161 respectively.
The Manhattan or character state matrix (Table 4) can be used to show relative distances between taxa on comparable taxonomic levels. The differences would range between 0 and 1 for the value generated. In a combined stuc^, part of that range would be taken up by the second level of taxa (in this case the genera of
Scytonotinae). If the second taxonomic level was removed from the calculation, the differences between the taxa would increase proportionally but still be based on a range between 0 and 1. On the subfamily level, the greatest distance is between Epanerchodinae and
Doratodesmidae, with a distance of 0.354, vhile the smallest character distance is between Polydesminae and Epanerchodinae 152
(0.138). In the subfamily Scytonotinae, the greatest distance is between Idahodesmus and Speodesmus (0.328), viiile the shortest distance is between Idahodesmus and Speorthus (0.062).
The patristic distance matrix (Table 5) would show the same proportionality as the character state distances mentioned above.
The greatest subfamilial distance is between Epanerchodinae and
Doratodesmidae and between Mastigonodesminae and Doratodesmidae
(0.354), vhile the smallest distance is bettfeen Polydesminae and
Epanerchodinae (0.138). In the subfamily Scytonotinae the greatest distance is between Idahodesmus and Speodesmus (0.668), vhile the shortest distance is between Idahodesmus and Speorthus (0.062).
The homoplasy matrix (Table 6 ) has the same proportionality as the other two matrices. The greatest amount of homoplasy on the subfamilial level is between Doratodesmidae and Mastigonodesminae and between Epanerchodinae and Mastigonodesminae (0.031) with a single homoplastic character. The other pairwise distances are
0 .000.
In the subfamily Scytonotinae the greatest amount of homoplasy is between Coronodesmus and Scvtonotus (0.123), and the lowest degree of homoplasy (0 .0 0 0 ) is shared with a number of pairwised taxa.
The Deltran test, vhich maximizes parallelism to reversal, lists no change in the tree length or consistency index. However, changes in the cladogram, with variations in the corresponding clades, lowered the f-value from 3.124 to 2.200, and the normalized f-value from 0.161 to 0.113. There were four charcters listed for 153 possible change in their character stater in the case of marginal ozodeme location (12), seen in Bidentogon and is probably a reversal of secondarily derive because the change would require the change of state for four taxa, unlikely; in small size (15), seen in the Mastigonodesminae and Speodesmus prdoably represent secondarily derive size, since both taxa appear to be well established troglodites; in leg modifications (18), are in most cases appear to be independently derived; in strictly asian in range (24), seen in the doratodesmids and the epanerchodines probably represent a primitive stock in the doratodesmid, the sister group for the polydesmids, vhile the epanerchodines a very recent taxon.
The Acctran test, vhich maximizes reversals to parallelisms, addressed the same characters mentioned above, but with a reversal of emphasis. The arguments are the same for the stated conditions.
Although the cladogram remained the same the variations lowered the
F-value from 3.124 to 2.460 and the normalized F-value from 0.161 to
1.117.
Monophvlv of the Genus Pseudopolydesmus
As is the case with almost all milliped taxa, there have been few systematic studies to determine diagnostic characters in polydesmids. Attems' (1898) original description distinguished
Pseudopolydesmus from Polvdesmus only in a key without further statement. He used the absence of the "haarpolster" (^endomerite) and "samenblase" (=seminal vesicle), vhich are present in
Polvdesmus. to separate the two genera. These two structures are 154
present in Pseudopolydesmus and he corrected this oversi^t in 1940.
The first question vhich must be answered here is the monophyly of
the genus Pseudooolvdesmus. Characters 1 to 9 below are the
suggested autapomorphies for Pseudooolvdesmus ; characters 10-35 are
for distinguishing clades or species in Pseudooolvdesmus.
Character 1; Presence of Process
In all Pseudooolvdesmus. presence of the process on the
tibiotarsus is autapomorphic. A comparable process is present on
several species of Polvdesmus but I believe that these are derived
states.
Character 2: Connected Medial Bosses
The plesiomorphic condition of two medial bosses (MB)
(anterior and posterior) is present in Polvdesmus (Fig. 11). In
Pseudooolvdesmus. the autapomorphic state is exhibited the fusion
of these two bosses into one (Fig. 12). It appears that the
remaining polydesmines are variable, but the majority have two medial bosses. All other members of the family appear to have the
plesiomorphic condition.
Character 3: Sternum 3 of Female Unmodified
The third sternum of Polvdesmus is modified with variously
shaped lobes, spines, and processes (ex. Demange, 1968). In the 155
autapomorphic condition, seen in Pseudopolydesmus. the third sternum
is flat and unmodified.
Character 4: Presence of cyphopod Plate
The autapomorphic condition is the presence of a distal,
flattened, and variously modified plate-like dorsal enlargement of
the cyphopod, present in Pseudooolvdesmus and not present in any
other polydesmids. In Polvdesmus this dorsal area may be someiÆiat
modified but not flattened.
Character 5: Sternum 7 of Male Modified
Althou^ the amount of modification is not large, a laterally
enlarged 7th sternum is present in all Pseudooolvdesmus and absent
in the genus Polvdesmus. making it a good autapomorphy.
Character 6 : Anterior Tibiotarsal Attachment
The attachment of the tibiotarsus to the gonocoxa is more
lateral in Polvdesmus. and more anterior in Pseudopolydesmus. This autapomorphy gives Pseudooolvdesmus the appearance of a more firmly attached tibiotarsus.
Character 7; Endomerite Enlarged
The large size of the endomerite, used by Attems (1940), appears to be a good autapomorphy for Pseudooolvdesmus. The endomerite is also more consistent in size and location than in
Polvdesmus. 156
Character 8 ; Cyphopcd Elongate
The longitudinal elongation of the cyphqpod is autapomorphic.
This character is seen in all Pseudopolvdesmus ; in Polvdesmus the
shape is more oval. The cyphqpod is secondarily reduced to a
sli^tly oval shape in minor, caddo. and probably paludicola c is a result of their small size.
Character 9: Tibiotarsus with Terminal Setae
The presence of a row of setae on the tip of the tibiotarsus
is autapomorîMc for Pseudopolvdesmus (Fig. 91). Setae may be absent
from specimens due to breakage. The plesiamorphic state has no terminal setae.
Character 10: Presence of Process M 3
Process M 3 is considered apomorphic because it is always present in the Group Canadensis (Fig. 70). In the remaining species of Pseudopolvdesmus. the process is absent.
Character 11: Cÿphopodal Plate Enlarged
The enlargement of the dorsal surface of the cyphopod is also apomorphic for the Group Canadensis (Fig. 103). The plesiamorphic condition, seen in the Group Serratus, is ejdiibited by a smaller plate. 157
Character 12: Numerous Sternal Setae
The number of sternal setae is characteristic. Grotç»
Canadensis has the apomorphic condition of numerous setae. In Group
Serratus, the number always appears to be less than 10.
Character 13: Presence of Process
Presence of process is apomorphic in the Groiç) Canadensis.
The process is absent in the Grovç) Serratus. Although in Polvdesmus
there are species with processes in a similar position as process
E^, it is accepted that the plesiamorphic condition is with the process missing. This is because character states have not been
determined in Polvdesmus and the simplest form is probably plesiamorphic. The variability of all process should increase the probability of homoplasies.
Character 14: Small Paranatal Denticles
The small size of the lateral denticles is apomorphic for the
Group Canadensis. In many cases these denticles are almost nonexistent. In Hoffman's description of paludicola he stiatzes that denticles are absent, however this not the case. They appear to be
reduced; however more specimens are needed to establish variability.
Character 15: Ipiproct Downtumed
In the species pinetorum the epiproct is d o w n t u m e d (Fig. 77).
This condition is not normal in any other Pseudopolvdesmus (Fig. 158
78) ; however, some specimens of other species showed an
uncharacteristic downtumed condition.
Character 16: Gonopod Thickened longitudinally
In the Groiç) Canadensis, the apomorphic condition of the
longitudinal thickening of the tibiotarsus around the endoraerite
characterizes the tallulanus-erasus clade (Figs. 80, 81). The
plesiomorphic condition has the typical thin tibiotarsus.
Character 17: Presence of Process E 3
The presence of process E 3 is apomorphic for two grorps of
taxa, the Group Serratus and the canadensis-collinus ccstplex. This
is probably a case of parallelism: the appearance of this process
twice in unrelated taxa. I speculate that the low number of genes
controlling these processes permits mutations, causing similar phenotypes.
Character 18: Presence of Process M 2
The presence of the M 2 process is autapomorphic for the canadensis-collinus complex.
Character 19: Presence of Process E 2
The loss of the E 2 process is autapomorphic for erasus.
Character 20: Montane Habitat
The use of habitat information is not customary but is included because of its distinctive influence. Here the generalized 159 habitat is assumed to be plesiomorphic. The mountain or cold- dwelling habitat can be seen in two species. In the erasus- tallulanus complex, tallulanus is found only in the central and southern i^palachians at hi^ e r elevations than erasus vAiich is found primarily on the Cumberland Plateau below 450 meters, west of tallulanus. There does not appear to be any overlap of the two species.
In the canadensis-collinus complex, canadensis is found only at hiiÿier elevations of the Appalachians and expands beyond them in the northern areas of its range. P. collinus appears to be a generalist of a different type. The species has been found from high elevations (over 2 0 0 0 meters) to muddy river flood plains, to sandy coastal regions.
Character 21: Presence of Process E 2 + 3
The presence of process E 2 + 3 is apomorphic in the Group
Canadensis except for pinetorum. It is however, attached differently in the erasus-tallulanus and canadensis-collinus complexes. In erasus-tallulanus the process is basally attached to the tibiotarsus
(Figs. 8 8 , 89) vhile in canadensis-collinus the process is elevated on a distinct lateral stalk.
Character 22: Anterior Edge of Gonocoxae Multilobed
This apomorphic condition is present in the Group Serratus, and with the exception of collinus. absent in the Group Canadensis
(Fig. 82). Here the apparent parallelism is probably caused by 160 inultiple gene loci, not ky a single locus. There appears to be a h i ^ degree of variation in the gonocoxae, ranging from monolobel to multilobed.
Character 23: RnDcesses M 2 and Connected Ventrally
The connection of these two processes along the posterior face of the tibiotarsus is autapomorphic in ttie Group Serratus. Both processes may be present in several other species, but the connecting ridge is not present. This ridge is most clearly seen in
Pseudopolvdesmus serratus (Fig. 99). The connecting ridge may also bear a small process; however, the latter appears sporadically throu^out the range, without discernable geographic pattern.
Character 24: Sternum 7 laterally Enlarged
As mentioned in character 5, the modification of the lateral edge of the 7th sternum is apomorphic for Pseudopolvdesmus. There is further modification of the structure in the Group Serratus into a moderately long, thin spine (Fig. 62). This contrasts with a very weak spine in the Group canadensis (Fig. 61).
Character 25: legs Short
Presence of relatively short legs is autapomorphic for the minor-caddo complex and paludicola. Although all three species are noticeably smaller than the others in the genus, the relative leg length to boc^ length is also smaller. In these species leg length to boc^ length ranges from 9-14% vhile in the remaining species this ratio is over 16%. 161
Character 26; Body Length Short
The overall smaller size of individuals in the minor-caddo
coitplex and paludicola is a good autapomorphic character. These
species have a length ranging from 9 to 14mm vhile the remaining
species range between 12 and 32mm. The reduction in boc^ size can be
seen in many milliped taxa.
Character 27: Dorsal Setae Present
The presence of dorsal setae is apomorphic for the minor-
caddo cortplex and paludicola. These small setae are associated with
the bosses and denticles. There are also cases vhere other species
have dorsal setae, for example in collinus. but this is rare. Each
boss and denticle will usually have a single seta, however two setae
are occasionally present.
Character 28: Antennae Short
The autapomorphic condition of short antennae is present for
paludicola and the minor-caddo complex. Again these three species
are small (character 25), but the relative length of the antennae is
also shorter. In these three species, the relative antennal length
ranges from 1 0 to 1 2 % of the total body length vhile the plesiomorphic condition ranges between 15 and 21%.
Character 29: Cÿphopodal Plate Margin Straight
As mentioned earlier (character 4), the presence of the cÿphopodal plate is apomorphic for the genus. However the condition 162
eidiibits two general shapes, quadrate and argular. The strai^t
plate margin is apomorphic, and can be seen in the minor-caddo
coirplex (Figs. 115, 116). The status of paludicola is unknown but I
believe that vdien females of this species have been examined, they
will be similar, so this character is used as an apomorphy for all
three species. To restrict the trait to minor and caddo would only move the apomorphy one branch distally, and change nothing in the
phylogenetic pattern.
Character 30: lowland Habitat
As in character 19 the use of geographical information may be
somevhat suspect due to inccarplete collecting. As before, the
generalized habitat is considered plesiomorphic. Three species: paludicola. minor, and caddo exhibit apomorphic lowland preferences,
inhabiting low, wet, flood-prone areas. Two of these, minor and
caddo. are usually found along the margins of major rivers vAiich are prone to flooding. P. paludicola has been found in lowland swaitps of
eastern Virginia and North Carolina, in very wet localities. These
localities are not generally regarded as good milliped habitats.
During personal collecting, the caddo type locality had flood debris
on overhanging power lines and water marks 9-12 meters up trees.
This habitat does not preclude the occasional occurrence of other species, as, for example, serratus in Louisiana. 163
Character 31: Process M 4 Absent
The absence of M 4 is apomorphic in two species, paludicola and
caddo. with the plesiamorphic condition present in minor (vhich is
a probable case of reversal). The plesiamorphic condition is present
in the remaining species of the genus as well as in Polvdesmus where
some type of terminal medial process is usually present.
Character 32: Dorsal Curvature Decreased
In minor and caddo the bo<^ is depressed dorsoventrally. This
apomorphic condition may be related to habitat preference (see
character 30). The width/thickness ratio of minor and caddo is less
than 80% as ccarpared to over 90% in the remaining species.
Character 33: Bunched cyphopod Setae
This condition is apomorphic for at least the minor-caddo
cdtplex but may include paludicola (no female paludicola have been
positively identified). This condition involves a distinct ventral bunching of marginal setae (Fig. 114) on the cyphopod as corrpared to
the regularly spaced setae seen in all other polydesmids (Fig. 107).
Character 34: Process Isolated from Endomerite
The general location of process is near the base of the
endomerite. The apomorphic condition is more proximal and separate
from the endomerite (Fig. 94). This is an autapomorphy for minor. 164
Character 35. Sternum II of Female Modified
The presence of small, smooth buirps on the second sternum of
females is an autapomorphy for Pseudonolvdesmus caddo (Fig. 76). The plesiamorphic condition is the absence of any modification on
sternum II.
Character 36. Dorsum Flattened
There appears to be a trend in many polydesmid taxa to flatten the bo(^. The extreme case can be seen in minor vhere the boc^ has a relative width/thickness ratio of approximately 70% as compared to
80% in caddo and over 90% in the remaining species. This is a further and noticable step beyond character 32.
The phylogenetic tree generated by the PAUP program for the genus Pseudopolvdesmus was based on the preceding 36 character states (Table 7). The data set listing the states present in each taxon is in Table 8 . The tree (Fig. 118) easily splits Polvdesmus from Pseudopolvdesmus based on 9 autapomorphies. From this point the tree is divided for clarity into two Groips, Canadensis and
Serratus. The Groip> Canadensis is composed of five species with pinetorum splitting off first. The next major clade sees the remaining four species divided into two-^^ecies complexes, tallulanus-erasus and canadensis-collinus. The Groip Serratus is composed of four species. The species serratus splits off first. The next derived species is paludicola leaving the final clade consisting of the caddo-minor complex. 155
The minimum tree length was 46 character steps. The difference between the absolute minimum of 35 and the determined minimum of 46 is due to reversals and parallelisms.
The consistency index (Cl) is a moderate 0.761. If the tree length was 35, without any reversals or parallelisms, this ratio would be 1 .0 0 0 .
The f-value and normal f-value are 3.296 and 0.196 respectively, vhich is also moderate.
The Manhattan or character state distance matrix (Table 12) is the pairwise distance between two taxa. The greatest distance for
Pseudopolvdesmus is between the minor-caddo and the tallulanus- erasus and canadensis-collinus corrplexes (0.629). The shortest distance is between minor and caddo. and tallulanus and erasus
(0.114).
The patristic distance matrix (Table 13) shows that the greatest distance is between minor and collinus (0.829) vhile the shortest distance is between canadensis and collinus (0.086).
The greatest amount of homoplasy as presented in the homoplasy matrix (Table 14) is between minor and collinus (0.286), vhile a number of taxa showed no homoplasy between them (0 .0 0 0 ).
The Deltran test, vhich maximizes parallelism to reversal, shows no variation in the above calculations. There were five characters listed for possible change in their character state: in the case of cÿphopodal shape (8 ), the parallelism of the oval shape seen in minor and caddo is probably related to size and therefore secondary; in antennal length (27), the same is true; in the M 2 -E 1 166 connection (2 2 ), this condition also requires a change in the state of both minor and caddo; the loss of process M 4 (30) is seen in caddo; and the multilobed condition seen in collinus and Grovç)
Serratus, may be due to gonocoxal variation.
The Acctran test, vhich maximizes reversals to parallelisms, was the same as the original Farris test.
CLASSIFICATION OF POLYDESMIDAE
My classification of the Polydesmidae is summarized in Table 1 and is es^jressed as a phyletic sequence scheme (Wheeler, 1979). In the phyletic sequence, each taxon is the sister groiç) of all taxa of equal or lesser rank listed below it. Also included are the tentative subfamilies and genera from the Old World that were not covered in this stuc^; these are marked with an asterisk.
In this classification, I retain the subfamily
Mastigonodesminae with 3 genera and propose new subfamily status for
Scytonotinae, with a new genus, Idahodesmus. and the new name
Coronodesmus. to handle the observed diversity. Also proposed are the subfamilies Polydesminae to include a number of palearctic genera and Pseudopolvdesmus. and Epanerchodinae to include from 3 to
8 Asian genera. ZOOGEOGElABffif
In a morphological cladistic analysis, insists into relationships among taxa correlate with the areas these taxa occupy.
Since a special effort was made to obtain locality information for the genus Pseudopolvdesmus. a moderately satisfactory picture of distribution of its species was obtained. In the absence of corrplete locality information, general distribution patterns, especially in the Scytonotinae, can only be surmised.
THE VICARIANCE MDDEL
Theories explaining the distribution of most animal groi:qps fall into two categories, dispersal and vicariance. The ideas dealing with dispersal have been chairpioned by Darlington (1957),
Brundin (1966), and Hennig (1966). The vicariance model is more recent, first presented in Croizat's Panbiogeocrraphv (1958) and later nKDdified by Croizat, et al. (1974), Platnick and Nelson
(1978), and Rosen (1978). The only difference lies in the type and
167 168 degree of isolating barriers. Vicariance is viien a widespread population, or species, has its range divided into smaller populations by developing barriers. Dispersal takes place if the barrier is alreacfy present, and success in crossing that barrier leads to the founding of a separate and new population. Both models have as their central point, the idea of allopatric spéciation
(Mayr, 1969), but because of the differences in environmental conditions and genetic make-iç), evolution will act differently on each isolated population.
Platnick and Nelson (1978) stated that by inference it is possible to develop a vicariant model based on the cladistic relationships of the areas. If an initial pattern can be established, and can be seen in other organisms, this is representative of a shared "generalized track" (Croizat, et al.,
1974). The idea is that if a barrier isolates a geographic region, many taxa, including millipeds, should also be isolated.
It is now apparent to most biogeographers that both dispersal and vicariance are itiportant in understanding the distribution of organisms. However the degree of importance varies with the group.
In dealing with flightless soil arthropods vhich spend their lives in a very controlled and relatively homeostatic environment, dispersal probably plays a minimal role. Terrperature, humidity, li^t, and moisture are critical, so barriers don't have to be large to be efficient. Because of the limited mobility of these organisms and their slow rate of dispersion, soil arthropods hold a great future as indicators of vicariant events. 1 6 9
DISTEÎIBünON PATTERNS
Below is a brief discussion of the distributions of the 4 subfamilies and the North American genera of Polydesmidae.
Subfamily Mastigonodesminae. Palearctic: Around the western basin of the Mediterranean Sea, ejctending into northern Italy and central France. Several species are troglodytic.
Subfamily Scytonotinae. Nearctic: In eastern North America north of Tennessee and south of Canada, and in the west from Texas,
%oming, and California to southern Alaska (Map 3).
1. Speodesmus. Nearctic: Caves of west-central Texas.
2. Scvtonotus. Nearctic: Two populations, one from Iowa and
Arkansas east to the Atlantic coast, and the other from vyoming and
Utah northwest into southern Alaska.
3. Utadesmus. Nearctic: Utah and New Mexico.
4. Bidentoaon. Nearctic: Coastal central California.
5. Coronodesmus. Nearctic: Central California.
6 . Speorthus. Nearctic: Caves of southern and central New
Mexico.
7. Idahodesmus. Nearctic: Two localities in northwestern
Idaho.
Subfamily Polydesminae. Holarctic: Palearctic from Europe to extreme western Siberia (although how extensive in Siberia is 170
uncertain) and the extreme northwestern tip of Africa (Morocco,
Algeria, and Tunisia). In the Nearctic, all except Mexico and the
Central Plains. All species are epigean.
1. Polvdesmus. Palearctic: Entire Eurasian continent north of
30°. Hipest concentration of species in the Balkan Mountains.
2. Pseudopolvdesmus. Nearctic: Eastern North America, from
eastern Texas and Wisconsin east to the Gulf and Atlantic coasts,
northern Florida to Hudson Bay, Canada.
Subfamily Epanerchodinae. Palearctic: Japan, Korea, and China.
Some species are trogloc^^ic.
The structures and exact relationships of the exotic genera of
Polydesmidae are uncertain with the exception of the genus
Epanerchodus which is well described in the literature. Although few
specimens and inadequate data made clarification of other central
Asian and European polydesmids impossible, the genus Epanerchodus. with several associated genera, should be elevated to subfamilial
ranking.
Spéciation Patterns- An inportant part of zoogeography is the
appraisal of spéciation patterns. All distributions are obviously
continental. Two patterns can be observed in the Polydesmidae: wide-
^read genera with numerous species (Polydeanus, Pseudopolvdesmus,
Epanerchodus. and Scr/tonotus) and genera with a limited distribution
and few species. 171
The species of Pseudopolvdesmus fall into three distribution and habitat patterns. Pseudopolvdesmus minor, caddo. and paludicola inhabit lowlands and major river flooc^lains or areas vàiere inundation by flood waters is hi^ily probable (for example trees at the type locality of minor in Louisiana had water marks thirty feet above the ground). The second type involves primarily montane and colder habitats. This is characterized fcy canadensis, tallulanus. airi possibly pinetorum althou^i to a lesser extent. The disjunct populations of canadensis in southern Mississippi and central
Florida probably represent isolated relicts from the last period of glaciation. The remaining species, serratus. erasus. and collinus appear to be generalists with little elevation preference, with serratus wide-spread and erasus and collinus less so. At least two species pairs are seen, canadensis-collinus and tallulanus-erasus. with one member northern or montane and the other primarily not. The non-montane species may represent young species dispersing since the last period of glaciation.
In the Scytonotinae, with its high species to genus ratio, adequate data for only one genus, Scvtonotus. are available. The eleven species are divided into eastern and western populations supported by a number of distinct morphological characters. Although disjunct based on present data, it is possible that they are continuous through southern Canada. I expect that the genus was wide-spread across the North but increasing aridity in the Great
Plains caused the extirpation of the genus there. It would be 172
interesting if relict populations occur in the gap between V^oming
and Iowa.
HISTORICAL ZOOCŒOGRAEHY
The patterns for the family Polydesmidae indicate a moderately
sinple model because of the Holarctic distribution. The family in
its present form didn't arise until after the breakup of Pangea (180
HKBP). The Holarctic was continuous until late Cretaceous (75 MYBP) vhen the Palearctic and Nearctic continents separated. The
distribution of the family basically conforms to the Arcto-tertiary
Nemoral Circunpolar Forest of the Northern Hemisphere (Itomas,
1972). This forest was continuous ip to the Miocene and possibly the
Pliocene (7 MÏBP) and then diminished to its present-day distribution. Only three major areas of this substantial deciduous
forest survive; eastern North America, western Europe, and the Far
East of Asia including China and Japan, each isolated by seas or intervening drier areas. All three areas are centers vhere radiation and spéciation has taken place, Polvdesmus in Europe, Epanerchodus in the Far East, especially Japan, and to some extent
Pseudopolvdesmus in North America. In west and central Asia and western North America only isolated populations remain with a few relict genera and species. BIBIiTOGRAHK
Aley, T. 1972. Ozark Underground Laboratory Newsletter. 5(2) :1. (Not
seen).
Attems, Carl. 1898. System der Polydesmiden. 1 Theil. Denkschr.
Akad. Wiss., Wien (Math.-naturwiss. Classe) 67:221-482, pis.
1-11, figs. 1-276.
------. 1901. Neue durch den Schiffsverkehr in Hamburg
eingeschleppte ly^iapoden. Mitt. Naturh. Mus. Hanburg. 18:109-
116, pi. 1 .
------. 1914. Die Indo-Australisdhen myriopoden. Arch.
Naturg. 80(4):1-398, pis. 1-7, figs. 1-125.
------. 1926. Myriopoda, b) Kukenthal und Knmibach, Handbuch
der Zoologie, Hamburg Zool. 4:1-402, figs. 1-477.
------1931. Die familie Leptodesmidae und andere
Polydesmiden. Zool., Stuttgart, 30(3-4):1-149, figs. 1-245.
------1940. Family Polydesmidae, Vanhoeffeniidae,
Cryptodesmidae, Oniscodesmidae, Sphaerotrichopidae,
Peridontodesmidae, Rhachidesmidae, Macellolophidae,
173 174
Pandircdesmicaae, in Das Tierreidi. Lief. 70 (Polydesmoidea
III);1-577, figs. 1-719.
Bailey, John W. 1928. The Chilopoda of New York State with notes on
the Diplopoda. New York State Mis. Bull. 276:1-50, figs. 1-15.
Baker, G. H. 1978. The population dynamics of the millipede,
Qmmatoiulus moreletii (Diplopoda: lulidae). J. Zool. London.
186:229-242.
Blower, J. Gordon. 1970. The millipedes of a Cheshire wood. J. Zool.
London. 160:455-496, figs. 1-12.
Blum, Murray S. 1981. Chemical Defenses of Arthropods. Academic
Press, New York. 562pp.
Bollman, Charles H. 1887a. Description of fourteen new species of
North American myriapods. Proc. U. S. Nat. Mus. 10:617-627.
------. 1887b. New genus and species of Polydesmidae. Ent. Amer.
3(1):45-46.
------1888. A preliminary list of the Myriapoda of Arkansas,
with descriptions of new species. Ent. Amer. 4:1-8.
------. 1893. The r^iapoda of North America. U. S. Nat. Mus.
Bull. 46:1-210.
Branson, Branley A., and Donald L. Batch. 1971. Records for and
annotations on some valley millipeds (Diplopoda) from Northern
Kentucky. Trans. Kentucky Acad. Sci. 31:79-83.
Brimley, Clement S. 1938. Insects of North Carolina, being a list of
the insects and their relatives. North Carolina Department
Agriculture, Div. Ent., Ralei^i. 560pp. 175
Brolemann, Henry W. 1916. Essai de classification des polydesmiens
(myriapodes). Ann. Soc. Ent. France. 84(4) : 523-608, figs. 1-
18.
Brundin, Lars. 1966. Transantarctic relationships and their
significance as evidenced by chironcmid midges. Kungliga
Svenska Vetenskapsakademiens Handlingar. series 4 11(1):1-472.
Buckett, John S. and Michael R. Gardner. 1968. A new genus and
species of milliped from northern California. Pan-Pacific Ent.
44(3):198-202, figs. 1-7.
Carl, Johann. 1902. Exotische polydesmiden. Rev. Suisse Zool.
10:563-679, pis. 10-12, figs. 1-109.
Causey, Nell B. 1943. Studies on the life history and the ecology of
the hothouse millipede, Orthomorpha gracilis (C. L. Koch).
Amer. Midi. Natur. 29(3):670-682, figs. 1-3.
------. 1950. On four new polydesmoid millipeds. Ent. News.
61(7):193-198, figs. 1-7.
------1952a. Some records and descriptions of polydesmoid
millipeds from the United States. Chicago Acad. Sci. Nat.
Hist. Misc. 106:1-11, figs. 1-8.
------. 1952b. New records of millipeds from southern Ontario.
Canadian Field Nat. 66:145.
------. 1954. New records and species of millipeds from the
western United States and Canada. Pan-Pacific Ent. 30(3):222-
227, figs. 1-5.
------. 1955. New records and descriptions of polydesmoid 176
millipeds (Order Polydesmida) from the eastern United
States. Proc. Biol. Soc. Washington. 68:21-30, figs. 1-7.
------. 1959. Two new troglodytic millipeds from Texas. Proc.
Biol. Soc. Washington. 72:69-74, figs. 1-6.
------. 1963. Additional records of Louisiana millipeds. Proc.
Louisiana Acad. Sci. 26:76-79.
Chamberlin, Ralph V. 1910. Diplopoda from the western states. Ann.
Ent. Soc. Amer. 3(4):233-262. fig. 16.
------. 1911. Notes on myriapods from Alaska and Washington.
Canadian Ent. 43:260-264, fig. 16.
------1914. Notes on myriapods from Douglas lake, Michigan.
Canadian Ent. 46:301-306.
------. 1918a. Four new western diplopods. Pomona Coll. J. Ent.
and Zool. 10(1):9-ll.
------. 1918b. Myriapods from Nashville, Tennessee. Psyche.
25(2):23-30.
------. 1920a. Canadian myriapods collected in 1882-1883 by J.
B. Tyrrell, with additional records. Canadian Ent. 52(6) : 166-
168, figs. 16, 17.
------. 1920b. A new diplopod from Texas and a new chilopod from
Alaska. Proc. Biol. Soc. Washington. 33:41-44.
------. 1925. Notes on some centipeds and millipeds from Utah.
Pan-Pacific Ent. 2(2):55-63.
------. 1930. On some centipeds and millipeds from Utah and
Arizona. Pan-Pacific Ent. 6(3):111-121, figs. 1-11. 177
------1940. On some chilopocSs and diplopods from North
Carolina. Canadian Ent. 72:56-59.
------. 1941. New American millipeds. Bull. Univ. Utah. biol.
ser. 6(4):1-39, pis.1-5, figs. 1-49.
------, 1942a. On a collection of myriapods from Iowa. Canadian
Ent. 74(1):15-17, figs. 1-4.
------. 1942b. New southern millipeds. Bull. Univ. Utah, biol.
ser. 6(8):1-19, pis. 1-4, figs. 1-40.
------. 1943a. On nine North American polydesmoid millipeds.
Proc. Biol. Soc. Washington. 56:35-40, pis. 1-2, figs. 1-11.
------. 1943b. A new Polvdesmus from Missouri and Oklahoma
(Diplopoda). Ent. News 54:15-16, figs. 1, 2.
------. 1943c. On some genera and species of American millipeds.
Bull. Univ. Utah, biol. ser. 8(2):l-20, figs. 1-42.
------. 1943d. Some records and descriptions of American
diplopods. Proc. Biol. Soc. Washington. 56:143-152, pis. 7-8,
figs. 1-15.
------. 1945. On some diplopods from the Indo-australian
archipelago. Amer. Mus. Novit. No. 1282:1-43, figs. 1-137.
------1946. On four millipeds from Georgia and Mississippi.
Proc. Biol. Soc. Washington. 59:139-142, figs. 1-5.
------. 1947. Some records and descriptions of diplopods
chiefly in the collection of the Academy. Proc. Acad. Nat.
Sci. Philadelphia. 99:21-58, figs. 1-73.
------. 1949. Some millipeds of the families Polydesmidae and
Xystodesmidae. J. Washington Acad. Sci. 39(3) :94-102, figs. 178
1-27.
------, 1951a. On ei^ t new southern millipeds. Great Basin Nat.
11(1-2):19-27, figs. 1-16.
------1951b. Records of American millipeds and centipeds
collected by Dr. D. Eldon Beck in 1950. Great Basin Nat. 11(1-
2);27-35, figs. 1-3.
------. 1952a. Three cave-dwelling millipeds. Ent. News. 63:10-
12.
— ------. 1952b. American polydesmoid millipeds in the collection
of the Chicago Museum of Natural History. Ann. Ent. Soc. Amer.
45:553-583, figs. 1-47.
------, 1959. Two new troglodytic millipeds from Texas. Proc.
Biol. Soc. Wa^ington. 72:69-74, figs. 1-6.
------and Richard L. Hoffman. 1950. On some genera and families
of North American diplopods. Chicago Acad. Sci. Nat. Hist.
Misc. 71:1-7, figs. 1, 2.
------. 1958. Checklist of the millipeds
of North America. U. S. Nat. Mus. Bull. 212:1-236.
------and Y. M. Wang. 1953. Records of millipeds (Diplopoda)
from Japan and other oriental areas, with description of new
genera and species. Amer. Mus. Novit. No. 1621:1-13, figs. 1-
6.
Cook, Orator F. 1896. Crvotodesmus and its allies. Brandtia 5:19-28.
------. 1911. Notes on the distribution of millipeds in southern
Texas, with descriptions of new genera and species from Texas, 179
Arizona, Mexico, and Costa Rica. Proc. U. S. Nat. Mus. 40:147-
167.
------and A. C. Cook. 1894. A monograph of Scvtonotus. Ann.
New York Acad. Sci. 8:233-248. pis. 6-9, figs. 1-71.
Croizat, Lson, 1958. Paribiogeography. 2 vols. Published by author,
N. V. Drukkerij Salland Deventer Netherlands. 2849pp.
------, Gareth Nelson and Bonn Rosen. 1974. Centers of origin
and related concepts. Syst. Zool. 23(2):265- 287.
Darlington, Philip J. 1957. Zoogeography: the geographical
distribution of animals. Wiley, New York. 675pp.
Demange, Jean-Paul M. 1968. Etudes des femelles appartenant aux
genres Polvdesmus et Brachydesmus pour servir a une faune des
myriapodes de France. Bull. Soc. Hist. Natur. Toulouse. 104(1-
2):276-305, figs. 1-78.
Elliott, William R. 1976. Morphometries and evolution of Speodesmus
in central Texas Caves (Diplopoda, Polydesmida). Ki. D.
Dissertation, Texas A. & M. Univ. 153pp.
Enghoeff, Henrik. 1981. A cladistic analysis and classification of
the millipede order Julida. Z. zool. Syst. Evol.-forsch.
19:285-319, figs. 1-13.
------. 1984. Phylogeny of millipedes - A cladistic analysis. Z.
zool. Syst. Evol.-forsch. 22:8-25, figs. 1-11.
Farris, James S. 1970. Methods for conputing Wagner trees. Syst.
Zool. 19:83-92.
------. 1972. Estimating phylogenetic trees from distance
matrices. Amer. Natur. 106:645-668. 180
Felsenstein, Joseph. 1981. A likelihood approach to character
w e i ^ t i n g and vÆiat it tells us about parsimony and
canpatibility. Biol. J. Linn. Soc. 16:183-193.
------. 1984. Distance Methods for inferring phylogenies: A
justification. Evol. 38(1):16-24.
Gardner, James E. 1986. Invertebrate fauna from Missouri caves and
springs, conservation Commission of the State of Missouri.
72pp.
Heller. 1858. Beitrage zur osterreichischengrotten fauna. Sitz. ber.
Akad. Wiss. Wien. 26:313-326, fig. 1-14.
Hennig, E. H. Willi. 1966. Phylogenetic Systematics. University of
Illinois Press, Urbana, Illinois. 370pp.
Hoffman, Richard L. 1947. The status of the milliped lasiolathus
vircfinicus. with notes on Scvtonotus granulatus. Proc. Biol.
Soc. Washington. 60:139-140.
------. 1950a. Records and descriptions of diplopods from the
southern i^palachians. J. Elisha Mitchell Sci. Soc. 66(1) : 11-
33, figs. 1-32.
------. 1950b. Notes on some Virginia millipeds of the family
Polydesmidae. Virginia J. Sci., new series, 1(3) :219-225,
figs. 1-4.
------. 1962. The milliped genus Scvtonotus in eastern North
America with the description of two new species. Amer. Midi.
Nat. 67:241-249, figs. 1-12.
------. 1969. The origin and affinities of the Southern 181
i^paladiian diplopoda fauna. Virginia Polytechnic Institute
Res. Div. Monog. 1:221-246, figs. 1-8.
------1974. A new polydesmid milliped from the southern
J^palachians, with remarks on the status of Dixidesmus and a
proposed terminology for polydesmid gonopods. Eroc. Biol. Soc.
Washington 87(31): 345-350, figs. 1-3.
------1977. Ihe ^stematic position of the diplopod family
Doratodesmidae, and description of a new genus from Malaya.
Pacific Insects. 17(2-3):247-255, figs. 1 - 8 .
------1978. Diplopoda from Papuan caves (zoological results of
the British Speleological Expedition to Papua-New Guinea,
1975. Intern. J. Speleol. 9(3-4):281-307, figs. 1-7.
------. 1979. Classification of the Diplopoda. Museum de Geneve
Switzerland. 237pp.
Johnson, Bert M. 1954a. A new species of the milliped genus
Dixidesmus from Michigan. Chicago Acad. Sci. Nat. Hist. Misc.
137:1-5, fig. 1.
------1954b. The millipeds of Michigan. Papers Michigan Acad.
Sci. 39:241-252, pls. 1-7, figs. 1-45.
Jong, R. de. 1980. Some tools for evolutionary and phylogenetic
studies. Z. zool. Syst. Evol.-forsch. 18:1-23.
Kevan, D. Keith MCE. 1983. A preliminary survey of known and
potentially Canadian millipedes (Diplopoda). Can. J. Zool.
61:2956-2975.
Kime, R. Desmond and Georges Wauthy. 1984. Aspects of relationships 182
between millipedes, soil texture and temperature in deciduous
forests. Pedobiologia 26:387-402, figs. 1-4.
KLuge, Arnold G. and James S. Farris. 1969. Quantitative phyletics
and the evolution of anurans. Syst. Zool. 18:201-205.
Koch, Carl L. 1847. System der Myriapoden, jji Herrich-Schaeffer,
Kritische Revision der Insectenfauna Deutschlands, Vol. 3, p.
350.
------. 1863. Die ly^iapoden. Getreu nach der natur abgebildet
undr beschrieben..., 1:1-134, pis. 1-60, figs. 1-123; 2:1-112,
pis. 61-119, figs. 124-234. Halle.
Kbmas, Jan. 1972. In Taxonomy, Riytogeography, and Evolution. Acad.
Press. New York. 431pp.
Latreille, Pierre A. 1802. Histoire naturelle. . . des crustacés et
des insectes. 3:1-467. latzel, Robert. 1884. Die myriapoden der Osterreichisch-Ungarischen
Monarchie. Die Syitphylen, Pauropoden und Diplopoden, Wien,
2:l-XII, 1-414. leach, William E. 1815. A tabular view of the external characters of
four classes of animal tAiich Linne arranged under Insecta. . .
Trans. Linn. Soc. London. 11:306-400.
Loksa, 1958. Eine neue form von Polvdesmus fAcanthtarsius)
edentatulus bidentalus. Verh. aus ungam, und beitrage zur
mikroskipltur der Polydesmiden. Opusc. zool. Budapest.
2(4):49-54, figs. 1-12.
Loomis, Harold F. 1939. The millipeds collected in Appalachian caves 183
by Mr. Kenneth Dearolf. Bull. Mus. Ccsnp. Zool. 86(4) : 165-193.
figs. 1-14.
------, 1943a. New cave and epigean millipeds of the United
States, with notes on some established species. Bull. Mas.
Conap. Zool. 92(7) :371-410, pi. 1, figs. 1-18.
------. 1943b. A new genus of Virginia millipeds related to
Scvtonotus and a new species from Florida. J. Washington Acad.
Sci. 33(10);318-320, figs. 1, 2.
------1944. Millipeds principally collected by Professor V. E.
Shelford in the eastern and southeastern United States.
Psyche. 51(3-4);166-177, figs. 1-6.
------, 1959. Millipeds collected enroute from Florida to San
Antonio, Texas and vicinity. J. Washington Acad. Sci.
49(5):157-163, figs. 1-23.
------. 1960. Millipeds of the Order Polydesmida from the
western states and Baja California. J. Kansas Ent. Soc.
33(2):57-68, figs. 1-17.
------1969. New and known millipeds from four southern
states. Florida Ent. 54(4):245-251, figs. 1-13.
------and Richard L. Hoffman. 1948. Synonymy of various
diplopods. Proc. Biol. Soc. Washington. 61:51-54.
Mayr, Ernst. 1969. Principles of systematic zoology. McGraw-Hill.
New York. 428pp.
McDaniel, V. R. and K. L. Smith. 1976. Cave fauna of Arkansas:
selected invertebrate taxa. Arkansas Acad. Sci. Proc. 30:57-
60. 184
Mikhaljova, E. V. and S. I. Golovatch. 1981. Polymorphism in a new
species of genus Eoanerchodus (Diplopoda, Polydesmidae) from
the Far East. Zool. Zh. 60(8):1183-1189, figs. 1-4.
Miley, H u ^ H. 1927. Development of the male gonopods and life
history studies of a polydesmid millipede. Ohio J. Sci. 27:25-
41, pis. 1, 2, figs. 1-9.
Miyosi, Y. 1954. Beitrage zur kenntnis japanischer ityriopoden. 10
Aufsatz: Uber zv;ei neue arten von Diplopoda. Zool. Mag. To] 63:288-291, figs. 1, 2. ------. 1956. Beitrage zur kenntnis japanischer ntyriopoden. 18 Aufsatz: Uber eine neue gattung von Cryptodesmidae und eine neue unterart von Crvptops. Dobutsugaku zasshi. 61:339-340, figs. 1 - 2 . Morse, Max. 1902. Fyriapods from Vinton, Ohio. Bull. Ohio Acad. Sci. 2:187-188. Morakami, Yoshiteru. 1966. Postembryonic development of the common myriapoda of Japan. XXII. Three new species of millipeds. Zool. Mag. Tokyo. 75:94-97. figs. 1-3. Nelson, Gareth and Donn Rosen. 1981. Vicariance Biogeography. Columbia University Press. New York. 593pp. Newport, George. 1844. A list of the species of myriapods in the collection of the British Museum. Ann. Mag. Nat. Hist., ser. 1, 13:263-270. O'Neil, R. V. 1967. Niche segregation in seven species of diplopods. Ecol. (Brooklyn) 48:983. 185 Packard, Alpheus S. 1877. On a new cave fauna in Utah. Bull. U. S. Geol. Geogr. Surv. Terr. (Hayden). 3(1), art 10:157-169, figs. 5-10. ------1886. larval form of Polvdesmus canadensis. Alter. Nat. 4:651. Peck, Stewart and James Lewis. 1978. Zoogeography and evolution of the subterranean invertebrate faunas of Illinois and southeastern Missouri. National Speleol. Soc. Bull. 40:39-63. Pembelton E. F. and E. L. Blake. 1967. The fauna of Maze Cave. Missouri Speleo. 9(1):8-13. (not seen). Peters, E. H. 1954. Millipedes Dinloiulus londinensis [sic] leach and Polvdesmus serratus (Say) : Ontario. Can. Insect. Pest Rev. 32:135. (not seen). Platnick, Norman and Gareth Nelson. 1978. A method of analysis for historical biogeography. Syst. Zool. 27:1-16. Pocock, R. I. 1894. Chilopoda, Syitphyla, and Diplopoda from the Malay Archipelago, in Weber Zoologische Ergebnisse Reise in Niederlandisch Ost-Indien. 3:307-404, pis. 19-22. Ramsey, James M. 1966. Vast migrating armies of the millipede Pseudopolvdesmus serratus (Say) in the Dayton region. Ohio J. Sci. 66(3)-.339. Rosen, Don E. 1978. Vicarient patterns and historical explanation in biogeography. Syst. Zool. 27(2):159-188. Sager, Abram. 1856. Description of three myriapods, Proc. Acad. Nat. Sci. Philadelphia. 8:109. 186 Say, Thomas. 1821. Description of the ]%riapodae of the United States. J. Acad. Nat. Sci. Philadelphia. 2:102-114. Shear, William A. 1972. The milliped genus Bidentooon (Diplopoda, Polydesmida, Tridhopolydesmidae). Eroc. Biol. Soc. Washington. 85(42):489-492. figs. 1-5. ------. 1974a. North American cave millipeds. II. An unusual new species (Dorypetalidae) from southern California, and new records of Speodesmus tugaribuis (Trichpolydesmidae). Occasional Papers California Acad. Sci. 112:1-9. figs. 1-8. ------. 1974b. Millipeds (Diplopoda) from Mexican and Guatemalan Caves. Acc. Naz. Lincei, Prob. Att. Sci. Cult. 171(2): 240-305. ------. 1975. Millipeds (Diplopoda) from caves in l\fexico and Central America. Species and Records of Glomeridae, Cleidogonidae, Trichopetalidae, Fuhrmannodesmidae and Sphaeriodesmidae. Assoc. Mex. Cave Stud. Bull. 8:145-160, figs. 1-54. Shelley, Roland M. 1974. Polydesmid diplopods of the eastern Piedmont of North Carolina. J. Elisha Mitchell Sci. Soc. 90:111-112. ------. 1978. Millipeds of the eastern Piedmont region of North Carolina U.S.A. (Diplopoda). J. Nat. Hist. 12:37-79, figs. 1-72. Silvestri, Fillippo. 1898. Viaggio del dott. e. Festa nella Republics dell' Ecuador. XI. Diplopodi. Boll. Mus. Zool. Anat. Coitp. Univ. Torino. 13(324) :1-11, figs. 1-29. 187 ------. 1910. Descrizioni preliininari di navi generi di Diplopodi. I. Polydesmoidea. Zool. Anz. 35:357-364. Strasser, Karl. 1974. Diplopodi chdlognati della Sardegna. Frag. Ent. 10(3): 231-293, figs. 1-73. ------. 1941. Versuch eines Siphonophoriden-systems und geographisch-phylogenetische beurteilung der gonopod. Zool. Anz. 134:212-224. figs. 1-8. Wheeler, Quentin D. 1979. Slime mold beetles of the genus Anisotoma (Leiodidae) classification and evolution. Syst. Ent. 4:251- 309. Wiley, Edward 0. 1981. Riylogenetics. The theory and practice of phlogenetic systematics. John Wiley and Sons, New York. Williams, Eliot C. Jr. and Robert A. Hefner. 1928. The millipedes and centipedes of Ohio. Bull. Ohio Biol. Surv. 4(3):93-l46. figs. 1-26. Withrow, Charles P. 1978. Millipeds of Cabell County, West Virginia. W. Va. Acad. Sci. 38:48-50. Wood, Horatio C. 1865. The I'^'riapoda of North America. Trans. Amer. Hhilos. Soc., new series, 13(2) art. 7:137-248, pis. 1-3, figs. 1-61. Wray, David L. 1950. Insects of North Carolina. Second Supplement, North Carolina Dept. Agriculture, Div. Ent., Raleigh. 59pp. Youngsteadt N. W. & N. 0. 1978. A survey of some cave invertebrates from northern Arkansas. Arkansas Cave Studies. 1:1-13. (not seen). 188 APPENDIX A TABLES 189 Table 1. Systematics of the Family Polydesmidae FAMILY POLYDESMIDAE Subfamily Mastigonodesminae new status * Mastiqonodesmus Galliocookia Heterocookia Subfamily Polydesminae new status Archiopolydesmus * Polvdesmus * Jaxarbes * Kirqisdesmus * Scfaizoturanus * Schizomeritius * Turanodesmus * Usbekodesmus * Cretodesmus * Serradium * Pseudopolvdesmus Subfamily Scytonotinae new status Speodesmus Scvtonotus Utadesmus Bidentocfon Coronodesmus Speorthus Idahodesmus Subfamily Epanerchodinae new status * Epanerchodus Subgenus Prionomatis Huzichodus Niponchodus Nipponesmus Riuechodus Styqoerchodus Anthrochodus Krenoerchodus * Old World Taxa 190 Table 2. List of generic character states for the Family Polydesmidae. Character Plesionorphic J^xxnorphic 1 . 2nd Paranotum normal flabellately enlarged 2 . Involution Ability absent present 3. Gonocoxal Ventral normal with microsetae Surface 4. Sternal Width narrow wide 5. Length/Width Ratio low(8-23%) h i ^ (15-32%) 6 . Interantennal Width narrow wide 7. Antennal Lsngtli ^ o r t long 8 . Seminal Groove subterminal subterminal on Tibiotarsus or terminal 9. Cÿphopodal Shape elongate ovate 1 0 . Metazonite plain with tubercles 1 1 . Mandibular Shape quadrate round 1 2 . Ozodeme Location medial marginal 13. Ozopore Location anterior posterior or medial 14. Tergal Shape rectangular trapezoidal 15. Size Large present absent 16. Tibiotarsal Attachment ventral lateral 17. Coloration pigmented unpigmented 18. leg Modifications absent present 19. Macrosetae on legs absent present 2 0 . Paraproct Setation few hirsute 2 1 . Sternum III typical modified 2 2 . Syncoxostemal Hooks absent present 23. Claw Length long short 24. Biogeography Asia Holarctic Nearc 25. Segments Numbers 19 or 20 19 26. Lergth of Gonopodal short long Prefemur 27. Gnathochilarium quadrate rectangular 28. Syncoxostemum with absent present Lateral Spines 29. Claw evenly constricted tapered 30. Leg Sphaerotrichomes most only tibia and podoraeres tarsus 31. Collum Width smaller larger than head than head 32. Head Setae few-long numerous-micro 33. Leg Thickness thin robust 191 Table 2. (continued). Character Plesiomorphic jpomorphic 34. Prefemur Setation normal non-hirsute 35. Anterior Serrations absent present on Gonopods 36. Gonopodal Femur absent present with Setal Row 37. Tibiotarsus with absent present Caudalad Process 38. Hypoproctal Setae few numerous 39. Epiproct Setae little many 40. Tergal Size large small 41. Setae shape strai^t hooked 42. Peritremata narrow wide 43. Femur sirrple branched 44. Anterior Edge unflared flared of Collum 45. Ozopore Shape flush elevated 46. Dorsal Tubercle variable 1 0 , 1 0 , 8 Formula 47. Gonopod arcuate elorgate 48. Endomerite X-section round flat 49. Dorsal Setae Shape filiform clavate or peg 50. Paranotal Denticles typical cephalad 51. Tibiotarsus branched unbranched 52. Anterior Row of caudad cephalad Dorsal Setae 53. Sternal Modifications absent present 54. Dorsal Row Number 3 4 or 5 55. Tibiotarsus Setae Row 56. Medium Sized absent present 57. Anterior Terga cephalad laterad (2-4) Direction 58. Seminal Bulb absent present 59. Tergal Texture tuberculate smooth 60. Endomerite Size large small 61. iperture Shape ovate triangular 62. Clivus absent present 63. Seventh Antennomere w/o lobe w/ secondary lobe 64. Cyphqpodal Plate absent present 65. Segment Number 2 0 19 or 20 w w w w to tototo tototo to to to l->H H H HHH H H w to H o 00 •J <3\U1 fa. w to HO VD(X)'O m U1 W to H O VD 00 C\ U1 W to H •••*• • ••• I w o o O O O o o O o O H o O O O O O o O OOOOOO o o o H H H H g H g OOOOOOOOOOOOHHHHOHHHHHOHHHHHOOOOO & H 8, 1 HI a§ i 1 1 ^ OHHHHHHOHtOHHOOOHOHHOOHHHHHHHOOOOO n 4 OHHHHHHHHtOHHOOOOOl-’HOOHHHHHHHOOOOO p . OHHHHHHHHtOHHOOOHOHHOOHOH H O O O O O g I ill.? OHHHHHHHHWHHOHOHOHHOOHHHI-'HHHOOOOO g I _ r OHHHHHHHHWHHOHHOMHHOHHHHHHHHOOOOO g I IIP OHHHHHHHHtOHHHHHOHHHOHHHHHHHHOOOOO & HOOOOOOOOHHOOOOOOOOOOHHHHHHHOOOOO H ? HOOOOOOOOOOOOOOOOOOOOHHI-’HHHHOOOOO (D T , H H 8 ë tSJ UlrfiWt0HOV0(»^0>UliP*Wt'JHOVDCa^CT>UlitiWt0HOV003vJ(J>Ulrf:» • •••••••••••t» • • •• ••••••••••a I w oooooooooooooooooooooooooooooooo? ooooooooooooooooooooooooooooooooI OOOOOOOOOOOOOOOOOOOOOOOOOOOHHHHH Ü' I H OOOOOOOOOOOHOOOOOOOOOHHHHHHOOOOO H H I ' OOOOOOOOOOOOOOOOHH-'HHHOOOOHHOOOOO OOOOOOOOOOOOOHHHOOHHHOOOOHHOOHOO I OOOOOOOOOOOOHHHHOOOOHOOOOHHHOOHO OOOOOOOOOHHHOOHHHOOOHOOOOHHOOOOO IH OOOOOOOOOHHHOOHHOOOHHOOHOPHOOOOO OOOOHOHHPOOOOOOOOOOOOOOHOOOOOOOO g H H HHHHOOHHHOOOHOOOOOOOOOOHOOOOOOOO H ë CO Table 4. Manhattan distance matrix for the Family Polydesmidae. OTU Dor Mas Spe Sçy Uta Bid Cor Spr Ida Pol Epa Doratodesmidae 0 . 0 0 0 Mastigonodesminae 0.323 0 . 0 0 0 Speodesmus 0.508 0.369 0 . 0 0 0 Scvtonotus 0.492 0.415 0.262 0 . 0 0 0 Utadesmus 0.492 0.415 0.231 0.185 0 . 0 0 0 Bidentoaon 0.562 0.391 0.266 0.250 0.141 0 . 0 0 0 Coronodesmus 0.569 0.400 0.215 0.231 0 . 2 0 0 0.109 0 . 0 0 0 Speorthus 0.585 0.385 0.292 0.246 0.185 0.172 0.169 0 . 0 0 0 Idahodesmus 0.625 0,391 0.328 0.250 0.203 0.175 0.203 0.062 0 . 0 0 0 Polydesminae 0.308 0.292 0.385 0.369 0.400 0.500 0.477 0.492 0.500 Epanerochodinae 0.354 0.308 0.462 0.446 0.477 0.578 0.523 0.569 0.578 Table 5. I^tristic distance matrix for the Family Polydesmidae. OIU Dor Mas Spe Sçy Uta Bid Cor Spr Ida Pol Fÿa Doratodesmidae 0 . 0 0 0 Mastigonodesminae 0.354 0 . 0 0 0 Speodesmus 0.508 0.369 0 . 0 0 0 Scvtonotus 0.585 0.446 0.262 0 . 0 0 0 Utadesmus 0.554 0.415 0.231 0.185 0 . 0 0 0 Bidentoaon 0.656 0.516 0.328 0.281 0.172 0 . 0 0 0 Coronodesmus 0.662 0.523 0.338 0.292 0 . 2 0 0 0.109 0 . 0 0 0 Speorthus 0.667 0.538 0.354 0.308 0.215 0.172 0 . 2 0 0 0 . 0 0 0 Idahodesmus 0 . 6 6 8 0.547 0.359 0.312 0.203 0.175 0.203 0.062 0 . 0 0 0 Polydesminae 0.308 0.292 0.446 0.523 0.492 0.594 0.600 0.615 0.625 Epanerochodinae 0.354 0.338 0.492 0.569 0.538 0.641 0.646 0.662 0.672 S U1 Table 6. Hcsmoplasy matrix for the Family Polydesmidae. OIU Dor Mas Spe Sçy Uta Bid Cor Ida Pol Epa Doratodesmidae 0 . 0 0 0 Mastigonodesminae 0.031 0 . 0 0 0 Speodesmus 0.000 0.000 0.000 Scvtonotus 0.092 0.062 0 . 0 0 0 0 . 0 0 0 Utadesmus 0.062 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 Bidentoaon 0.094 0.125 0.062 0.031 0.031 0 . 0 0 0 Coronodesmus 0.092 0.123 0.123 0.062 0.000 0.000 0.000 Speorthus 0.092 0.154 0.062 0.062 0.031 0 . 0 0 0 0.031 0 . 0 0 0 Idahodesmus 0.062 0.156 0.031 0.062 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 Polydesminae 0 . 0 0 0 0 . 0 0 0 0.062 0.154 0.092 0.094 0.123 0.123 0.125 Epanerochodinae 0 . 0 0 0 0.031 0.031 0.123 0.062 0.062 0.094 0 . 0 0 0 0 . 0 0 0 ë a\ 197 Table 7. List of specific character states for the genus Pseudopolvdesmus. Character Plesiomorphy i^xomorphy 1. Process absent present 2. Medial Boss split coitplete 3. Sternum III (Female) modified unmodified 4. Cÿphopodal Plate absent present 5. Sternum 7 (Male) unmodified modified 6 . Tibiotarsal Attachment lateral medial 7. EMomerite small expanded 8 . Cÿphopod oval elongate 9. Tibiotarsus with absent present Terminal Setae 10. Process M 3 absent present 11. Cÿphopod Plate Margin small large 12. Sternal Setae few numei;T:'S 13. Process absent present 14. Paranotal Denticles large small 15. Epiproct normal d o w n t u m e d 16. Gonopodal Thickness thin thick 17. Process E 2 + 3 absent present 18. Process M 2 aksent present 19. Process E 2 absent present 2 0 . Habitat-Mountains generalized montane 21. Process E 3 absent present 22. Ant. Gonocoxae Edge sirrple multilobed 23. M 2 -E 2 Connection absent present 24. Posterior Sternum 7 small large 25. Legs long short 26. Size large small 27. Dorsal Setae absent present 28. Antennae long small 29. Cÿphopodal Plate angled straight 30. Habitat-Lowlands generalized lowlands 31. Process M 4 present absent 32. Dorsal Curvature arched sli^tly (>90%) arched 33. Cÿphopodal Setae evenly bunched spabed 34. Process Position near away from endomerite endomerite 35. Sternum II (Female) unmodified modified 36. Dorsum slightly flattened arched CO s & HHHHHrHrHrHHOOOOOOOrHOOHOOHOHHHHHHHHHOOH I HHHHHHHHHOOOOOOOHOOHOOHOHHHHHHHHHOHO rMrHrHrH«-!fHrHrHrHOOOrHOOOrHOOfHOOOOr“lrHrHrHpHrHrHrHOrHOO HHHHHHHHHHOOOOOOHOOHHOHHHOOOOOOOOOOO 8 HHHHHHHHHHOOHHOHHOOOOOHOOOOOOOOOOOOO I HHHHHHHHHOOHHOHHOHOHHOOOOOOOOOOOOOO &> HHHHHHHHHOOHHr-tOOOOHOOOOOOOOOOOOOOOO ij g HHHHHHHHHHOOHHHOOHHOOOOOOOOOOOOOOOOO il HHHHHHHHHHHHOOOOOOOOOOOOOOOOOOOOOOOO oooooooooooooooooooooooooooooooooooo CO £ #*#######$#*#######***####$ • ...... oH{v3r>'S*invot^co Table 9. Length and Width measurements of Pseudopolvdesmus species, males and females, with range, mean, standard deviation, and number measured. (pin-pinetorum; tal-tallulanus : era-erasus; can- canadensis; col-collinus; ser-serratus; cad-caddo; min- minor) . length Width Range Mean SD-1 Range Mean SD-l pin M 14.0-25.2 18.62 2.40 2.0- 4.3 3.12 0.50 N=112 F 13.6-25.6 18.68 2.67 2.1- 4.9 3.10 0.53 N=110 tal M 18.1-26.5 23.27 3.13 3.5- 4.8 4.28 0.52 N=33 F 17.6-26.8 23.25 2.95 3.5- 4.3 3.95 0.49 N=20 era M 15.8-24.0 20.38 3.27 2.6- 4.4 3.62 0.47 N=77 F 17.8-31.8 19.57 3.92 2.8- 3.7 3.20 0.50 N=ll can M 11.8-27.3 22.79 2.88 2.1- 5.0 4.03 0.53 N=100 F 15.5-28.6 21.15 2.92 2.6- 4.8 3.74 0.52 N=42 col M 13.4-24.7 19.73 2.57 2.7- 4.8 3.92 0.85 N=22 F 12.9-25.3 19.43 2.39 2.6- 4.6 3.60 0.73 N=17 ser M 16.5-32.0 23.86 4.88 2.9- 5.6 4.14 0.43 N=250 F 13.2-29.1 21.50 2.82 2.3- 5.1 3.72 0.45 N=250 cad M 7.5-13.3 10.21 0.78 1.0- 1.8 1.42 0.21 N=14 F 7.6-12.9 9.75 0.74 1.1- 1.7 1.37 0.21 N=13 m m M 8.8-12.5 10.34 0.80 1.1- 1.8 1.34 0.20 N=17 F 9.1-12.7 10.78 0.92 1.1- 1.6 1.34 0.14 N=14 200 Table 10. W/L (width/length) and WL (width X length) measurements of Pseudopolvdesmus species, males and females, with range, mean, standard deviation, and number measured, (pin- pinetorum; tal-tallulanus; era-erasus; can-canadensis; col- collinus; ser-serratus; cad-caddo; min-minor). W/L W L Range Mean SD-1 Range Mean SD-1 pin M 10.6-20.5 16.45 1.51 12.8-108.4 57.65 16.77 N=112 F 13.3-20.7 16.60 1.32 28,6-125.4 59.37 18.68 N=110 tal M 17.0-20.1 18.39 1.05 63.2-123.6 99.60 23.41 N=33 F 15.2-17.9 16.99 1.13 80.5-101.2 91.84 18.31 N=20 era M 15.9-20.4 17.66 1.21 40.3- 99.7 73.37 19.75 N=77 F 15.3-17.9 16.35 1.17 49.0- 80.0 62.62 18.82 N=ll can M 15.4-20.1 17.76 1.09 24.8-132.2 93.77 21.31 N=100 F 16.2-20.3 17.60 1.16 39.7-135.6 80.89 21.34 N= 42 col M 13.2-19.7 16.95 1.22 36.8-124.4 78.45 20.85 HN22 F 14.4-19.2 17.15 1.09 36.5-118.7 68.75 20.46 N=17 ser M 12.2-21.3 17.31 1.04 38.0-164.8 99.42 19.42 N=250 F 13.6-22.0 17.90 1.72 38.3-147.3 86.99 23.22 N=250 cad M 10.2-17.4 13.04 1.42 9.4- 24.7 15.22 2.42 N=14 F 9.8-16.8 12.87 1.32 9.2- 21.1 14.37 2.79 1^13 min M 10.7-16.2 12.95 1.68 10.2- 22.4 13.92 2.81 N=17 F 9.7-14.9 12.45 1.25 11.1- 20.3 14.59 2.36 N=14 201 Table 11. Distance from telqpodite tip to base of mesial and ectal (Ei_4 ) processes as a percentage of total telopodite length. M 4 M 3 M 2 Ml pinetorum 11.2-16.0 20.1-23.9 52.3-60.0 tallulanus 9.4-12.4 20.0-24.9 49.9-55.3 erasus 9.7-12.4 17.1-21.9 52.8-58.1 canadensis 2.3- 5.7 11.1-15.6 49.8-55.8 collinus 3.9- 7.2 - —--- -—— 24.4-28.9 54.0-61.9 serratus 10.2-15.7 34.9-41.0 52.7-60.0 paludicola ------2 0 .1 - 2 2 . 2 caddo 8.3-11.2 ------27.3-32.3 51.9-55.4 minor ------18.4-21.2 35.7-39.8 55.0-60.1 E 4 % E2 El pinetorum 6 .8 — 8.4 13.1-19.5 24.9—30.2 ——----- tallulanus 7.2-10.1 20.1-25.7 30.2-35.4 39.6-45.1 erasus 6.2- 9.8 ------28.0-32.4 37.6-43.4 canadensis 2.6- 5.4 2 0 .] 27.3 35.1-40.6 collinus 4.8- 7.8 18.S 25.2 38.3-42.5 serratus 6 .0 -1 0 . 1 33.8-40.3 ------ paludicola 5.0- 6.1 - —----- caddo 7.2- 9.8 ------32.1-44.6 minor 5.8- 9.5 ------30.1-34.6 Table 12. Manhattan distance matrix for the species in the genus Pseudonolvdesmus. OKI Pol pin tal era can col ser pal min cad Polvdesmus 0.000 pinetorum 0.314 0.000 tallulanus 0.400 0.143 0.000 erasus 0.400 0.143 0.057 0.000 canadensis 0.457 0.200 0.114 0.171 0.000 collinus 0.429 0.229 0.200 0.200 0.086 0.000 serratus 0.400 0.257 0.343 0.286 0.343 0.257 0.000 paludicola 0.625 0.406 0.438 0.375 0.438 0.469 0.281 0.000 minor 0.629 0.543 0.629 0.600 0.486 0.514 0.343 0.216 0.000 caddo 0.629 0.543 0.629 0.600 0.486 0.514 0.343 0.281 0.114 0.000 to 8 Table 13. Patristic distance matrix for the species in genus Pseudopol ydP-gffm . OTU Pol pin tal era can col ser pal min cad Polvdesmus 0.000 pinetorum 0.314 0.000 tallulanus 0.400 0.143 0.000 erasus 0.400 0.143 0.057 0.000 canadensis 0.457 0.200 0.171 0.171 0.000 collinus 0.486 0.229 0.200 0.200 0.086 0.000 serratus 0.400 0.257 0.343 0.343 0.400 0.429 0.000 paludicola 0.688 0.531 0.625 0.625 0.688 0.719 0.281 0.000 minor 0.800 0.657 0.743 0.743 0.800 0.829 0.400 0.216 0.000 caddo 0.743 0.657 0.686 0.686 0.743 0.771 0.343 0.218 0.114 0.000 to wo Table 14. Homoplasy matrix for the species in the genus Pseudopolvdesmus. OTU Pol pin tal era can col ser pal min cad Polvdesmus 0.000 pinetorum 0 . 0 0 0 0 . 0 0 0 tallulanus 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 erasus 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 0 . 0 0 0 canadensis 0.000 0.000 0.057 0.000 0.000 collinus 0.057 0.000 0.000 0.000 0.000 0.000 serratus 0.000 0.000 0.000 0.057 0.057 0.171 0.000 paludicola 0.062 0.125 0.188 0.250 0.250 0.250 0.000 0.000 minor 0.171 0.114 0.114 0.171 0.171 0.229 0.057 0.000 0.000 caddo 0.114 0.057 0.057 0.114 0.114 0.286 0.000 0.000 0.000 0.000 to g APPENDIX B FIGURES 205 206 Figure 1. Figure 2 •v. Figure 3. Figures 1-3. Dorsum, midbody, dorsal view, male. 1. Cerastelachvs cavemicola. Thailand, Gua Tanan, Biserat; 2. Mastiqonodesmus viqnai. Italy, Sardinia, Carbonia; 3. Epanerchodus bidens Japan, Yamanashi Eref., Saiko- komori-ana Cave. 207 Figure 4. ^^irrrri Figure 5. Figure 6 . Figures 4-6. Dorsum, midbody, dorsal view, maie. 4. Speodesmus echinourus. Texas, Kerr Co.; 5. Scvtonotus qranulatus. Ohio, Fairfield Co.; 6 . Utadesmus henriensis. Utah, Henry Mountains. 208 Figure 7. Figure 8 . / y Y V v Y r V Y Y y ' -f t y r Y r y r r v v r \ \ /,A y r II t jr r ^ y y v r ry t y i r r V y y y v y y M Figure 9. Figures 7-9. Dorsum, midbody, dorsal view, maie. 7 . Bidentogon helferorum. California, Mendocino Co.; 8 . Coronodesmus bituberculatus. California, Alameda Co.; 9 . Speorthus tuqanbius. New Mexico, Eddy Co. 209 Figure 10. Figure 11. :l‘î Figure 12. Figures 10-12. Dorsum, dorsal view, male. 10. idahodesmus dentatus. Idaho, Latah Co. ; 11. Polvdesmus inconstans, Ohio, Franklin Co.; 12. Pseudopolvdesmus serratus. West Virginia, Cabell Co. 210 Figure 13. Figure 15. Figures 13-15. Right gonopod, lateral view. 13. Cerastelachvs cavemicola Thailand, Gua Tanan, Biserat; 14. Mastiqonodesmus vionai It^y, Sardinia, Carbonia; 15. Epanerchodus bidens Japan, Yamanashi Bref., Saiko- komori-ana Cave. 211 Figure 17. Figure 16. Figure 18. Figures 15-18. Left gonopod of Soeodesmus echinourus. Texas, Kerr CO.; 16. medial view; 17. lateral view; 18. ventral view. 212 i i s i muiu cf Figure 19. Figure 20. Figures 19-20. Dorsal view. 19. Pseudopolvdesmus erasus. male; 20. Scvtonotus qranulatus. Ohio, Fairfield CO., male and female. 213 Figure 21. Figure 22. Figure 23. Figures 21-23. R i ^ t gonqpod of Scvtonotus qranulatus. Ohio, Fairfield Co.; 21. medial view; 22. lateral view; 23, ventral view. 214 Figure 24. Figure 25. Figure 26. Figures 24-26. R i ^ t gonopod of Utadesmus henriensis. Utah, Henry Mountains; 24. medial view; 25. lateral view; 26. ventral view. 215 Figure 27. Figure 28. Figure 29. Figures 27-29. R i ^ t gonopod of Bidentogon helferorum. California, Medocino Co.; 27. medial view; 28. lateral view; 29. ventral view. 216 Figure 30. Figure 31. Figure 32. Figures 30-32. Ri ^ t gonopod of Coronodesmus bituberculatus, California, Alameda Co. ; 30. medial view; 31. lateral view; 32. ventral view. 217 Figure 33. Figure 34. Figure 35. Figures 33-35. Right gonopod of Speorthus tuoanbius. New Mexico, Eddy Co.; 33. medial view; 34. lateral view; 35. ventral view. 218 Figure 37. Figure 36. Figure 39. Figure 38. Figures 36-39. Right gonopod of Idahodesmus dentatus. Idaho, latah CO.; 36. medial view; 37. lateral view; 38. ventral view; 39. sternum III, posterior view, female. 219 Figure 41. Figure 40. / Figure 42. Figures 40-42. R i ^ t gonopod of Polvdesmus inconstans. Ohio, Franklin Co.; 40. medial view; 41. lateral view; 42. ventral view. Mast Spe Scy Uta Ida Spr Bid Cor Epan Poly 60 42 36,47 62-651 49 53 [35 38,53 46 541 52 49 49 45 46 39 40-441 47 50,51 17“ 21 |\|33~38l 57-61 13 145 53 2339,55 22-32 r J51 Doratodesmidae 11-16 1 -5 6-10 Figure 43. Cladogram for the Family Polydesmidae including the subfamilies and the genera of Scytonotinae (Mast-Mastigonodesminae; Epa-Epanerchodinae; Pol-Polydesminae; Spe- Speodesmus; Scy-Scvtonotus; Uta-Utadesmus; Ida-Idahodesmus; Spr-Speorthus; Bid- Bidentoqon; Cor-Coronodesmus. to CO o 221 Figure 44 Figure 45. Figures 44-45. Head and tergites 1-4, dorsal view. 44. Scvtonotus qranulatus. Ohio, Fairfield Co.; 45. Pseudopolvdesmus serratus. West Virginia, Cabell Co. # Figure 46 Figure 47 Figures 46-47. Diplosegment, lateral view, laidbody. 46. Pseudopolvdesmus serratus. West Virginia, Cabell Co.; 47. Scvtonotus qranulatus, Ohio, Fairfield Co. to ro to 9 Figure 48. Figure 49. Figures 48-49. Diplosegment, cross section, midbody. 48. Pseudopolvdesmus serratus. West Virginia, Cabell CO.; 49. Scvtonotus qranulatus. Ohio, Fairfield CO. M-male, F-female. to CO w 224 Figure 50. Figure 51. Figure 52. Figures 50-52. Dorsal setae, lateral view. 50. Pseudopolvdesmus caddo. Louisiana, Iberville Par.; 51. Bidentogon helferorum. California, Mendocino Co.; 52. Speodesmus echinourus. Texas, Kerr Co. 225 Figure 53. Figure 54. Figures 53-54. Legs, posterior view, male. 53. Utadesmus henriensis. Utah, Henry Mountains, right, third leg; 54. Scvtonotus qranulatus. Ohio, Fairfield Co., ri^t posterior leg, 13th segment. 226 Figure 55. Figure 56. Figures 55-56. Claw, posterior view. 55. Pseudopolvdesmus canadensis. Tennessee, Caitpbell Co.; 56. Scvtonotus qranulatus. Ohio, Fairfield Co. 227 SC Figure 57. Figure 58. Figures 57-58. Antennae, anterior view. 57. Pseudopolvdesitius serratus. West Virginia, Cabell Co.; 58. Scvtonotus qranulatus. Ohio, Fairfield Co. bs-bacilliform setae, SC-sensory cones. 228 Figure 59. Figure 60. Figures 59-60. Midbocty sternum, ventral view. 59. Pseudopolvdesmus serratus. West Virginia, Cabell Co.; 60. Scvtonotus qranulatus. Ohio, Fairfield Co. 229 Figure 61. Figure 62. Figures 61-62. Posterior sternum and coxa of 7th segment, posterior view, male; 61. Pseudopolvdesmus canadensis. Tennessee, Caitpbell Co.; 62. P. minor. Arkansas, Desha Co. 230 ' ' I# Figure 63. Figure 64. Figures 63-64. Syncxjxostemum, posterior view. 63. Pseudopolvdesmus serratus. West Virginia, Cabell Co.; 64. Scvtonotus qranulatus. Ohio, Fairfield Co. 231 Figure 65. Figure 66. Figures 65-66. Sternum III, posterior view, female. 65. Polydesmus inconstans. Ohio, Franklin Co.; 66. Pseudopolydesmus serratus. West Virginia, Cabell Co. 232 !/, 'i' ; Figure 67. Figure 68. Figures 67-68. Cyphopod, lateral view. 67. Pseudopolydesmus serratus. West Virginia, Cabell Co. ; cp-cyphopod plate, v-valve, o-operculum; 68. Scvtonotus qranulatus. Ohio, Fairfield Co. cl ta 233 sp Figure 69. Midbody leg of Pseudopolydesmus serratus. West Virginia, Cabell Co., anterior view, male and female, semidiagrammatic, setae omitted. Double-headed arrows indicate length as measured, c-coxa, tr-trochanter, prf- preferaur, f-femur, ti-tibia, ta-tarsus, cl-claw, sp- sphaerotrichomes. 234 2!; % Figure 71. Figure 70. Figure 73. Figure 72. Figures 70-73. Ri^t gonopod of Pseudopolydesmus canadensis, Tennessee, Canpbell Co., male; 70. mesal view; 71. lateral view; 72. ventral view; 73. in situ. 235 Figure 74. Figure 75. Figure 75. Figure 77. Figures 74-77. Dorsum, dorsal view, midbody- 74. Pseudopolydesmus pinetorum. Arkansas, Madison Co.; 75. P. tallulanus. North Carolina, Jackson Co.; 76. P. canadensis. Tennessee, Cairpbell Co.; 77. P. caddo. Louisiana, Iberville Par. 236 Figure 78. V/ Figure 79 Figures 78-79. Female sternum III, ventral view. 78. Pseudopolydesmus serratus. West Virginia, Cabell Co. ; 79. P. minor. Arkansas, Desha Co. 237 Figure 80. Figure 81. Figures 80-81. E$)iproct, lateral view. 80. Pseudopolydesmus pinetorum. Arkansas, Madison Co. ; 81. P. serratus. West Virginia, Cabell Co. 238 Figure 82. Figure 83. 1 Figure 84. Figure 85. Figures 82-85. Right gonopod of Pseudopolydesmus spp., Group canadensis, lateral view. 82. P. pinetorum. Arkansas, Madison Co.; 83. P. tallulanus. North Carolina, Jackson Co.; 84. P. erasus. Tennessee, Anderson Co.; 85. P. collinus. West Virginia, Cabell Co. 239 Figure 86. Figure 87. Figure 88. Figure 89. Figures 86-89. Riÿit tibiotarsus of Pseudopolydesmus spp., Groip canadensis, medial view. 86. P. pinetorum. Arkansas, Madison Co.; 87. P. tallulanus. North Carolina, Jackson Co.; 88. P. erasus. Tennessee, Anderson Co.; 89. P. collinus. West Virginia, Cabell Co. 240 Figure 90 Figure 91. Figure 92. Figure 93. Figures 90-93, Right tibiotarsus of Pseudopolydesmus spp., Group canadensis, ventral view. 90. P. pinetorum. Arkansas, Madison Co. ; 91. P. tallulanus. North Carolina, Jackson Co.; 92. P. erasus. Tennessee, Anderson Co.; 93. P. collinus. West Virginia, Cabell Co. 241 Figure 94. Figure 95. ?>- Figure 96. Figure 97. Figures 94-97. Right gonopod of Pseudopolydesmus spp., Group serratus, lateral view. 94. P. serratus. West Virginia, Cabell Co. ; 95. P. paludicola. Virginia, Princess Anne Co.; 96. P. caddo. Louisiana, Iberville Par. ; 97. P. minor. Arkansas, Desha Co. 242 Figure 98. Figure 99. Figure 100. Figure 101. Figures 98-101. R i ^ t tibiotarsus of Pseudooolvdesmus spp.. Group serratus, medial view. 98. P. serratus. West Virginia, Cabell Co.; 99. P. paludicola. Virginia, Princess Anne Co.; 100. P. caddo. Louisiana, Iberville Par. ; 101. P. minor. Arkansas, Desha Co. 243 Figure 102. Figure 103. Figure 104. Figure 105. Figures 102-105. Ri«ÿit tibiotarsus of Pseudopolydesmus ^p.. Group serratus, ventral view. 102. P. serratus. West Virginia, Cabell Co.; 103. P. paludicola. Virginia, Princess Anne Co.; 104. P. caddo. Louisiana, Iberville Par. ; 105. P. minor. Arkansas, Desha Co. 244 Figure 106. Figure 107. Figure 108. Figure 109. Figure 110. Figures 106-110. Right cyphopod of Pseudopolydesmus spp., Group canadensis, lateral view. 106. P. pinetorum. Arkansas, Ifedison Co.; 107. P. tallulanus, North Carolina, Jackson Co.; 108. P. erasus. Tennessee, Andersen Co.; 109. P. canadensis. Tennessee, Cairpbell Co. ; 110. P. collinus. West Virginia, Cabell CO. 245 Figure 111. Figure 112. Figure 113. Figure 114. Figure 115. Figures 111-115. R i ^ t cyphopod of Pseudopolydesmus spp., Group canadensis, ventral view. 111. P. pinetorum. Arkansas, Madison Co.; 112. P. tallulanus. North Carolina, Jackson Co.; 113. P. erasus. Tennessee, Anderson Co.; 114. P. canadensis. Tennessee, Cairpbell Co. ; 115. P. collinus. West Virginia, Cabell Co. 246 Figure 116. Figure 117. Figures 116-117. Right cyphopod of Pseudopolydesmus spp., Group serratus, lateral view. 116. P. caddo. Louisiana, Iberville Par. ; 117. P. minor. Arkansas, Desha Co. 247 Figure 118. Figure 119. Figure 120. Figures 118-120. Right cyphopod of Pseudopolydesmus spp., Group serratus, ventral view. 118. P. caddo. Louisiana, Iberville Par. ; 119. P. minor. Arlcansas, Desha Co.; 120. P. serratus. West Virginia, Cabell Co. 248 pin era can col ser mm cad / 13 ]20 \ 34,351 136 31-33 22 2 4 -3 0 Polydesmus 2 0 - 2 3 17 2-8 Figure 121. Cladcgram for the genus Pseudopolydesmus. (pin- pinetorum; tal-tallulanus; era-erasus; can-canadensis; col-collinus; ser-serrratus; pal-paludicola; min-minor; cad-caddo. Length (ntn) 10 15 20 25 30 35 pinetonmi H F talliilanua M F J- erasus M F canadensis M F collinus M F serratus M F caddo . M F minor H - c F Figure 122. Graph of length measurements of Pseudopolydesmus species, male and female. Range- horizontal line, means-vertical line, one standard deviation-rectangle. to VD Width (mm) 0.0 0.5 1.0 1.5 2.0 2:5 3.0 3.5 4.0 4.5 5.0 5.5 6.0 pinstortnn M F tallulanus M F erasus H F canadensis M F collinus M F c = ± serratus M F M F caddo M F - 0 = H Figure 123. Graph of width measurements of Pseudopolydesmus species, male and female. Range- horizontal line, means-vertical line, one standard deviation-rectangle. NJ ui o ■Width/length (%) 9 10. 11 12 ' 13 14 15 16. 17 18 19 20 2 1 22 23 pinetorum ; ' M -c F tallulanus H F erasus . M F canadensis M F -c collinus M F serratus M -c= t; F minor .M F caddo M F Figure 124. Graph of W/L measurements.of Pseudopolydesmus species, male and female. Range- horizontal line, means-vertical line, one standard deviation-rectangle. Length X Width (mm^) 20 40 60 80 100 120 140 160 180 pinetorum M F tallulanus M ■ F erasus M F canadensis M F collinus M F serratus M F minor M - C | D - F caddo M ■ 1— j— F 4 Figure 125. Graph of WL measurements of Pseudopolydesmus species, male and female. Range- horizontal line, means-vertical line, one standard deviation-rectangle. tv) 8 Mesial Processes 10 15 20 25 30 35 4 0 45 50 55 GO Ot pinetorum tallulanus ,______,_ i___ , I ^ I ' ÿ ' erasus I— =— ( e ^ canadensis collinus I— : — I------I------z ------1 I - 4 ? '1 serratus ;---- :— paludicola minor______,_ 2 caddo ^ 2 ■ ' 1 • Ect.al P'rocessps 60 pinetorum tallul.tallulanus ,____ , ' 3 '^ 2 ^ erasus 4 3 2 '_____ ^ ______1 canade:canadensis , , ^ ^ ,_____ L, collinus 4' 2 * 1 i serratus paludicola minor caddo Figure 126. Graph of distances from telopodite tip to bases of mesial and ectal processes as a percentage of total telopodite length. to U) APPENDIX C MAPS 254 ^ : ■ ■ Map 1. Distribution of families Polydesmidae and Doratodesmidae. Doratodesmidae ( — ) ; Polydesmidae ( ). to oi Ul 256 Map 2. Distribution of Scytonotinae. Scvtonotus Bidentoqon ( □ ) ; Utadesmus ( A ) ; Speodesitius ( |H ) ; Speorthus { — )> Coronodesïïius ( @ ) ; Idahodesitius ( ® ) . 257 Map 3. Distribution of Pseudopolvdesimis. tv to Map 4. Distribution of Pseudooolvdesmus pinetorgn. s 259 9 m 9 Map 5. Distribution of Pseudopolvdesmus tallulanus ( g ) and erasus ( ® ) • 260 Map 6. Distribution of Pseudopolvdesmus canadensis. 261 Map 7. Distribution of Pseudopolvdesmus collinus. 262 -H Map 8. Distribution of Pseudooolvdesmus serratus. Ifep 9. Distribution of Pseudooolvdesmus paludicola ( @ ), caddo ( a ), and minor ( H ). sto APPENDIX D IDCALITY DAITA 264 265 This is a list of localities and specimens ejraiained. Those marked with a * are literature sites where the identifications are assumed to be correct. Distances are copied and have not been converted to metric units. The following abbreviations were used in the list: ^ p r o x i mately- circa Between- btwn County- Co. County Road- CR. Creek- Ck. Female- F Great Smoky Mountain National Park- GSMNP Junction- jet Juvenile- J Male- M Mountain- Mt. Near- nr Parish- Par. River- R. Road- Rd. State Forest- S. F. State Park- S. P. State Route- SR. United States Forest Service- USFS All relative directions are abbreviated (ex. : north of Clarkesville as N. Clarkesville). Pseudopolvdesmus pinetorum ALABAMA: Walker Co., Jasper, 800 Airport Rd., IM, IV-28-1978, B. R. Wall Jr, ( N Œ H ) . ARKANSAS: Benton Co., Vough, IM, X-13-1962, H. M. Bevel, (FSAC) ; Vough, IF, XI-4-1960, H. M. Bevel, (FSAC) ; Logan Cave, 2M, 1-27-1958, T. Barr, (FSAC); Siloam Springs, IF, III-1-1955, (FSAC); 266 Bella Vista, 3M IF, IV-13-1962, H. Hart, (FSAC). *Clark Co., Arkadelphia, (Bollman, 1888). Columbia Co., Magnolia, 2M 3F, XI-26- 1961, R. Rogers, (FSAC). Dallas Co., E. of Princeton, 2M, VII-1-54, N. B. Causey, (FSAC). Faulkner Co., Conway, IF, XII-24-1952, M. A. Jackson, (FSAC) ; Bridge on Warren Ck., Rt. 64, IF, XI-18-1966, M. Hite, (FSAC). Franklin CO., Spirit Ck. Drainage, Sec.7 R28W, T13N, 8M 8F 4J, XI-6-1958, J. R. Preston, (FSAC) ; Shores Lake Area, IM 3F, XI-20-1958, J. R. Preston, (FSAC). Garland Co., *llmi. W. Hot Springs, IM, III-21-1938, K. P. Schmidt, (Chamberlin, 1952) ; US Rt. 70, 13.3mi. E. jet US Rt. 70B, IM IF, VI-23-1983, C. P. Withrow, (CEWC). Independence Co., Batesville, 19M 4F 22J, IX-X-1959, O. D. Brown, (FSAC); 1.3mi. W. Cushman, vicinity Blowing Cave, IM, III-5- 1973, B. M. Blaney, (FSAC). Izard Co., Mount Pleasant, 2M 2F, XII-2 3-1954, N. B. Causey, (FSAC). Jackson Co., Elmo, IM, IV-26-1952, (FSAC). Jefferson Co., 2mi. E. Pine Bluff, 2M, X-1954 & V-VI-1955, Kirkwood, (FSAC). *Iawrence Co., IM, IV-3-1937, K. P. Schmidt, (Chamberlin, 1952). Madison Co., St. Paul, IF, X-24-1954, N. B. Causey, (FSAC) ; Huntsville, IF, V-19-1962, C. Caby, (FSAC); Withrow Springs S. P., IM IF, VI-22-1983, C. P. Withrow, (CEWC). Montgomery Co., lOmi. NW. Langley, AR at Albert, IM IF, 1979, H. W. Robinson, (NCMH) ; IM, VIII-25-1950, N. B. Causey, (FSAC). Nevada Co., Jackson Township, 2M IF, XII-30-1954, R. Delaney, (FSAC). Newton Co., SR. 59, 2.9mi. S. Harrison, 2F U , VI-22-1983, C. P. Withrow, (CEWC); US Rt. 74, 12.0mi. E. jet SR. 21, IM IF, VI-22-1983, C. P. Withrow, (CEWC); Glenwood, 3M, 1-8-1954, N. B. Causey, (FSAC) ; IM, VIII-25- 1950, N. B. Causey, (FSAC). North Crawford Co., 0.75mi. S. No. 12 Fire Tower, 2M, X-28-1958, J. R. Preston, (FSAC). Pike Co., Mena, Rich M t . , IF, Redman, (FSAC) ; Murfreeboro, 3M, IV-15-1953, W. S. Carter, (FSAC) ; Bear Ck., IV-13-1953, IM, N. B. Causey, (FSAC). EOlk Co., SR. 88, l.Omi. W. Queen Wilhelmina S. P., 2M IF, VI-20-1983, C. P. Withrow, (CEWC). Prairie Co., XI-1958, IF, (FSAC). Scott Co., 3.0 mi. E. Woldrove, U , IV-4-1961, D. Combs, (FSAC). Searcy Co., IM, VIII-24-1950, N. B. Causey, (FSAC) ; *cave, (Youngsteadt & Youngsteadt, 1978). Stone Co., W. of Mt. View, IM IF, IV-26-1952, N. B. Causey, (FSAC). Washington Co., Cove Ck. Valley, 4M 4F, IX-1- 1956, M. Hite, (NCMH) ; IM IF U , XII-1958, G. Ogden, (FSAC) ; Habberton, 3M, IV-4-1953, B. Johnson, (FSAC) ; Fayetteville, IM, X- 30-1962, F. Clayton, (FSAC); Fayetteville, Mt. Kessler, IF, V-4- 1955, (FSAC) ; (Cove Ck. Valley, IM 3J, 1st week 11-1955, Mr. & Mrs. 0. Hite, (FSAC) ; 7M 14F, Spring 1956, (FSAC) ; IM IF U , XII-1958, G. Ogden, (FSAC); Clear Ck., IF, 11-14-1955, Hastings, (FSAC). *Caves, (McDaniel & Smith, 1976). ILLINOIS; Cjhairpaign Co., SR. 49, 3.8mi. S. Odgen at US Rt. 150, IM 2F U , VI-14-1983, C. P. Withrow, (CPWC). *Macoupin Co., (Causey, 1952a). Madison Co., Alton, 4M 6F, X-1963, J. H. (Serard, (FSAC). *Mason Co., (Causey, 1952a). *EUlaski Co., (C^ausey, 1952a). Randolph Co., btwn Modoc and Roots, IM, IV-14-1936, K. P. Schmidt, (USNM), holotype of Polvdesmus modocus. Tazewell Co., US Rt. 74, 13.6mi. E. SR. 121, rest area, IM IF, VI-14-1983, C. P. Withrow, (CEWC). *Cave, (Peck & Lewis, 1978). IOWA: ^ p a n o o s e Co., jet SR. T30 & SR. 5, 4M 2F, C. P. Withrow, (CEWC). 267 KANSAS: Marshall Co., Marysville, 3M 2F, VIII-9-64, I. Schroller, (FSAC). *Riley Co., IV-2-1952, L. 0. Warren, (Causey, 1955). KENIUCKY: *Fayette Co., Lexington, III-17-1922, A. Stone, University of Kentucky, (Causey, 1955). *Jefferson Co., Louisville, IV-4-1922, F. E. Merriman, Universily of Kentucky, (Causey, 1955). LOUISIANA: Ascension Par., Gonzales, IF, V-11-1954, H. S. Eybas, (EMNH). Beauregard Par., DeRidder, 2J, VII-29-1969, H. H. Rhame, (FSAC). Bienville Par., SR. 9, 0.5itii. N. Lucky, IM 6J, VI-27- 1983, C. P. Withrow, (CPWC). Caddo Par., Shreveport, 3M, lLL-24- 1962, N. B. Causey, (FSAC). Shreveport Fair Park H. S., IM, 1-25- 1969, D. C. Marizot, (FSAC). Caldwell Par., 6.3mi. ENE. Colombia Heists on SR. 4, 2M, IV-3-1966, R. E. Tan^, (FSAC). Catahoula Par., 6.5mi. SSE. Aimwell, SR. 126, IM IF, ILI-27-1966, R. E. Tandy, (FSAC). *Claibome Par. (Causey, 1963). Evangeline Par., Boggs Bayou, 2.7mi. W. jet SR. 106 & 13, 6M IF, R. E. Tanc^, (FSAC) ; Lake Chicot, IM, IIL-20-1965, M. Kordisch, (FSAC) ; 2.5itd. S. Chicot S. P. entrance, 2M IF, 1-30-1968, E. D. Keiser, (FSAC). Grant Par., Dry Prong, U , X-18-1953, H. S. Pitas, (LMNH). Iberia Par. Avery Id., IM, IV-27-1965, R. E. Tanc^, (FSAC). Jackson Par., 1.5mi. N. Clay, IM, IV-12-1936, L. Babricht, (USNM), holotype of Polvdesmus paroicus. La Salle Par., 10.3 mi. SSW. Midway on SR. 772 & SR. 8, Trout Ck., IM, IV-2-1966, R. E. Tandy, (FSAC); La Salle, IM 2F, ILI- 27-1966, R. E. Tandy, (FSAC); Jenks, 3M 4F, (FSAC). *Lincoln Par., (Causey, 1963). Livingston Par., Springfield, IM 2F 16J, XII-7-1963, Harman, (FSAC). *Morehouse Par., nr Miller Landing, IM, 1909, C. B. Moore, (Chamberlin, 1947); Qiemin-a-Haut S. P., IM, IIL-15-1967, J. Cooper, (WASC) ; SR. 142, 2.1mi. S. Arkansas line, 4M 2F, III-16- 1967, J. & M. cooper, (NCMH). Natchitoches Par., 2mi. S. Saline, 2M 2F, IV-12-1936, L. Hubricht, (USNM), holotype of Polvdesmus natchitoches; Dam N. S. C., 3M 2F U , 1-20-1955, C. Morehead, (FSAC) ; 3M 3F, 11-28-1956, (FSAC) ; Grand Ecore, 3M, 11-15-1955, R. Hernandez, (FSAC). Ouachita Par., Monroe, 1410 Forsythe Ave., 2M IF, XII-31-1974, M. R. Cooper, (NCMH); IM, IV-26-1976, M. R. & J. E. Cooper, (NCMH). St. Charles Par., Norco, IM IF, VIII-31-1944, H. S. Dybas, (EMNH). St. Martin Par., SR. 70, St. Maiy Par. line, IM, IV- 7-1967, R. E. Tandy, (FSAC). MISSISSIPPI: Attala Co., SR. 35, 13.7mi. S. US Rt. 51, U , VII-2-1983, C. P. Withrow, (CEWC). Marshall Co., 2mi. E. Slayden, mixed woods, 4M IF, 11-27-1961, L. Hubricht, (RIHC). Neshoba Co., SR. 16, 20.6mi. E. SR. 35, 14J, VI-31-1983, C. P. Withrow, (CEWC). Panola Co., Como, 3M 3F, XI-30-1963, W. D. Longest, (FSAC). Wilkenson Co., circa 1 air mi. SE. Fort Adams, IM IF, III-30-1974, D. A. Rossman, (FSAC). MISSOURI: Adair Co., Thousand Hills S. P., IM 4F, VI-17-1983, C. P. Withrow, (CEWC). Barry Co., Brook Cave, 3mi. NNE. Shell Knob, IM IF, III-15-1979, J. E. Gardner, (NCMH) ; 5.25mi. NW. Eagle Rock, Trailside Spring Cave, IF, V-6-1982, J. E. Gardner, (CEWC). Roaring R. S. P., 3M 3F, VI-21-1983, C. P. Withrow, (CEWC). Boone Co., Camden Co., 5.5mi. ENE. Linn Ck., Ozark Caverns, IF, III-17-1982, J. E. Gardner, (CEWC) ; Columbia, 3M 2F, III-13-1966, W. W. Dowc^, (FSAC); 2M IF, III-18-1962, 0. Holliday, (FSAC). Callaway Co., 268 Jefferson City, 2M, XII-28-1965, W. W. Dowdy, (ESAC). Camden Co., Bunch Cave, IF, II-9-1957, O. Howksley, (ESAC). Cape Girardeau Co., Imi. N. Eriedheim, 2M, IX-21-1946, H. S. Djlaas, (FMNH). Carter Co., Big Spring S. P., IE, X-31-1954, R. E. Crabill, (RIHC); Spout Springs Annex Cave, 17mi. S. Van Buren, 3M, XI-10-1978, J. E. Gardner, (RIHC). *Chadwick Co., IM, 1900, H. A. Pilsbry, (Chamberlin, 1947). Christian Co., Camp Ridge Cave, 2mi. SW. Chadwick, 2M, IV-18-1979, J. E. Gardner, (NCMH); Smaelin's Cave, 4E, XII-8-1958, B. Ostting, (ESAC). COle Co., SR. 54, 2M 3E, X-25-1965, W. W. Dowdy, (ESAC) ; SR. 54, IX-24-1965, W. W. Dowdy, (ESAC) ; Neilhom Place, 4M, 11-12-1966, W. W. Dowt^, (ESAC) ; flood plain, 2M, VI-27-1968, W. W. Dow<^, (ESAC) ; IM 2E, V-8-1968, W. W. Dowdy, (ESAC). Crawford Co., 5mi. W. Berryman, 2M IE, IV-2-1955, R. E. Crabill, (RIHC). Dade Co., Carrico Cave, IM 3E, III-31-1965, (ESAC) ; Maze Cave, 2M U , IV-25-1965, E. Pendleton & E. Blake, (Pendleton & Blake, 1967), (ESAC). Douglas Co., SR. 4, 21.8mi. E. SR. 5, IE, VI- 20-1983, C. P. Withrow, (CPWC). Howell Co., Spring Ck. Cave, lOmi. S. Wiramow A. Springs, IM, XII-20-1979, J. E. Gardner, (NCMH). Iron Co., Taum Sauk Mt., 5M 4E, X-21-1954, R. E. Crabill, (RIHC); llmi. E. Bixby, Peter Cave #2, IF, VI-2-1982, J. E. Gardner, (CPWC). Johnson Co., Robins, 4M IE, IV-23-1959, J. N. Brooks, (ESAC) ; 3M 4E, IV-1959, J. N. Brooks, (ESAC). Lincoln Co., 4.75mi. NE. Troy, Sherwood Forest Cave, IM, J. E. Gardner, (CEWC) ; Cuivre R. S. P., 2M IE 4J, VI-18-1983, C. P. Withrow, (CPWC). McDonald Co., CR. K, 4.3mi. E. US Rt. 71, Huckleberry Ridge, S. E., IE U , VI-21-1983, C. P. Withrow, (CEWC). *Miller Co., (Causey, 1952a). Monroe Co., CR. FF, 1.5mi. from jet CR. CC & PP, 4M 3E 2J, VI-17-1984, C. P. Withrow, (CEWC). Montgonfôry Co., 2.25mi. W. Danville, Graham Cave, IV-16-1982, IM IE, J. E. Gardner, (CEWC). Oregon Co., White's Ck. Cave, nr Wilderness, 2M, XI-18-1978, J. E. Gardner, (NCMH) ; Alton, 5M 6E, IX-1 to X-31-1954, W. E. Rushton, (ESAC) ; 6M 8E, I-III-1955, W. E. Rushton, (ESAC) ; 6M IF, IV-V-1955, W. E. Rushton, (ESAC). Pettis Co., 4.25mi. N. Sedalia, Stony Ridge Cave, IE U , IV-23-1982, J. E. Gardner, (CEWC) ; 4.25mi. N. Sedalia, Quarry Pit Cave, IE, J. E. Gardner, (CPWC) ; 13mi. SE. Birch Tree, Big Dome Cave, 2M, VI-8- 1982, J. E. Gardner, (CEWC). Phelps Co., 5mi. S. Nevhurg, Coon Cave #1, IM IE, V-1-1982, J. E. Gardner, (CEWC). Pike Co., 4mi. S. Erankford, Du Chein Cave, IM, IV-16-1982, J. E. Gardner, (CPWC). Pulaski Co., 9mi. SSW. Waynesville, Knife Cave, 2F, VE-3-1982, J. E. Gardner, (CEWC). Reynolds Co., SR. 34, 9.3mi. E. US Rt. 60, 2M 4F, VI-19-1983, C. P. Withrow, (CEWC). St. Charles Co., Anhauser Busch Nature Preserve, IM, X-6-1955, R. E. Crabill, (RIHC). St. Francois Co., Imi. S. Dislodge, Sam Hildebrand's Cave, IM, IV-8-1982, J. E. Gardner, (CEWC) ; *Libertyville, IM, IV-2-1937, K. P. Schmidt, (Chamberlin, 1952b). St. louis CO., University City, IM, III-29- 1936, L. Hubricht, (USNM A2541), holotype of Polvdesmus hubrichti; bank of Mississippi R. at lemay, 2M, V-19-1955, R. E. Crabill, (RIHC). Shannon Co., 8mi. WNW. Birch Tree, Den Cave, 2M IE U , X-30- 1982, J. E. Gardner, (CEWC) ; 0.25mi. E. Round Springs, Bootlegger cave, IM, IV-20-1983, J. E. Gardner, (CPWC). Tandy Co., 2M IE, IV- 1962, H. H. Leake, (ESAC) ; *Ozark Underground laboratory, (Aley, 269 1972). Texas Co., 3.75mi. N. Mt. View, B a m Hollow Cave, IM, XI-15- 1982, J. E. Gardner, (CPWC). OKEAHŒA: *Mair Co., (Causey, 1952a). *Cleveland Co., (Causey, 1952a). *Iatimer Co., (Causey, 1952a). le Flore CO., lOmi. SW. Heavener Winding Stair, watdhtower, IM, VI-10-1969, S. Peck, (WASC) ; SR. 1, 10.4mi. E. US Rt. 259, IM 3F, VI-25-1983, C. P. Withrow, (CPWC). logan Co., Guthrie, 5M 7F, X-25-1961, R. C. Harrel, (FSAC). Murray Co., cave, 2M 2F 2J, 11-28-1959, R. C. Harrel, (FSAC) ; *Arbuckle M t . , nr Latimer Co., 3M & F, IV-26-1936, L. Hubricht, (Chamberlin, 1943d). *Payne Co., (Causey, 1952a). Pittsburg Co., 3mi. E. Krebs, 9M 9F, X-22-1961, R. C. Harrel, (FSAC). Pontotoc Co., 2J, IX-23-1958, R. C. Harrel, (FSAC). TENNESSEE: Blount Co., Friendsville, IM, V-1970, R. H. Perry, (RIHC). Franklin Co., US Alt. Rt. 41, 1.5mi. N. SR. 555, 2J, VII-5- 1983, C. P< Withrow, (CEWC). Obion CO., Reelfoot lake S. P., 1.5mi. SE. Samburg, 2M 2F, VII-4-1983, C. P. Withrow, (CPWC) ; Reelfoot lake, 2M, IV-28-1956, F. J. Etges, (FSAC). Shelby C o . , Mertphis, IM 2F, IV-8-1960, B. & L. Dirkson, (FSAC). Sullivan Co., Kingsport, IM 2J, VIII-7-1971, D. A. Rossman, (FSAC). TEXAS: *Bowie Co., (Causey, 1952a). Harris Co., Sheldon, IF, VIII-10-1961, R. O. Albert, (FSAC). "Texas", IM, Hutcherson, (ŒINH), holotype of Polvdesmus americanus. Pseudopolvdesmus tallulanus GEORGIA: Gilmer Co., at Camp Cherry Log, off SR. 5, 2.4mi. S. Lucius, IM, VIII-8-1978, R. M. Shelley & WBJ, (NCMH). *Habersham Co., N. & NW. Clarkesville, MM & FF, IV-27-1943, W. Ivie, (USNM), holotype of Dixidesmus penicillus. Hall Co. Gainesville, 2M 2F, IV- 24-1943, W. Ivie, (USNM), holotype of Dixidesmus humilidens. Lumpkin Co., B l a c W o u m S. P . , nr Dahlonega, 3M IF, VI-4-1975, R. L. Hoffman, (RIHC). Rabun Co., btwn Clayton & Tallulah Falls, IM, IV-28-1943, W. Ivie, (USNM), type of Dixidesmus tallulanus; NW. Clayton, IM, IV-27- 1943, W. Ivie, (USNM), paratype Polvdesmus tallulanus; NW. Clayton, many M & F, IV-28-1943, W. Ivie, (USNM). White Co., Andrews Cave, SR. 17-75, 4.8mi. N. Robertstown, 4M, VII-10-1978, R. M. Shelley & WBJ, (NCMH); Tallulah R., Coleman R. Scenic Area, IM, VII-6-1978, R. M. Shelley & WBJ, (NCMH). NORTH CAROLINA: Haywood Co., N. side Soco Gap, IM IF, VII-21- 1961, R. L. Hoffman, (RIHC); W. side Soco Gap, US Rt. 19, IF, IX-23- 1961, R. L. Hoffman, (RIHC) ; Beech Gap, S. off Sunburst, 4M, VI-14- 1953, L. Hubricht, (RIHC); Waterrock Kiob, summit, 6292', IF, X-30- 1969, W. A. Shear, (WASC) ; US Rt. 276, just S. off Waynesville, 2M 2F, X-19-1974, R. M. Shelley, (NCMH) ; 6mi. S. Sunburst, Beech Gap Rd. at West Fork Pigeon R., 17M 12F, US Rt. 276, 4.Imi. N. Transylvania Co. line, 2M, X-19-1974, R. M. Shelley, (NCMH); 4mi. NW. Maggie along CR. 1300, IM, VII-9-1976, R. M. Shelley, (NCMH). Jackson Co., Whiteside Cave, 4mi. SE. Highland, IM, VII-26-1958, R. L. Hoffman, (RIHC) ; Soco Gap, 14M 9F, V-20-1956, R. L. Hoffitan, Ifeeton, & Lund, (RIHC) ; Coyle Farm, 1.5mi. SW. Webster, 2200', IM, 270 VIII-8-1970, F. A. Cqyle, (NCMH) ; 4M 2F, IV-16-1970, W. A. Shear, (WASC) ; Brushy Fork Hollow, nr Cullovdiee, IM, V-12-1970, W. A. Shear, (WASC) ; Western Carolina University Preserves, Cullotdiee, CullotAee Mt. Rd., 2M IF, X-25-1969, W. A. Shear, (WASC); 12.2mi. ESE. CallotAee, SR. 218 & CR. 1762, IM, VII-10-1976, R. M. Shelley, (NCMH); 7.4mi. S. Cashiers, along CR. 1100, 2M, VII-11-1976, R. M. Shelley, (NCMH). Jackson & Swain Cos., Heintooga Ridge, 5M 2F, VI- 18-1958, R. L. Hoffman & T. Howell, (RIHC). Macon Co., Hi^ilands, 7F, VII-7/13-1958, R. L. Hoffman, (RIHC) ; IM, VIII-2-1949, J. M. Valentine, (RIHC); IM, VIII-2-1949, R. L. Hoffman, (RIHC) ; Hi^ands, 1400', IM, XE-16-1961, R. L. Hoffman, (RIHC); Highlands, 4000', IM, XI-16-1961, R. L. Hoffman, (RIHC); Highlands Biological Station, IF, VIII-lO-1955, J. T. Darlington, (RIHC) ; Little Yellow Mt., 2M, VIII-25-1955, J. T. Darlington, (RIHC) ; Wayah Bald, 3M 6F, VI-13-1958, L. Hubricht, (RIHC); Wayah Bald, 5335', IM, VI-13-1958, L. Hubricht, (RIHC) ; 5mi. NWN. Highlands, 13M 3F, VII-9-1958, R. L. Hoffman, (RIHC) ; W. side Wesser Bald on Otter Ck., IM, VII-29-1949, R. L. Hoffman, (RIHC) ; Blue Valley Rd., 2.4mi. W. SR. 28 at 7.2mi. S. H i ^ a n d s , 35M 22F, VII-11-1960, S. D. Malaik, (RIHC) ; Coweeta Hydrologie Station, IM, VI-16-1973, R. Duffield, (RIHC), 9.9mi. SW. Hi^ilands, dirt rd. at SR. 106, Georgia St. line, IM, VII-31-1974, R. M. Shelley, (NCMH) ; Dry Falls, Hi^ilands Biological Station, IM, VII-22-1949, R. L. Hoffman, (RIHC) ; Turtle Pond Rd., 5mi. W. Highlands, 18M 13F, X-26-1969, W. A. Shear, (WASC); Ellicott Ck., clearcut, 7.5mi. SE. Hi^ilands on Bull Pen Rd., 5M 3F 4J, VI-22- 1976, F. A. Ccyle, (NCMH) ; Cherokee, IM, IV-25-1979, R. M. Shelley, (NCMH). Macon & Jackson Cos., Walnut Ck. Gap, Cowee Mt., IF, VII-15- 1958, R. L. Hoffman, (RIHC). Swain Co., along unk. Rd., 4.4itii. NW. Smokemont, 3.25mi. NW. jck Rt. 441, GSMNP, IM, V-16-1978, R. M. Shelley & WBJ, (NCMH). Transylvania Co., Thompson R. Gorge, SE. Lake Toxaway, IM, IX-6-1961, R. L. Hoffman, (RIHC); Balsam Grove, nr Rosman, IM, VII-20-1961, R. L. Hoffman, (RIHC) ; Pinks Beds Recreational Area, 8mi. NW. Brevard, IF, VII-30-1958, R. L. Hoffman, (RIHC) ; 5.7mi. NW. Brevard, Gov. R d . , l.8mi. W. Fish Hatchery, 6M, VIII-29-1973, R. M. Shelley, (NCMH) ; 3.2mi. NW. Brevard, US Rt. 276, 4.9mi. N. US Rt. 64, 7M, VIII-29-1973, R. M. Shelley, (NCMH) ; 12.Imi. WSW. Rosman, CR. 1152, O.Simi. N. CR. 1151, 14M lOF, VIII-28- 1973, R. M. Shelley, (NCMH) ; 0.4mi. N. Brevard, along USFS R d . , Pink Beds Recreational Area, IM, IX-12-1977, R. M. Shelley, (NCMH). Transylvania & Buncombe Cos., Pisgah Mt. Campground, 3M, VI-15-1953, L. Hubricht, (RIHC). SOUTH CAROLINA: Oconee Co., Welcome Center, US Rt. 85, IM, V- 2-1976, M. R. & J. E. Cooper, (NCMH). TENNESSEE: Sevier Co., GSMNP, nr Summit, IM, VII-5-1933, W. J. Gertsch, (AMNH) ; GSMNP, 3M 2F, VII-8-1933, Gertsch, (RIHC); 7mi. SW. Sugarlands, GSMNP, 5M IF, X-24-1969, W. A. Shear, (WASC). Pseudopolvdesmus erasus AIABAMA: Barbour Co., 6mi. SW. Eufaula, 2M IF, IV-29-1983, R. M. Shelley & P. D. Nader, (NCMH). Clark Co., 8mi. NW. Jackson, SR. 271 69 at Jackson C k . , IM IF, IV-22-1983, R. M. Shelley & B. D. Nader, (NCMH). Clay Co., lake Chinnabee Recreational Area, 13.5mi. NW. Ashland in Tennessee N. F., 2M, V-21-1980, R. M. Shelley & MSM, (NCMH). Cleburne Co., 9.5mi. N. Heflin at Coleman Lake Recreational Area in Tennessee N. F . , IM, V-20-1980, R. M. Shelley & MSM, (NCMH) ; Imi. from Rt. 431 on Rt. 49 in Tennessee N. F., 3M, V-20-1980, R. M. Shelley & MSM, (NCMH). Conecuh Co., alorg US Rt. 84, btwn Belleville and Repton, 2M IF, IV-20-1976, M. A. Cooper, (NCMH). De Kalb Co., Little Ck. Cave, IM IF, VII-6-1968, J. Cooper, (WASC). Hale Co., 15mi. NE. Greensboro, Payne Lake Recreational Area, Tennessee N. F., IM, VII-8-1980, R. M. Shelley & MSM, (NCMH). Henry Co., 2.8mi. NE. Abbeville, SR. 47, 2.Imi. E. SR. 95, 2M, IV-29-1983, R. M. Shelley & P. B. Nader, (NCMH). Houston Co., 5.7mi. W. Dothan, SR. 59 at ChoctavAiatchie R., 2M, IV-30-1983, R. M. Shelley & P. D. Nader, (NCMH). Lawrence Co., entrance of Saltpeter Cave, IM IF, V-27-1987, C. P. Withrow, (CPWC). Madison Co., Monte Sano S. P., E. of Huntsville, IM 3F, VII-22-1939; Huntsville, 2M 2F, IV-24-1943, (MCZ), holotype of Polvdesmus erasus; Huntsville, Monte Sano S. P., 9M IF, V-23-1980, R. M. Shelley & MSM, (NCMH) ; Monte Sano S. P., IM, V-17-1972, S. Peck, (WASC). Mobile Co., 3.Imi. N. Citronelle, US Rt. 45, 2.imi. N. SR. 96, IM, IV-23-1983, R. M. Shelley & P. B. Nader, (NCMH). Monroe Co., 19mi. NE. Monroeville, SR. 47, IM, IV-22-1983, R. M. Shelley & P. B. Nader, (NCMH). Morgan Co., 5.Imi. SE. Falkville, 2M, V-20-1983, R. M. Shelley & Staton, (NCMH). Pickens Co., 4.8mi. E. Ethelsville, US Rt. 82, IM, V-18-1983, R. M. Shelley & Staton, (NCMH). GEORGIA: Chattooga Co., Chattooga, IM, IV-28-1979, R. M. Shelley, (NCMH). Murray Co., Fort Mt. S. P., 2M IF, IV-16-1978, R. M. Shelley & R. E. Ashton, (NCMH). ILLINOIS: *Pope Co., Herod, IM, Faunistic Survey, (Causey, 1952a). KENIUCKY: Letcher Co., US Rt. 119, 32.Imi. W. US Rt. 460, 3F 8J, VII-12-1983, C. P. Withrow, (CPWC). Pulaski Co., Alpine Area on US Rt. 27, nr McCrary Co. line, 3M, VI-14-1976, R. M. Shelley, (NCMH). Warren Co., SR. 101, at Barren R., 4M, VI-15-1976, R. M. Shelley, (NCMH). MISSISSIPPI: Tishomingo Co., J. P. Coleman S. P., 2M, V-25- 1980, R. M. Shelley & MSM, (NCMH). NORLH CAROLINA: Haywood Co., Richland, Balsam summit, 4M IF, X-2-1973, W. A. Shear, (WASC). Jackson Co., n r CullovAiee, IM, VI-24- 1970, A. Weaver, (WASC). Transylvania Co., US Rt. 276, nr Looking Glass Falls, Pisgaii N. F . , 5mi. N. Brevard, 2M, IX-10-1978, W. B. Jones, (NCMH). TENNESSEE: Anderson Co., Oak Ridge, Union Carbide Park, IM, VII-7-1983, C. P. Withrow, (CPWC); SR. 116, 14.5mi. W. SR. 25W, 7M 3F 6J, VII-12-1983, C. P. Withrow, (CPWC). Benton Co., Nathan B. Forrest S. P., 2M, V-26-1980, R. M. Shelley & MSM, (NCMH). Bledsoe Co., Falls Creek S. P., U , VIII-4-1956, M. Engelman, (EMNH) ; Rt. 30, 2.3mi. E. Rt. 28, Rt. 127, 2M IF, V-12-1979, R. M. Shelley, (NCMH); SR. 30, 1.2mi. W. US Rt. 127, IM, VII-6-1983, C. P. Withrow, (CPWC). Carroll Co., SR. 22A, 1.3mi. N. US Rt. 40, Natchez Trace S. P., IM IF, VII-5-1983, C. P. Withrow, (CEWC). Cumberland Co., CR. at 272 Caney Fork R., 11.1 air mi. SW. Crossville, 2M, V-8-1979, R. M. Shelley, (NCMH) ; SR. 68, 9 air mi. SE. Crossville, IM, V-8-1979, R. K. Tarde & R. M. Shelley, (NCMH). Fentress Co., 6.5mi. W. Jamestown, SR. 52, 1.9mi. W. East Fork Obey R., 2M, VI-9-1979, R. M. Shelley & R. K. Tarde, (NCMH). Hardin Co., Pickwick Landing S. P., IM, V-25- 1980, R. M. Shelley & MSM, (NCMH). Henderson Co., Natchez Trace S. P., Fern Nature Trail, 7M, V-26-1980, R. M. Shelley & MSM, (NCMH). Henry Co., Paris Landing S. P., IM, V-26-1980, R. M. Shelley & MSM, (NCMH). Marion Co., 6.5mi. E. Jasper, US Rt. 72, 2M, V-24-1983, R. M. Shelley & Staton, (NCMH) ; 4M 3F, IV-26-1979, R. E. Ashton, A. Braswell & D. Franz, (NCMH); 3.5mi. N. Whitwell, SR. 108, 0.9mi. S. CR. 4303, IM IF, V-23-1983, R. M. Shelley & Staton, (NCMH). Morgan Co., llmi. W. Wartburg on dirt rd., 2.25mi. N. CR. 4252 at Clear Ck., IM IF, VI-7-1979, R. M. Shelley & R. K. Tarde, (NCMH); SR. 62, 1.7mi. S. SR. 116, IF U , VI-12-1983, C. P. Withrow, (CPWC). *Overton Co., Standing Rock S. P., 2M, VII-5-1942, (Loomis, 1944). Rhea Co., vicinity Dayton, IF, summer 1969, W. Henning, (RIHC). Robertson Co., Imi. N. Adams, II, VEII-15-1949, H. S. Dybas, (EfflSH) ; circa 7mi. SW. Springfield, SR. 49, 2.9mi. N. US Rt. 24, IM, VI-12- 1979, R. M. Shelley & R. K. Tarde, (NCMH). Scott Co., 7mi. S. Huntsville, CR. 2342, 3mi. SE. Mt. Pleasant nr Slick Rock, IM, VI-8- 1979, R. M. Shelley & R. K. Tarde, (NCMH). Sequatchie Co., SR. 8, 4.Imi. S. SR. Ill, 8M 7F 4J, VIII-6-1983, C. P. Withrow, (CEWC) ; 5.2mi. SSE. Dunlap, US Rt. 127, Imi. S. CR. 4264, IM, V-24-1983, R. M. Shelley & Staton, (NCMH) ; SR. Ill, 1.5mi. N. CR. 4350, IM, V-12- 1979, R. M. Shelley, (NCMH). Van Buren Co., cave, along SR. 30, E. Spencer, IM, VI-17-1976, R. M. Shelley, (NCMH) ; Fall Creek Falls S. P., 25M IF, VI-17-1976, R. M. Shelley, (NCMH). White Co., Rock Island S. P., entrance, IM, V-8-1979, R. K. Tarde & R. M. Shelley, (NCMH). Wilson Co., US Rt. 70N, 0.8mi. from Smith Co. line, 2M, V- 28-1980, R. M. Shelley & MSM, (NCMH). Pseudopolvdesmus canadensis CANADA QUEBEC: Montreal, IF, IV-11-1964, A. Nimmo, (RIHC). ONTARIO: Hudson's Bay, Albany River, IF, (BMNH), holotype of Polvdesmus canadensis ; Capreol, IF, V-10-1964, A. Nimmo, (RIHC). UNITED STATES FLORIDA: Clay Co., Gold Head Branch S. P., IM, IV-18-1963, R. L. Hoffman, (WASC) ; Sprong Black Ck. at SR. 21, S. SR. 16, IM IF, XI-11-1977, R. M. Shelley & L. R. Franz, (NCMH). Escambia Co., Pensacola, IM of 15 reportedly collected, C. H. Bollman, (USNM), holotype of Polvdesmus nitidus. Levy Co., 6.8mi. SE. Otter Ck., US Rt. 19-98, Imi. S. Waccasassa R., IM IF, VI-11-1983, R. M. Shelley & J. L. Staton, (NCMH). Suwannee R., (USNM), holotype of Dixidesmus phanus. GEORGIA: Charlton Co., Stephan Foster S. P., IM, IX-13-1979, R. M. Shelley, (NCMH). Screven Co., 7mi. N. Sylvania, IM, IV-12- 273 1943, W. Ivie, (USNM), holotype of Dixidesmus svlvicolens. Thomas Co., Thomasville, Charkri, IM IF, IV-2-1940, F. Field, (FMNH). HAWAII; Maui, SB10242, IJM, III-21-1968, W. Sedgwick, (WASC). KENTUCKÏ: Bell Co., Cumberland Mt. nr Cubage, 2M IF, VI-13- 1976, R. M. Shelley & J. K. Ettman, (NCMH) ; Pine Mt. S. P . , Asher Lumber Yard, 2M IF, VI-12-1976, R. M. Shelley & J. K. Ettman, (NCMH); Pine Mt. S. P., nr residences, 2M, VI-11-1976, R. M. Shelley, (NCMH); Pine Mt. S. P., Fonde M t . , IM, VI-12-1976, R. M. Shelley & J. K. Ettman, (NCMH); Pine Mt. S. P., 35M 23F, VI-12-1976, R. M. Shelley & J. K. Ettman, (NCMH) ; US Rt. 119, 14.5mi. W. SR. 421, IM IF U , VII-12-1983, C. P. Withrow, (CPWC). *Fayette Co., (Causey, 1952a). Jackson Co., 7.7mi. S. Estill Co. line, SR. 89, 4M IF, V-11-1978, C. P. Withrow, (CPWC). Lawrence Co., US Rt. 23, 0.7mi. S. SR. 3, 8M 12F, VII-12-1983, C. P. Withrow, (CPWC). Perry Co., Buckhom lake S. P., 3M 2F, V-11-1978, C. P. Withrow, (CPWC). Powell Co., Natural Bridge S. P., 2M IF 2J, VII-9-1966, B. Branson, (WASC). MARYLAND: Frederick Co., Cunningham Falls S. P., nr Thurmond, 4F, XII-6-1956, Hilton, (RIHC). Garrett Co., Crabtree Cave, IF, X- 22-1966, R. Franz, (FSAC) ; nr Jennings, SR. 495, IM 4F, XI-4-1956, R. Highton, (RIHC) ; circa 3mi. W. Bloomington, SR. 135, IM IF, XI-4- 1956, R. Hi<ÿiton, (RIHC); along Savage R . , at first bridge, circa Imi. from SR. 135 on Savage R. Dam Rd., 7M IIF, XI-4-1956, R. Highton, (RIHC) ; nr Jennings, SR. 495, 2JM 4F, XI-4-1956, R. Hilton, (RIHC). MICHIGAN: *Alcona, Alpena, Antrim, Benzie, Charlevoix, Cheboygan, Chippewa, Crawford, Dickinson, Emmet, Grand Traverse, Kalkaska, Leelanau, Mackinac, Montmorency, Oscoda, Otsega, Presque Isle, Wexford Cos., (Johnson, 1954b). Mackinaw Co., W. side Garnet Lake on Old US Highway 2, T34N-R8W-S4, 2M 2F, VII-31-1949, B. M. Johnson, (USNM 2117), holotype of Dixidesmus qausodicrorhachus. MISSISSIPPI: Harrison Co., Pass Christian, IM 3F several J, 11-15-1946, J. & W. Rap, (USNM), holotype of Dixidesmus christianus. Jackson Co., 4mj.. E. Ocean Springs, U , X-15-1953, H. S. D^tas, (FMNH); Ocean Springs, around Gulf Hills, IM IF, X-2-1958, (FSAC). NEW YORK: Cattaraugus Co., Allegheny S. P., Anderson Trail, IM 3F, V-10-1958, W. B. Mudhmore, (RIHC). Greene Co., Catskill, (ANSP), holotype of Dixidesmus catskillus; along 23A nr Jarrett, IF, VI-15- 1975, R. M. Shelley, (NCMH) ; nr Lanesville, IM 2F, VIII-15-1978, T. L. McCabe, (NCMH); 412 meters from Lanesville, 3M, VIII-18-1978, R. M. Shelley & T. L. McCabe, (NCMH) ; Diamond Notch nr Lanesville, IM, VIII-19-1978, R. M. Shelley & T. L. McCabe, (NCMH). Monroe Co., Mendon Pond, 2M, X-7-1954, W. B. Machmore, (RIHC). Ulster Co., nr Poenicia, IM, VIII-18-1978, R. M. Shelley, (NCMH). NORTH CAROLINA: Alexander Co., 5.3mi. SW. Taylorsville, CR. 1128, O.lmi. E. CR. 1121, IM, VIII-13-1975, R. M. Shelley & J. C. Clamp, (NCMH). Ashe Co., Imi. S. Grayson, IF, V-17-1964, R. L. Hoffman, (RIHC) ; 8.8mi. WSW. Jefferson, CR. 1100, l.Omi. N. CR. 1125, 2J, VII-21-1972, R. M. Shelley, (NCMH) ; 7.2mi. WSW. Jefferson, CR. 1125, l.Omi. E. CR. 1100, 5J, VI-21-1972, R. M. Shelley, (NCMH) ; 8.6mi. SSE. Jefferson, CR. 1183, 0.6mi. N. CR. 1003, 6J, VII-21- 1972, R. M. Shelley, (NCMH) ; along rd to Mt. Jefferson S. P. btwn 274 1st & 2nd gates, in open, IM, V-21-1978, R. E. Ashton & J. E. Cooper, (NCMH). Avery Co., Linville, US Rt. 221, IF, X-19-1969, L. S. Kni^t, (RIHC); 4.4mi. N. Elk Park, CR. 1305, 1.7mi. N. CR. 1307, IM, IX-8-1973, R. M. Shelley, (NCMH); 2.8mi. N. Elk Park, along CR. 1305 at Elk R. Falls, IM IF U , X-29-1977, R. M. Shelley, (NCMH) ; 13.5mi. N. Newland at jet Rt. 321 & CR. 1316, IM IF, VI-19-1980, R. M. Shelley & MSM, (NCMH). Buncombe Co., 8.8mi. WSW. Asheville, CR. 1220, at Pole Ck., IM, IX-7-1977, R. M. Shelley, (NCMH); 8.8mi. SE. Asheville, Povdiatan Recreational Area, Pisgah N. F., 2M, IX-5-1977, R. M. Shelley, (NCMH); 6.8mi. SW. Asheville, SR. 191, 0.2mi. N. Blue Ridge Parkway, Bent Ck. Forest Experimental Station, 4M, X-20-1974, R. M. Shelley, (NCMH) ; 12.6itii. SE. Asheville along US Rt. 74, 0.4mi. W. Henderson Co. line, 3M, VII-12-1976, R. M. Shelley, (NCMH) ; 2.7mi. W. Black Mt., CR. 2468, Imi. N. CR. 2435, 2M, IX-5-1977, R. M. Shelley, (NCMH) ; 5.2mi. NE. Asheville, CR. 2424, hni. W. CR. 2419, IM, IX-5-1977, R. M. Shelley, (NCMH) ; 2.5mi. NE. Bamardsville, CR. 2027, 0.9mi. NE. SR. 197, IM, IX-6-1977, R. M. Shelley, (NCMH). Burke Co., 4.8mi. SE. Morganton, CR. 1922, 0.4mi. S. CR. 1936, IM, VII-26-1975, R. M. Shelley & J. C. Claitp, (NCMH) ; Morganton, 3M, XII-3-1982, J. Rivers, (NCMH). Caldwell Co., 5.7mi. NW. Kings Creek, CR. 1507, 0.2mi. N. CR. 268, IM, VII-21-1976, M. Filka & F. Scott, (NCMH) ; llmi. N. Lenoir, CR. 1371, 3.0mi. on CR. 1370, IM, VI-22-1980, R. M. Shelley & MSM, (NCMH) ; 11.6mi. ENE. Lenoir, CR. 1730, 0.6mi. S. CR. 1772, IM, VIII-13-1975, R. M. Shelley & J. C. Claitp, (NCMH). Cleveland Co., 5.7mi. SW. Kings Mt., CR. 2283, 0.8mi. NE. SR. 216, 2F, VII-8-1976, M. Filka & W. Thomas, (NCMH); 3mi. S. Kings Mt., CR. 2289, 0.6mi. SR. 161, IM, X-18-1976, M. Filka & G. Wicker, (NCMH) ; 3.9mi. SE. Kings Mt., SR. 161, imi. S. CR. 2289, IF, X-18-1976, M. Filka & G. Wicker, (NCMH); 4.3mi. SW. Kings Mt., CR. 2245, 0.2mi. W. CR. 2283, IF, X-18-1976, M. Filka & G. Wicker, (NCMH) ; 3.Imi. S. Kings Mountain (town) along rd., 0.4mi. S. CR. 2289, 0.7mi. W. 2nd exit SR. 161, 15M 3F, IV-30-1976, R. M. Shelley, (NCMH) ; 3.9mi. S. Kings Mmxntain (town), SR. 161, 0.2mi. S. CR. 2354, IM, IV-30-1976, R. M. Shelley, (NCMH) ; 4mi. SW. Kings Mountain (town), CR. 2245, 0.2mi. N. CR. 2283, IM, IV-30-1976, R. M. Shelley, (Na»IK) ; 5.Imi. SW. Kings Mountain (town), CR. 2286, 0.8mi. SW. CR. 2283, U , IV-8-1977, W. Filka, (NCMH); 4.75mi. SW. Kings Mountain (town), CR. 2245, 1.4mi. SE. 2283, IF, X-18-1976, M. Filka & G. Wicker, (NCMH) ; jet 161 & SR. 85, FDA, 1.2mi. SE. Kings Mt., 2M 2F U , X-18-1976, M. Filka & G. Wicker, (NCMH) ; Cumberland Co., Fayetteville, CR. 1016, Nhrtindale Dr., 2M, Fall 1977, M. Dennis, (NCMH). Gaston Co., 7.0mi. SE. Bessemer City, CR. 1103, jet CR. 1112, U , IV-9-1976, M. Filka, (NCMH) ; 6.2mi. SE. Bessemer City, O.Smi. SW. CR. 1113, IM 4J, X-16-1976, M. Filka & G. Wicker, (NCMH) ; U , IV-9-1977, M. Filka, (NCMH); 3.4mi. SE. Bessemer City, CR. 1131, O.lmi. NW. CR. 1133, IM 3J, X-16-1976, W. Filka & G. Wicker, (NCMH) ; 2.8mi. S. Bessemer City, CR. 1122, 0.4mi. E. CR. 1120, 4M IF U , X- 16-1976, M. Filka & G. Wicker, (NCMH) ; 6.9mi. S. Bessemer City, CR. 1112, 0.2mi. E. CR. 1125, U , X-17-1976, M. Filka & G. Wicker, (NCMH); W. Gastonia, CR. 1106, 1.5mi. E. CR. 1236, IM IF, X-16-1976, M. Filka & G. Wicker, (NCMH) ; 4.8mi. NE. Gastonia, CR. 2201, l.Smi. N. SR. 7, IF, X-31-1976, M. Filka, (NCMH) ; 4.8mi. SW. Gastonia, CR. 275 1131, O.lmi. NW. CR. 1131, IF, VII-9-1976, M. Filka & W. Thomas, (NŒH) ; 2.5mi. S. Bessemer City, CR. 1125, O.lmi. SE. Rt. 74-9, 3M 2F, IV-9-1977, M. Filka, (NŒH) ; 6.9mi. S. Bessemer City, CR. 1122, 0.4mi. E. OR. 1125, 2M IF, IV-9-1977, M. Filka, (NCMH). Graham Co., Joyce Kilmer Memorial F., 3M, V-20-1970, W. A. Shear, (WASC). Haywood Co., 7.0mi. NW. Waynesville, CR. 1318, jet CR. 1319, IM, VII-8-1976, R. M. Shelley, (NCMH) ; GSMNP, 12mi. NW. Waynesville, rd. to Cosky, 0.6mi. W. Catalcodhee Ed. nr campground, IM, VII-8-1976, R. M. Shelley, (NCMH) ; 4mi. SE. Canton, CR. 1901, 0.4mi. E. CR. 1854, IM, IX-12-1977, R. M. Shelley, (NCMH); 14mi. NW. Waynesville, GSMNP, CR. 1397, 3.3mi. W. Catalcodhee Rd., IM, VII-8-1976, R. M. Shelley, (NCMH); Richlands, Balsam summit, 6400', IM, X-10-1971, W. A. Shear, (WASC); Waterrock Knob, 6292', 3M IF, X-13-1971, W. A. Shear, (WASC). Henderson Co., 6mi. E. Fletcher, CR. 1569, Imi. W. CR. 1594, IM, IX-13-1977, R. M. Shelley, (NCMH) ; Tuxedo, IM IF, XI- 30-1975, R. M. Shelley, (NCMH) ; Flat Rock, Bayou Dr. & Rue de Choiseaul., ]M, X-18-1974, R. M. Shelley, (NCMH) ; Flat Rock, 2M 3F, X-18-1974, R. M. Shelley, (NCMH) ; jet CR. 1127 & 1128, 5.6mi. SE. Etowah, IM, VI-8-1978, R. M. Shelley & WBJ, (NCMH). Jackson Co., Long branch C k., IM, VII-17-1970, F. A. Coyle, (NCMH); 7.6mi. ESE. Cullovdiee along Rt. 218, l.Omi. N. CR. 1137, IM, VII-10-1976, R. M. Shelley, (NCMH) ; Western Carolina University Preserve, CullcRiÆiee, CullovAiee Mt. R d . , 3M 3F 2J, X-25-1969, W. A. Shear, (WASC) ; Richland Balsam on Blue Ridge Parkway, 2M, VIII-9-1971, F. A. Coyle, (NCMH) ; 12.5mi. N. Franklin off Rt. 441, IM 4F, X-8-1973, R. Duffield, (RIHC) ; 9.2mi. ESE. Cullo&hee, SR. 218, O.lmi. N. CR. 1138, 3M 5F, VII-10-1976, R. M. Shelley, (NCMH) ; 9mi. NNE. Sylva, rest area, US Rt. 19, 1.2mi. WSW. Haywood Co. line, IM 3F, XI-25- 1978, A. L. Braswell & D. L. Stephan, (NCMH). McDowell Co., Black Mt. Campground nr entrance, IM, X-11-1975, J. C. Clamp, (NCMH) ; 3.2mi. S. Little Switzerland along CR. 1443, IM, X-28-1977, R. M. Shelley, (NCMH) ; along CR. 1566, 0.2mi. jet Blue Ridge Parkway, 3.5mi. N. Pitts, IM, V-23-1978, R. M. Shelley & WBJ, (NCMH) ; Marion, 2M, XII-14-1979, A. Etyrd, (NCMH). Macon Co., 5mi. NW. Rainbow Springs along USFS Rd. 437, IM IF, VI-2-1977, A. L. Braswell, (NCMH) ; 5mi. NNW. Rainbow Springs along USFS rd. 437, 2M 2F 8J, IV- 5-1978, A. L. Braswell, (NCMH). Madison Co., 9.6mi. SW. Marshall, SR. 63, 3.5mi. S. SR. 209, 4M, IX-11-1977, R. M. Shelley, (NCMH) ; 2.4mi. N. Hot Springs, along 25-70, 0.7mi. S. Governor Rd., 3M, R. M. Shelley & J. C. Clamp, (NCMH). Martin C o . , ravine rd, at end CR. 1347, 0.7mi. N. CR. 903, 4mi. N. Oak City, 8M IF, X-18-1979, R. M. Shelley & P. T. Hertl., (NCMH). Mecklenburg Co., Charlotte, IM, XI- 24-1976, R. M. Shelley, (NCMH). Mitchell Co., 12mi. NW. Bakersville, CR. 1323, 0.7mi. N. CR. 1321, IM, VII-24-1975, R. M. Shelley & J. C. Clamp, (NCMH) ; 11.5mi. NW. Bakersville, CR. 1323, 0.7mi. W. CR. 1321, IM, VII-24-1975, R. M. Shelley & J. C. Clamp, (NCMH) ; along CR. 1349, 0.9mi. jet CR. 1304, 3.25mi. SE. Popular, 2M IF, V-21- 1978, R. M. Shelley & WBJ, (NCMH) ; 7.5mi. NW. Spruce Pine, CR. 1178, 1.2mi. on SR. 80, IF, VI-21-1980, R. M. Shelley & MSM, (NCMH). Polk Co., 5mi. NE. Saluda, along Cove Ck., 3M 3F, X-15-1973, R. M. Shelley, (NCMH) ; 3.2mi. WNW. Tryon, along US Rt. 176, just S. Pacolet R . , IM, X-16-1974, R. M. Shelley, (NCMH); 2mi. NW. Columtbus, 276 along CR. 1136, 3M, X-15-1973, R. M. Shelley, (NŒH). Richmond Co., 6.7mi. NW. Ellerbe, SR. 73, O.Smi. W. CR. 1005, IM IF, V-1-1974, R. M. Shelley, (NŒH). Rutherford Co., 3.Imi. SW. Rutherfordton, SR. 108, 0.5mi. N. CR. 1193, 2M IF, X-15-1973, R. M. Shelley, ( N Œ H ) . Stokes Co., 2.4itd. W. Danbury, CR. 1001, 0.4mi. W. SR. 8-89, 2F, V- 30-1973, R. M. Shelley, (NŒH). Surry Co., Elkin, jet rd to hospital and CR. 1167, 0.3mi. W. CR. 1144, IM IF, X-12-1975, R. M. Shelley, (NCMH). Swain CO., 8.4mi. NW. Smokemont along 441, 6.5 air mi. NW. Smokemont, GSMNP, IM, V-16-1978, R. M. Shelley & WBJ, (NCMH) ; rd across Fontana Dam, ri ^ t turn, 0.5mi. from over jet, IM, V-16-1978, R. M. Shelley & WBJ, (NCMH) ; GSMNP, above Fontana Dam, Appalachian Trail, IM IF, VI-23-1970, A. Weaver, (WASC) ; GSMNP, NW. Bryson City, 8.0mi. W- Park, IM, V-9-1980, R. M. Shelley & MSM, (NCMH); GSMNP, 8.4mi. NW. Smokemont along 441, 6.5 air mi. NW. Smokemont, IM, V-16- 1978, R. M. Shelley & WBJ, (NCMH). Transylvania Co., 10.4mi. N. Brevard, along USFS Rd., Pink Beds Area, IM, IX-12-1977, R. M. Shelley, (NCMH) ; 5mi. N. Brevard, US Rt. 276 nr Looking Glass Falls, Pisgah N. F., IM, IX-10-1978, W. B. Jones, (NCMH); US Rt. 276, nr Sliding Rock, Pisgah N. F . , IM, IX-10-1978, W. B. Jones, (NCMH). Watauga Co., 2.2mi. N. Blowing Rock, CR. 1541, 1.5mi. N. CR. 1552, 2M, IV-26-1974, R. M. Shelley, (NCMH) ; Blowing Rock, Gonth R d . , 0.5mi. N. US Rt. 321, 8M IF, IX-8-1973, R. M. Shelley, (NCMH) ; 1.9mi. S. Boone, CR. 1549, 0.7mi. N. 1550, IM 5F, IV-26-1974, R. M. Shelley, (NCMH) ; Valle Crucis, SR. 194, 0.6mi. S. 1113, 12M, X-11- 1975, J. C. Claitp, (NCMH). Wilkes Co., 7.2mi. NW. North Wilkesboro, CR. 1541, O.lmi. E. CR. 1544, IM, VIII-12-1975, R. M. Shelley & J. C. Clairp, (NCMH) ; lOmi. SW. Moravain Falls, CR. 1130, 0.5mi. N. CR. 1129, IM, VIII-12-1975, R. M. Shelley & J. C. Clamp, (NCMH). Yancey Co., Oakwoods, 3.0mi. NW. Spruce Pine, IF, VI-4-1954, R. L. Hoffman, (RIHC); along CR. 1340, 0.5mi. jet CR. 1417, 3.0mi. NE. Ramsey Town, IM, V-22-1978, R. M. Shelley & WBJ, (NCMH) ; Mt. Mitchell, 6600', 2M 3F, V-13-1970, F. A. Cpyle, (NCMH); 13.4mi. S. Burnsville, Gov. Rd., 2.3mi. NW. Blue Ridge Park^y, IM, VII-20-1975, R. M. Shelley & J. C. Clamp, (NCMH) ; 7.7mi. SW. Burnsville, SR. 197, 3mi. W. CR. 1101, IM IF, IX-10-1977, R. M. Shelley, (NCMH) ; Mt. Mitchell, btwn Stepp's Gap & Blue Ridge Parkway, IM, VI-31-1972, R. L. Hoffman, (RIHC); Mt. Mitchell, camping area nr top, 4M IF, VII-31-1972, R. L. Hoffman, (RIHC); IM, VII-31-1972, R. L. Hoffman, (RIHC) ;_Mt. Mitchell S. P., camping area, 4M 5F, VIII-2-1968, Bayless, (WASC). PENNSYLVANIA: Bedford Co. ^ New Paris Sink, 6M 3F, 1961, N. D. Richmond, (FSAC); Bedford Township, S. Bedford, service plaza on turrpike, IM, VI-24-1973, C. B. Stein, (0S2M). Butler Co., New Cherry Township, 1.5mi. S. Boyer, 6.5mi. ESE. Harrisville, 2F, IX-6- 1975, C. B. Stein, (OSZM). Luzerne Co., Shawanese (Harvey's Lake), 2M 2F, F. C. Paulmier, (USNM), holotype of Polvdesmus echinoaon. Potter CO., Dir^man Run, sawmill, IM 2F, IX-3-1965, W. A. Shear, (WASC). SOUTH CAROLINA: Charleston Co., Wadmalaw Island, Bears Bluff Lab, IM, 1958, N. B. Causey, (NCMH). Cherokee Co., 4.6mi. SE. Blacksburg, SR. 5, 0.3mi. S. % . 68, IM, V-10-1977, R. M. Shelley, (NCMH). Chesterfield Co., Cheraw S. P., 6M 2F, XI-5-1979, J. F. Cornell, (NCMH). Laurens Co., Laurens City Park, IM, V-8-1977, R. M. 277 Shelley, (NŒH). Spartanburg Co., Spartanburg, U, IX-18-1943, R. L. Wenzel, (FMNH). Croft S. P., IM, VIII-20-1976, R. M. Shelley, (NCMH). TENNESSEE; Anderson Co., 4mi. N. Clinton, IM, (WASC). Blount Co., dirt rd. off CR. 2422, Blount-Sevier Co. line, 14mi. E. Maryville, IM, X-11-1978, R. M. Shelley & WBJ, (NŒH) ; creek off SR. 73, 4mi. W. Townsend, IM, V-17-1978, R. M. Shelley & WBJ, (NCMH) ; Cade Cove Rd., W. prong, 1st bridge after tunnel, GSMNP, 2M, V-17- 1978, R. M. Shelley & WBJ, (NCMH) ; Cades Gove, Vista Trail, at caitpground, 2M, VE-18-1976, R. M. Shelley, (NCMH) ; Cades Cove, vista trail, at caiipground, 2M, VI-18-1976, R. M. Shelley, (NCMH); SR. 321, 4.6mi. S. jet SR. 73, 5M 3F 5J, VII-9-1983, C. P. Withrow, (CPWC). Carter CO., SR. 143, 2.2mi. S. Roan M t . , 2M IF, X-11-1975, J. C. Clairp, (NCMH); SR. 143, 8.5mi. S. Roan Mt., IM, X-11-1975, J. C. Clairp, (NCMH). Cocke Co., 7.5 air mi. SW. Newport, dirt rd. off SR. 32, 2M, X-9-1978, R. M. Shelley & WBJ, (NCMH); nr jet SR. 160 & CR. 2487, 4.75 air mi. NE. Newport, IM, X-10-1978, R. M. Shelley & WBJ, (NCMH) ; GSMNP, Cosby Pienie Area & Trail, IM, V-19-1978, R. M. Shelley & WBJ, (NCMH) ; rd. to Catow's Grove, 11.5 air mi. S. Newport, IM, V-19-1978, R. M. Shelley & WBJ, (NCMH) ; Ck. off SR. 73, 4mi. W. Townsend, IM, V-17-1978, R. M. Shelley & WBJ, (NCMH). Greene Co., 1st rd off SR. 70, along NC. St. line, IM, V-20-1978, R. M. Shelley & WBJ, (NCMH) ; Paint Ck. Reereational Area, 4mi. NE. Wolf Ck., IM, VIII-1-1962, R. L. Hoffinan, (RIHC). Hamblen Co., Panther Creek S. P., 6.5 air mi. W. Morristown, IM, X-15-1978, R. M. Shelley & WBJ, (NCMH). Jefferson Co., Jefferson City, Mossy Ck., IM, C. B. Branner, (USNM), holotype of Polvdesmus branneri; along unnumber rd., 11.6mi. jet SR. 32, 8 air mi. NE. Danridge, IM, X-15-1978, R. M. Shelley & WBJ, (NCMH). Knox Co., *Kioxville, Branner, (Bollman, 1893); University Tennessee, woodlot, 4M IF, 11-15-1972, W. Tolbert, (RIHC). McMinn Co., CR. 4371, 0.3mi. from jet CR. 4276, 11.2 air mi. SE. Athens, IM, X-13-1978, R. M. Shelley & Ï-3BJ, (NCMH) ; Cherokee N. F., along CR. 4371, Q.3mi. jet CR. 4276, 11.2 air mi. SE. Athens, IM, X-13-1978, R. M. Shelley & WBJ, (NCMH). Morgan Co., Frozen Head S. P., 5M 8F, III-24-1984, C. P. Withrow, (CEWC). Pickett Co., Jamestown, Jordan Motel, IF 3J, VI-16-1962, H. R. Steeves, (FSAC). Seott Co., lOmi. NE. Huntsville, SR. 2451, O.Smi. E. South Fork Cumberland R . , IM, VI-9-1979, R. M. Shelley & R. K. Tarde, (NCMH). Sevier Co., GSMNP, Porter's Flat, Greenbrier Cove, IM 19J, VI-14-1939, D. C. Lowrie, (FMNH) ; Trillium Gap, 4700 ft, 4J, VI-15-1939, D. C. Lowrie, (FMNH) ; Gatlinburg, IM, several collected, VI-13/19-1942, (USNM), holotype of Polvdesmus conlatus; GSMNP, 3M, (WASC) ; GSMNP, Newfound Gap, IM IF, (WASC) ; 2.5mi. above Chimney Carrpground, 2J, H. S. D^Pas, (FMNH) ; entrance to Chimney Campground, 2F, VI-17-1963, J. LLoyd, (OSEM) ; 5000', 2M, X-13-1970, W. A. Shear, (WASC) ; GSMNP, n r Cosby Campground, 2M, V-14-1966, T. & B. Hlayac, (WASC); 3M 9J, IX-5/14-1966, T. & B. Hlayac, (WASC) ; GSMNP, trail. Newfound Gap to Mt. LeConte to Alum Cave Bluffs, IM, VI-5/6-1972, W. A. Shear, (WASC) ; 2mi. NNW. Newfound Gap, 3M, X-13-1970, W. A. Shear, (WASC). GSMNP, Elkmont Campground, IM, VI-19-1976, R. M. Shelley, (NCMH) ; GSMNP, Chimney Picnic Area, 7.25mi. S. Gatlinburg, 3M IF, V-16-1978, R. M. Shelley & WBJ, (NCMH); GSMNP, Roaring Fork 278 Natural Trail, btwn 3rd & 4th bridge, 2.9mi. SE. Gatlinburg, 4M IF, V-16-1978, R. M. Shelley & WBJ, (NŒH) ; GSMNP, Junglebrook Camp, 2 mi. N. Gatlinburg, IM, V-8-1980, R. M. Shelley & MSM, (NŒH) ; North Carolina & Tennessee, GSMNP, Newfound Gap, IM IF, X-24-1969, W. A. Shear, (WASC) ; GSMNP, Huskey Gap Trail, at Rt. 441, 2M 3F, X-24- 1969, W. A. Shear, (WASC),* GSMNP, B.Omi. NW. Bryson City, West Park, IM, V-9-1980, R. M. Shelley & MSM, (NŒH). Unicoi Co., Rock Creek Recreational Area, Cherokee N. F., CR. 2457, 3mi. E. Erwin, IM, V- 20-1978, R. M. Shelley & WBJ, (NŒH). VIRGINIA: Alle^iany Co., 3mi. N. Clifton Forge, 3M, III-30- 1947, R. L. Hoffinan, (RIHC); McGraw's Gap, 8M, IV-13-1947, R. L. Hoffinan, (RIHC); Stull's Cave, Richpatch, IM IF, VI-16-1947, R. L. Hoffman, (RIHC). Augusta Co., Humpback Mt., IF, VI-17-1957, R. L. Hoffitnan, (RIHC) ; Reddish Knob, 6M 3F, Hil t o n , Jones, & Littleford, (RIHC). Bedford Co., Blue Ridge Parkway, milepost 80.9, 2M IF, X-20- 1956, R. L. Hoffman, (RIHC). Bland Co., East R. Mt., above Cove Ck., 7M 5F, VI-3-1971, W. A. Shear, (WASC) ; 3mi. NE. South Gap, 3F, IV- 20-1962, R. L. Hoffman, (RIHC). Botetourt Co., N. side Bearhollow Gap, circa 2100', 3mi. S. Buchanan, Rt. 43, IM IF, V-27-1962, R. L. Hoffman, (RIHC). Buchanan Co., 2.7mi. SW. Vansant, 2M IF U , VTI-l- 1951, R. L. Hoffman & Newman, (RIHC). Carroll Co., along Rt. 92, 0.9mi. N. Fries, 2F, VI-20-1962, R. L. Hoffman, (RIHC). Craig Co., Top of Potts Mt., E. of Paint Bank, 3M 4F, III-13-1962, R. L. Hoffman, (RIHC). Dickenson Co., along Cranes Nest R., circa 5mi. W. Haysi, IM 3F, IV-21-1962, R. L. Hoffitnan (RIHC); Break Interstate Park, nr Haysi, 2M, VI-14-1961, R. L. Hoffman, (RIHC). Flcyd Co., Rocky Kiob Park, SE. Floyd, IM, VII-3-1947, R. L. Hoffman, (RIHC) ; Goose Ck., circa 3mi. N. Simpson, 1850', IF, X-9-1971, R. L. Hoffman & Knight, (RIHC). Giles Co., 5mi. N. Perhbroke, Cascades Little Stony Ck., 3M IF, IX-26-1970, W. A. Shear, (WASC); IF, V-17-1961, R. L. Hoffman, (RIHC) ; Mt. Lake, IM, VII-2-1947, R. L. Hoffman, (RIHC) ; Mt. Lake Biological Station, 16M, vTII-27-1947, H. H. Hobbs, Jr., (RIHC). Grayson Co. , Mt. Rogers, 2M, VI—19—1954, k . o . Horrman & Fowler, (RIHC); Mt. Rogers, 5400', 2M, VI-10-1962, R. L. Hoffman, (RIHC) ; 3M IF, VI-4-1963, Covell, (RIHC). Highland Co., locust Spring Recreational Area, circa 8mi. IQNW. Hi(ÿitown, 3900', iM IF, IV-28-1972, R. L. Hoffman & Kiight, (RIHC). Lee Co., Cave Spring Recreational Area, 2mi. N. Dryden, 2M, IX-2/3-1972, R. L. Hoffman, (RIHC). Montgomery Co., Blacksburg, IV-13-1958, R. L. Hoffman & Crabill; 5mi. NE. Blacksburg, 3M 2F, IV-10-1969, W. A. Shear, (WASC) ; Ipper Poverty C k . , 2020', IM IF, IV-16-1966, Scholz & Brownell, (RIHC). Rockbridge Co., ipple Orchard Mt., circa 3200', IM IF, X-14-1962, Radford College Biol. Club, (RIHC). Russell Co., Imi. NW. Lynn Springs Post Office, 2M IF, IV-20-1962, R. L. Hoffman, (RIHC). Scott Co., llmi. E. Hilton, US Rt. 58, 4M IF, VIII-29-1962, R. L. Hoffman, (RIHC). Tazewell Co., Bear Mt., 4F, VI-29-1947, R. L. Hoffman & KLeinpeter, (RIHC); Burkes Garden, 4000', IF, III-20-1954, R. L. Hoffman, (RIHC). Washington Co., 4mi. SW. Konnarock, 4M 4F, IV-28-1951, L. Hubricht, (RIHC) ; Abram's Falls, circa 5mi. NW. Bristol, IF, rv-13-1974, D. W. Ogle, (RIHC). Wise Co., Irai. N. Kent, circa 3mi. E. Tppalachia, SR. 603, IM, VI-26-1971, R. L. Hoffman & 279 (RIHC) ; Vesuvius Table Rock, 4M, VIII-13-1954, M. Caskie, (RIHC). WEST VIRGINIA: Berkeley Co., SR. 9, at Opequon R. Bridge, IM IF, VIII-23-1977, C. P. Withrow, (CPWC). Gilmer CO., SR. 17, 1.4itii. S. entrance Cedar Creek S. P., IM 6J, VIII-13-1977, C. P. Withrow, (CEWC). Greenbrier Co., SW. end of Kate's Mt., summit, picnic area, IM 2F, VE-24-1967, W. A. Shear, (WASC); North Anthony Ck., 5itii. NNE. Neola, 2M 2F, X-1-1972, W. A. Shear & W. Ash, (WASC); Greenbrier S. F. along 3mi. Young's Nature Trail, IM, IV-28-1973, W. A. Shear, (WASC); Kate's Mt., 4M 6F, IV-20-1968, W. A. Shear, (WASC); same, many, IV-20-1971, W. A. Shear (WASC) ; SW. end Kate's M t . , summit picnic area, IM 2F, VE-24-1967, W. A. Shear, (WASC) ; SR. 16/2, 5.2mi. E. Anthony, 2M 2F, VII-31-1977, C. P. Withrow, (CEWC) ; Dawson, IF U , IV-27-1966, (WASC). McDowell Co., Windmill Gap nr McComas, IM, IV-15-1973, W. McKenzie, (WASC). Mercer Co., Athens, IM, X-24-1972, W. A. Shear, (WASC) ; IM, III-12-1967, W. A. Shear, (WASC) ; Old Puirp House, IM, III-23-1967, W. A. Shear, (WASC) ; Brush Ck. Falls, 2M, V-9-1970, W. A. Shear, (WASC) ; Caitp Ck. S. F . , IM, X- 18-1970, W. A. Shear, (WASC). Mineral Co., US Rt. 50, l.Smi. W. Clayville, IM IF, VIII-7-1977, C. P. Withrow, (CEWC). Monroe Co., 2mi. N. Ballard on SR. 12, 7M 8F, IX-14-1962, R. L. Hoffman, (RIHC) ; 0.7mi. into Cacapon S. P., IM IF, VIII-23-1977, C. P. Withroif, (CEWC). Nicholas Co., SR. 39, 2.5mi. E. Nettie, IM 4J, VIII-5-1977, C. P. Withrow, (CEWC). Pendleton Co., SR. 14, off US Rt. 33, White's Run, 2M IF, X-1-1977, C. P. Withrow, (CEWC). Pocahontas CO., Hills Ck. Falls Scenic Area, 2F, V-5-1970, W. A. Shear, (WASC) ; Droop Mt. Battlefield S. P., 2M, X-17-1971, R. L. Hoffman & Knight, (RIHC) ; IM, IV-30-1967, W. A. Shear, (WASC) ; IM IF, VI-19-1971, W. A. Shear, (WASC) ; Kate's Mt., summit, IF, IV-1-1967, W. A. Shear, (WASC) ; 3M 6F, IV-20-1968, W. A. Shear, (WASC) ; Cranberry Glades Natural Area, IM 2F, V-5-1970, W. A. Shear, (WASC) ; same, 2M IF, V-20-1967, W. A. Shear, (msc) ; 3F, VI-18-1963, R. L. Hoffman, (RCHC) ; Monongahela N. F., Cranberry Mt., Visitor Area, 5M, IV-30-1972, W. A. Shear, (WASC) ; l.Omi. NNE. Gaudineer Kiob, IM 2F, IX-24-1960, Highton & Savage, (RIHC) ; Ambassadors for Christ Campground nr Huntersville, 3M IF, X-12-1974, E. Burkhalter, (RIHC) ; 2M, (RIHC). Preston Co., US Rt. 50, Cathedral S. P., 3M 7F 51, VIII-22-1977, C. P. Withrow, (CEWC) ; US Rt. 50, 0.3mi. E. Brookside, 2M 6F, VIII-7-1977, C. P. Withrow, (CEWC). Randolph Co., SR. 14, 5.2mi. S. Whitmer, IM, IX-15- 1977, C. P. Withrow & G. Wamsley, (MUIC) ; SR. 219/16, Kurribrabow S. F . , W. Oxley lîun, 2M 4F, VIII-14-1977, C. P. Withrow, (CEWC) ; l.Smi. NW. Gaudineer Kidb, 3M 3F, IX-24-1960, Highton & Savage, (RIHC). Summers Co., Bull Falls Recreational Area, IM, IV-2-1968, (WASC) ; IM, (WASC). Tucker Co., Dolly Sods, Imi. N. park entrance, IM 7F, IV-15-1977, R. M. & P. K. Blaney, (R. M., Blaney) ; 0.2mi. N. Lanesville, IM, VI-18-1977, C. P. Withrow, (CEWC) ; 4M 4F, X-1-1977, C. P. Withrow & G. Wamsley, (CEWC). Hpshire CO., SR. 20, 0.9mi. S. Kanavha Head, 3M 3F, VIII-15-1977, C. P. Withrow, (CEWC). Webster Co., jet SRs. 101 & 76, IM, VIII-5-1977, C. P. Withrow, (CEWC) . Wirt CO., SR. 5, 2.2mi. S. Elizabeth, IM 2J, VIII-15-1977, C. P. Withrow, (CEWC). 280 Pseudopolvdesmus collinus KENIUCKZ: Carter Co., 5mi. E. Grayson, US Rt. 460, IM, IV-19- 1961, R. L. Hoffman, (RIHC). MARYIAND: Calvert Co., 3.7mi. SW. jet SR. 2 & 231, nr Bowens, IF, XI-3-1957, H i l t o n & Jones, (RIHC). NORTH CAROLINA.: Gaston Co., 4.8mi. SW. Gastonia, CR. 1130, 0.2mi. W. CR. 1133, 12M, IV-29-1976, R. M. Shelley, (NCMH). Martin Co., 11.7mi. N. Rt. 903, nr Roanoke R., 4.4mi. NW. Oak City, IM, X- 19-1979, R. M. Shelley & P. T. Hertl., (NCMH). Rutherford Co., 17.7mi. NE. Rutherford, CR. 1732, l.Smi. N. CR. 1733, 3M IF, R. M. Shelley & MSM, (NCMH). OHIO: Montgomery Co., Taylorsville Dam, NE. side, IM, IV-10- 1969, G. A. Coovert, (EMNH). Pike Co., Pike S. P., IF, IV-1-1969, G. A. Coovert, (EMNH) ; IM, IV-12-1969, G. A. Coovert, (EMNH). Preble Co., Hueston Woods, IM, IV-16-1969, G. A. Coovert, (EMNH). SOUTH CAROLINA: Lancaster Co., 8.4mi. SW. Lancaster, CR. 296, CatavhaR., IM, V-1-1977, R. M. Shelley, (NCMH). TENNESSEE: Sevier Co., end Greenbrier Rd., at Porter Ck., parking area, 2M, V-8-1980, R. M. Shelley & MSM, (NCMH). VIRGINIA: Alleghany Co., Jordan Mines, 2M, III-12-1951, R. L. Hoffman, (RIHC); Floyd Co., SR. 8, circa 6mi. NW. Floyd, IM, V-30- 1951, R. L. Hoffman, (RIHC); Buffalo Mt., 3500', IM 2F, X-24-1958, R. L. Hoffman, (RIHC) ; Rocky Ihdb Park, 6mi. S. Floyd, IM, TV-13- 1951, R. L. Hoffman, (RIHC) ; 5mi. SW. Simpsons, IM, VII-2-1975, R. L. Hoffman, (RIHC). Franklin Co., 2mi. N. Algoma, 2M, IV-12-1958, R. L. Hoffitnan & R. E. Crabill, (RIHC); NE. Algoma, 2M IF, VII-15-1953, R. L. Hoffman & W. B. Newman, (RIHC). Henry Co., along ck. nr leatherwood, 3M 3F, R. L. Hoffman & R. E. Crabill, (RIHC) ; nr Ridgeway, 4M IF, XI-27-1961, R. L. Hoffiman, (RIHC). Montgomery Co., Roanoke R. Bluff, Imi. E. Shawsville, 2M 2F, TV-22-1956, R. L. Hoffman, (RIHC). Patrick Co., Pinnacles Dan, 4mi. WSW. Vesta, 2M IF, IV-19-1957, R. L. Hoffman & R. E. Crabill, (RIHC) ; Pinnacles of Dan, 4itii. WSW. Vesta, 4M 8F, IV-8-1959, Hubricht, (RIHC) ; Pinnacles of Dan, IV-22-1972, R. L. Hoffman & Fhight, (USNM), holotype of Pseudopolvdesmus collinus; Rock Castle Gorge, 2mi. W. Woolwine, 2M, X-7-1973, R. L. Hoffman, (RIHC). Pittsylvania Co., Staunton R. Bluff, 3mi. NW. Brights, 2M, 1-28-1951, L. Hubricht, (RIHC) ; Staunton R. Bluff, 3mi. NW. Motley, 4M 7F, 11-24-1951, L. Hubricht, (RIHC). Pulaski Co., nr Radford, IM, X-23-1975, R. M. Shelley, (NCMH). WEST VIRGINIA: Cabell Co., SR. 7/1, 2.4itii. from SR. 2, 7M, IV- 27-1977, C. P. Withrow, (CPWC). Mason Co., SR. 53, 3.Imi. E. Elizabeth, 3M 2F, VIII-13-1977, C. P. Withrow (CEWC) ; Mud Run, 0.75mi. off SR. 2, 4M 2F, XI-7-1977, C. P. Withrow, (CPWC) . McDowell CO., Coalwood, IM, X-23-1949, T. G. Vas, (RIHC) ; Windmill Gap, nr McComas, IM, IV-15-1973, W. McKenzie, (WASC). Monroe Co., top of mt. btwn Waiteville & Gap Mills, IM, VIII-14-1958, Hilton, (RIHC). Ralei^ Co., Grandview S. P., picnic area no. 2, IM, VI-8-1971, W. A. Shear, (WASC). Randolph Co., l-2mi. W. Bear Heaven Picnic Area on rd. to BickLe's Knob nr Alpena, IF, VI-12-1957, Highton, (RIHC). 281 Webster Co., N. Fork of Cranberry R., 5mi. SW. Three Forks, IM, VI- 18-1963, R. L. Hoffman, (RIHC). PseudoDolvdesmus serratus CANADA. ONTARIO: London, U , VII-24-1964, W. W. Judd, (FSAC) ; IF, V- 27-1964, W. W. Judd, (FSAC); * (Peters, 1954), QUEBEC: *Gaspe Peninsula, IM, 1883, R. Wells, (Chamberlin, 1920a); *Ottawa, 6 specimens (Chamberlin, 1920a); *Ottawa and Chelsea, 1918, F. Johansen, (Chamberlin, 1920a). UNITED STATES AIABAI'ôA: Lowndes CO., 8.4mi. NE. Fort Deposit, US Rt. 31, Sandy Ridge, IM, IV-19-1983, R. M. Shelley & P. D. Nader, (NCMH). ARKANSAS: C r a i ^ e a d Co., Jonesboro, 2F, IX-29-1966; IF, XI-26- 1966; 6M IF U , IX-29-1965; IF, IX-11-1966; 2M IF, XI-25-1966; U , IX-23-1966; IF, XI-3-10 1966; 2M, X-22-1966, IM 2J, IX-19-1966; IM, IX-19-1966; 3M, X-30-1966; IM, XI-26-1966, Hite, (FSAC); Jonesboro, 18M 18F 25J, III-17-1962, N. B. Causey, (FSAC). Jefferson Co., 2mi. E. Pine Bluff, 2M 2F, X-1954 & V-VI-1955, Kirkwood, (FSAC) . Hiillips Co., Helena, 31M 28F U , XI-29-1951, W. Benton, (FSAC); 57M 67F, IT- 1956, W. Benton, (FSAC). Poinsett Co., IM 2F, IV-17-1954, G. Doty, (FSAC). CONNECTICUT: New Haven Co., along Rt. C96, 6mi. S. Waterbury, U , VI-1-1975, R. M. Shelley, (NCMH) ; Woodbridge, U , V-17-1951, P. F. Bellinger, (RIHC) ; IM, IV-24-1958, C. L. Remington, (RIHC) ; Masoboro Sound, 3J, III-9-1978, A. B. Cary, (NCMH). INDIANA: Grant Co., IM 2F, 1951, C. J. Bushley, (FSAC). Greene Co., Richland Ck., IM, III-30-1952, (FSAC); 2M, III-25-1952, (FSAC). Monroe Co., Morgan, IM, X-20-1953, Owen, (FSAC) ; Morgan S. P., IM IF, IV-15-1952, (FSAC). Owen Co., Richland Ck., 2M 3F, III-20-1952, (FSAC). Porter Co., Indiana Dunes S. P., U , VIII-22-1946, H. S. Dybas, (FMNH). Tippecanoe Co., West Lafayette, 3M IF, IX-20-1956, L. Chandler, (FSAC). Vanderburg Co., Evansville, IF, VI-21-1943, H. S. P^tas, (PMNH). ILLINOIS: Adams Co., 5mi. NE. La Praire, IM, IV-15-1946, B. Patterson, (EMNH) ; IM, IV-19-1946, B. Patterson, (FMNH) ; NE. township, U , VIII-5-1943, B. Patterson, (FMNH). Calhoun Co., McNabb Hollow Cave, 3mi. W. Hardin, 5M IF, XI-25-1965, S. Peck, (FSAC). Champaign Co., SR. 49, bridge just N. Honer, 15mi. SE. Chaitpaign/Urbana, IM IF, IV-8-1967, D. H. Stansbery & C. B. Stein, (OSZM); SR. 49, 3.7mi. S. Ogden & US Rt. 150, IM IF 2J, VI-14-1983, C. P. Withrow, (CPWC). Cook Co., Palos Park, 2F, X-29-1938, M. A. Quait, (EMNH) ; Palos Park, Saganaskee Slough, woods, 5M, VI-lO-1961, J. Culbertson & W. Suter, (FSAC). Grundy Co., Praire, nr Lorenzo Will Co., 2M, VI-1-1962, (FSAC). Jackson Co., E. Carbondale, Holiday Inn, IM, IV-8-1967, C. B. Stein, (OSZM). Lake Co., Volo, Sayer Bog, sphagnum bog, IM 2F, W. Suter, (FSAC) ; Volo, Sayer Bog, sphagnum bog, IF 3J, rx-25-1959, W. Suter, (FSAC) ; Antioch Bog, IM 2F U , XI- 12-1959, W. Suter, (FSAC); Antioch, U , VI-3-1960, W. Suter, (FSAC). Rockford Co., Camp Grant, U , VI-2-1944, H. S. Dybas, (FMNH). 282 IOWA: Blackhawk Co., Grant Prairie nr Finchfort, IF U , IX-19- 1973, Nixon & Wilson, (FSAC). Boone Co., ledges S. P., 6itii. S. Boone, IM, V-19-1941, R. V. Chamberlin, (USNM), holotyP® Polvdesmus scopus; *3mi. W. Boone, IF, R. V. Chamberlin, (Chamberlin, 1942a). Story Co., Ames, IM, Spring 1941, R. V. Chamberlin, (USNM), holotype of Polydesmus planicolens. KANSAS: Johnson Co., 3mi. E. Eudora, IM IF, X-4-1966, D. Clark, (FSAC). KENIUCKï: Adair Co., Fanny Hollow Cave, IM, VII-28-1964, S. Peck, (FSAC). *Anderson Co., Tyrone, (Causey, 1963). Bell Co., Cumberland Mt. nr Cubage, IM, VI-13-1976, R. M. Shelley & J. K. Ettman, (NCMH) ; Pine Mt. S. P., Wildflower Garden Area, IM IF, 11 & 111-1976, J. H. Ettman, (NCMH). Carter Co., Carter Caves S. P., IM, X-10-1971, A. Weaver, (WASC). *Christian Co., Hopkinsville, (Causey, 1963). Estill Co., Eentud^y R. floodplain at Irvine, 2M 2F, V-9- 1954, R. L. Hoffman, (RIHC). *Fayette Co., Lexington, H. Garman, (Causey, 1963). *Fulton Co., Hickman, (Causey, 1963). Holmes Co., Lexirgton, IM, X-10-1958, (FSAC). Jackson Co., 3mi. NE. Sandgap, IM 2F, V-11-1967, T. G. Marsh, (FSAC). Jessamine Co., Indian Falls, 5mi. S. Nicholasville, IM, lV-16-1967, J. Reddell, (FSAC). McCreary Co., Cumberland Falls, IF, lV-20-1968, J. R. Tripp & F. J. Moore, (OSEM). Morgan Co., West Liberty, 2M, X-13-1961, O. Murphy, (FSAC). *Powell Co., Red R. Drainage, (Branson & Batch, 1971) ; Natural Bridge, ]M, lV-21-1968, (OSEM). Whitey Co., SR. 92, 27.8mi. W. US Rt. 25E, IF U , Vll-12-1983, C. P. Withrow, (CPWC). *Wolf Co., Red R. Drainage, numerous, 1970, (Branson & Batch, 1971). LOUISIANA: *Allen Par., (Causey, 1963) ; *US Rt. 190 at Kinder, IM 12F, XII-20-1958, K. F. Loomis, (Loomis, 1959). Ascension Par., Hope Villa, IF, X-3-1963, Calaway, (FSAC) ; nr. Praireville, 14M 5F U , 1-20-1966, W. Longest, (FSAC). Avoyelles Par., 0.7mi. N. Bunkie, 2M IF, V-24-1963, W. J. Harmon, (FSAC); Bunkie, IM, lV-17-1965, M. Kordisch, (FSAC) ; Evergreen, IM IF, Xll-23-1965, M. Kbrdisch & J. Walls, (FSAC) ; Evergreen, 4M, X-28-1965, M. Kordisch, (FSAC). Bossier Par., 7.7mi. W. Sawepta, IM, 111-12-1966, R. E. Tandy, (FSAC). Caddo Par., Shreveport, IF, 111-24-1962, N. B. Causey, (FSAC) ; 1.8 air mi. SE. Rodessa, on Black Bayou, Clyde Parker's Camp, 2M 2F, 111-11-1966, R. E. Tandy, (FSAC) ; 4M IF, 111-1966, R. E. Tanc^, (FSAC); Mira, 2M 4F, (FSAC). Caldwell Par., SR. 559, 7.4mi. N. Catahoula Par. line, IM, Xl-28-1965, R. E. Tandy, (FSAC). Catahoula Par., nr bridge on SR. 124, 15.2mi. S. Caldwell Par. line on SR. 559 & 124, IF, XE-28-1965, R. E. Tandy, (FSAC). De Soto Par., 2.5mi. N. Mansefield, Rt. 171, IF, 111-13-1966, R. E. Tandy, (FSAC). East Baton Rouge Par., Baton Rouge, 240 Richmond Ave., IM, Vlll-17- 1964, P. Harmon, (FSAC) ; Prophet Id., IM, IV-3-1965, R. E. Tandy, (FSAC). Highland Road, IM, IV-1-1969, (FSAC) ; IM 2F, XE-7-1964, (FSAC) ; Imi. ENE. KLeinpeter, 3M, lV-3-1971, D. A. & N. J. Rossman, (FSAC) ; Imi. ENE. KLeinpeter, 3M IF, lV-27-1971, D. A. Rossman, (FSAC) ; Baton Rouge, IM, 111-23-1962, K. Arnold, (FSAC). East Carroll Par., Transylvania, IM, lV-24-1971, D. A. Rossman, (FSAC) ; 2.7mi. S. Transylvania over Mississippi R. Levee, 4M 2F, Vll-11- 1966, R. E. Tandy, (FSAC) ; over Mississippi R. levee at Brunett, 3M IF, Vll-11-1966, R. E. Tandy, (FSAC). East Feliciana Par., 4.5mi. 283 WNW. Wilson, m , III-24-1971, P. Kimmich, (FSAC) ; T15 R4W Sect 26 & 35, IM IF, IV-17-1971, D. A. Rossman, (FSAC). Evangeline Par., Pine Prairie, IM, IV-19-1965, R. E. Tandy, (FSAC). *Iberia Par., (Causey, 1963). Jefferson Davis Par., Harahan, IM 2F, III-21-1951, G. H. Penn, (FSAC) ; Elton, IM IF, III-1-1965, C. Seal, (FSAC). Lafayette Par., lafayette, IM, VII-1965, N. Albert, (FSAC). *Lincoln Par., (Causey, 1963). Livingston Par., Springfield, IM 2F, Harman & Ecology Field Trip, (FSAC); 1.5mi. W. Springfield, 2M 2F, XII-19- 1964, R. E. Tan^. Madison Par., 1.5mi. W. 713.65, along spur, SR. 577, 2F U , IV-24-1971, Rossman, (FSAC); #6, IM IF, VII-10-1966, R. E. Tandy, (FSAC) ; 2 . 6 m . W . , 7mi. S. Tallulah, IM IF, IV-24-1971, D. A. Rossman, (FSAC). Morehouse Par., Cemin-a-haut S. P., 2F U , IV- 25-1971, D. A. Rossman, (FSAC) ; *Miller landing, IM, 1909, C. B. Moore, (Chamberlin, 1947). Natchitoches Bar., IIM 16F, 11-28-1956, (FSAC) ; Grand Ecore, 12M 2F, 11-15-1955, R. Hernandez, (FSAC) ; 6M 3F, 11-15-1955, C. Moréhead, (FSAC); College Farm, 3M, 11-14-1955, R. Hernandez, (FSAC) ; Northwestern S. C . , 7F, X-6-1954, WGL, (FSAC) ; Northwestern S. C. Farm, 5M IF, IV-11-1955, C. Mor^ead, (FSAC) ; 2F, II-23-1955, R. Hernandez, (FSAC) ; Old R . , 6M 5F, 11-23-1955, R. Hernandez, (FSAC) ; 5mi. S. Old R., 3M 5F U , 11-23-1955, R. Hernandez, (FSAC); Pacheco, 16M 15F 7J, II-7-1956, A. Coco, (FSAC); Kisatchie, 7M 5F, X-30-1954, W. T. Harman, (FSAC). Plaquemines Par., Boothville, 5M 5J, 1-22-1950, GHB, (FSAC). Pointe-Coiç)ee Par., Marganza btwn levee & river, 2J, VII-3-1965, R. E. Tandy, (FSAC) ; Mbrganza, btwn levee & R., IM 2F, VII-13-1965, R. E. Tandy, (FSAC). Rapides Par., 4.3mi. NE. Echo, SR. 1, 2M IF, 1-7-1965, R. E. Tandy, (FSAC) ; 0.3mi. SE. Lamatier, SR. 457, IM IF, XI-7-1965, R. E. Tandy, (FSAC) ; S.ami. E. Gardner, SR. 28, 2M 2F, XI-7-1965, R. E. Tandy & L. D. Wilson, (FSAC). Red R. Par., 4mi. NW., Imi. S. Coushatta, 3M IF, IV-10-1971, P. Kimmich, (FSAC) ; 1.7mi. N. Armistead, SR. 84, 2M 4F, III-13-1966, R. E. Tandy, (FSAC). Sabine Par., Hedges Gardens nr Many, IF U , III-26-1965, R. E. Tandy & Gates, (FSAC). St. Helena Par., Greensburg, 4M IF, 1-7-1939, N. B. Causey, (FSAC); 1.5mi. S. Grangeville, U , XI-1-1964, (FSAC) ; 8mi. E. Grangeville, 3J, XI-1- 1964, Romano, (FSAC). St. Landry Par., 4mi. SE. lebeau, 2M 2F, V-11- 1963, L. C. Binford, (FSAC) ; 14.2mi. SE. Lsbeau, US Rt. 71, IM, XI- 6-1965, R. E. Tanc^, (FSAC) ; 2mi. W. Melville, SR. 10, sawmill, 2M, III-24-1965, R. E. Tandy & G. E. Gates, (FSAC). St. Martin Par., 17mi. N. Henderson, W. levee of Atchafalaya Spillway, 2M IF, 11-20- 1965, R. E. Tancty & Romano, (FSAC) ; 4.6mi. S. Henderson, o n levee, 2.Imi. E. Diçjuis R d . , 2M 3F 2J, X-31-1965, R. E. Tandy, (FSAC). St. Tammany Par., T45, RllE, Sect 35, 2M, Rossman & Kimmich, (FSAC) ; Hickory, IM, XII-1-1950, G. H. Penn & E. W. Smith, (FSAC) ; 5mi. N. Hickory, 11-21-1965, Longest, (FSAC) ; Oaklawn, nr cypress Bayou, IM IF, V-2-1971, Kimmich, Rossman, Guthan, & Hebert, (FSAC) ; 2mi. W. on Bayou Liberty R d . , Sidell, 12M 8F, XI-10-1961, R. Hepburn, (FSAC) ; Hickory, 2M, IX-15-1950, G. H. Penn, (FSAC) ; Talisheek C k . , Talisheek, 4M, XII-12-1965, L. D. Wilson, (FSAC) ; 3mi. E. Pearl R . , IM IF, II-5-1966, La Val, (FSAC) ; 0.8mi. NNE., 0.6mi. ENE. Mondeville, SR. 1088, IF, III-6-1971, D. A. Rossman ard Herp. Class, (FSAC). Tangipahoa Par., 3.imi. S. Hammond, IM, XII-5-1964, Ecol. Class, (FSAC) ; Imi. N. Hammond, IM, XII-23-1964, R. E. Tandy, 284 (ESAC); Qiapp^jeela Ck., SR. 40, 2M IF, XII-23-1964, R. E. Tandy, (FSAC) ; l.OSmi. E. Holton, 8R. 16, IM, XII-23-1964, R. E. Tandy, (FSAC); 2.4mi. W. Independence on Natalbany R., Rt. 40, IF, R. E. Tandy, (FSAC) ; flooc%)lain, nr Robert, IM, X-26-1976, J. E. & M. R. Cooper, (NCMH). Tensas Par., Tensas R., 2M, X-5-1953, B. Stanberxy, (FSAC) ; Yucatan Lake, Newallton, 2M, IV-16-1966, R. E. Tarx^, (FSAC). Terrebonne Par., Schriever, 2M, X-1954, H. L. Whitten, (FSAC) ; Schriever, 3M, winter 1954-1955, H. L. Whitten, (FSAC) ; IM IF, 1954, H. L. Whitten, (FSAC); IM, VII-1953, H. L. Whitten, (FSAC); Houma, 5M 7F 17J, V-1954, H. L. Whitten, (FSAC). Vermilion Par., Kaplan, 0.2mi. S. SR. 14, SR. 35, IM IF, III-31-1967, R. E. Tanc^, (FSAC). Vernon Par., 2.4mi. ENE. Pitkin, SR. 113, 2M 2F, XI- 20-1966, R. E. Tanc^. Washington Par., Angie, V-8-1958, J. L. Crain, (FSAC) ; 2M, V-2-1958, J. L. Crain, (FSAC); 5M IF, IV-23-1958, J. L. Crain, (ESAC) ; 3M 3F, IX-15 to 26-1958, J. L. Crain, (FSAC) ; 7mi. S. Angie, X-23/24-1958, IF, J. L. Crain, (FSAC) ; 2M, XI-16-1958, 7th Grade Science Club, (FSAC); Bogalusa, IM, 11-24-1953, (FSAC); 6mi. SW. Bogalusa, IM 2F, 1-21-1965, N. B. Causey, (FSAC); Days Methodist Church, Rt. 438, l.lmi. N. Hackley, IM IF, R. E. Tandy, (FSAC); lawrence Ck., SR. 10, Imi. E. Franklinton, IM, III-2-1969, B. G. Mbrizot, (FSAC); 0.7mi. S. Angie, 34M 25F 2J, X-23/24-1958, J. L. Crain, (FSAC) ; 22M 16F, XI-1958, J. L. Crain, (FSAC) ; 4M IF, (FSAC). West Baton Rouge Par., over Mississippi R. levee at Addis, 3M, X-23- 1965, R. E. Tarx^, (FSAC). West Carrol Par., 2.5mi. W. Kilboume on Macon Bayou, IM, VII-31-1966, R. E. Tandy, (FSAC). West Feliciana Par., 6mi. S. & E. jet SR. 418 & Old R. Levee across from Batchelor, 7M IF U , (FSAC) ; Tunica, IF, IV-29-1972, M. A. Tidwell, (FSAC) ; nr S. gates Tunica Prison, IF, 1970, (FSAC) ; Bains, 4M IF 3J, 1-20- 1965, (FSAC). MARYLAND: Allegany Co., Dans Mt. Recreational Area, circa Imi. E. lonaconing, IF, XI-3-1956, Highton, (RIHC) ; Imi. W. crest Green Mt., US Rt. 40, circa 15mi. E. Cumberland, IV-27-1957, IM, Ellis, MacLeod, & Hilton, (RIHC). Anne Arundel Co., *nr Mayo, III-29-1916, H. W. Fowler, (Chamberlin, 1947); SR. 450, 3.Imi. E. jet US Rt. 301 at Patuxent R., IM IF, IV-25-1958, Hilton, Messel, & MacLeod, (RIHC). Calvert Co., 3.7mi. SW. jet SR. 2 & 231, nr Bowens, IF, XI- 3-1954, Highton & Jones, (RIHC); Battle Ck. Swanp at SR. 506, circa 2mi. S. Prince Frederick, 2F, VI-21-1957, Highton & Barry, (RIHC) ; nr Huntingtown on SR. 510, 1.6mi. S. SR. 262, 3M 2F, III-16-1958, Hilton & Barry, (RIHC). Caroline Co., nr Hillsboro, picnic area on SR. 404, 1.6mi. W. SR. 312, IM, III-16-1958, Hilton & Jones, (RIHC); IF, HI- 3 1-1957, Highton, (RIHC); 0.6mi. SW. Baltimore C o m e r on SR. 312, III-31-1957, (RIHC); 3mi. S. Ardersontown on SR. 313, 3mi. S. SR. 404, IF, III-16-1958, Highton & Jones, (RIHC). Charles Co., Indian Head, IM 2J, IV-12-1953, W. Harman, (FSAC). Dorchester CO., 2.5mi. N. Eldorado on SR. 313, III-16-1958, Highton & Jones, (RIHC) ; 3mi. S. Preston, IF, III-16-1958, H i l t o n & Jones, (RIHC). Frederick Co., Cunnin^iam Falls S. P., IM, IV-28-1958, H i l t o n fit E. MacLeod, (RIHC) ; Cunnin^iam Falls S. P . , nr Thurmcnt, 1000-1300', 2F 2J, XII-6-1956, R. Highton, (RIHC); 16M 17F, VIII- 20-1957, Hilton, (RIHC); 15M 8F, IV-28-1958, R. Hilton fit E. MacLeod, (RIHC) ; vicinity Cummingham Falls S. P., 5M 8F, VI-7-1957, 285 Barry & Hilton, (RIHC) ; 1.4mi. NW. Yellow Springs Catoctin Mt., IM 3F, VI-22-1958, Hiÿiton, (RIHC) ; 5M IF, V-14-1958, H i l t o n , (RIHC) ; foot of E. slope Sugarloaf Mt., IM, X-6-1951, Hilton & Barry, (RIHC) ; Frederick & Washington Cos. US Rt. 40 at county line, crest of South Mt., IM 2F, X-6-1957, Hilton & Barry, (RIHC). Garrett Co., circa 3mi. W. Bloomington on SR. 195, IM, XI-4-1956, Hil t o n , (RIHC). *Harford Co., Swan Ck., V-20-1914, H. W. Fowler, (Chamberlin, 1947). *Mbntgomery Co., Bethesda, 2 specimens, XI-4- 1949, E. Beck, (Chamberlin, 1947). Prince Georges Co., IF, V-3-1953, W. Harmon, (FSAC) ; 1.9mi. W. Glendale, 2M, IV-25-1958, Hilton, MacLeod, & Wessel, (RIHC) ; along NW. branch of NH. Ave., circa 2mi. NW. College Park, IM IF, IV-5-1958, Hilton, Ellis, & MacLeod, (RIHC) ; jet Adelphi Rd. & Metzerott Rd., circa 2mi. NW. College Park, IF, IV-5-1958, Hilton & MacLeod, (RIHC); Imi. E. Lanham, IM 2F, IV-14-1958, Hilton & MacLeod, (RIHC) ; University of Maryland Campus, College Park, IF, IV-26-1958, Hi<ÿïton, (RIHC) ; 2mi. NW. Piscataway, SR. 210, 2F, XI-24-1957, Hilton, (RIHC). Queen Annes Co., 4.3mi. NE. Centreville, US Rt. 213, IM IF, IV-14-1957, Highton, (RIHC) ; jet SR. 18 & US Rt. 50, Queenstown, 4M 4F, III-31-1957, Hilton & Barry, (RIHC). St. Marys Co., 4.5mi. NW. leonardtown, SR. 5, 2F, IV-7-1957, Hilton, (RIHC). Someret Co., 4mi. lŒ. Crisfield, 2F, XI-13-1956, Hilton, (RIHC). Talbot CO., 3.6mi. S. jet SR. 404, US Rt. 50, 5M 5F, IV-14-1957, Barry & Highton, (RIHC); 6.3ml. SSW. Easton, SR. 333, IM, IV-14-1957, Barry & H i l t o n , (RIHC) ; 2mi. N. Traope, 11-17-1956, Hilton, (RIHC). Washington Co., Gathland S. P., IM 3F, III-7-1958, Hilton, (RIHC). MASSACHUSETTS; Middlesex Co., MCZ Reserve nr Bedford, IM IF, X-6-1968, W. A. Shear, (WASC). MICHIGAN: *Lake Douglas, University of Michigan Biological Station, VII/VIII-1913, G. F. Sutherland, (Chamberlin, 1914). MINNESOTA: *Winona Co., Winona, 1886/1887, J. M. Holzinger, (Bollman, 1893). *Fort Snelling, 1886/1887, W. D. Messrs. & G. M. Howe, (Bollman, 1893). MISSISSIPPI: Adams Co., nr St. Catherine Ck., 2mi. S. Natchez, 2M, L. Hubricht, (RIHC). George Co., jet Pascagoula R. & SR. 26, IM, IV-11-1966, L. D. Wilson, (FSAC). Hinds Co., Jackson, 2M 3F, XII-11- 1962, G. Catching, (FSAC). Jefferson Co., 6.7mi. W. Fayette, 3.8mi. ESE. Chunk Hill off Natchez Trace, 2M, (OSZM). Lincoln Co., 12mi. E. Brookhaven, IM, XII-25-1953, N. B. Causey, (FSAC). Oktibbeha Co., state line, 2M, XI-16-1959, Shoford, (FSAC) ; lOmi. S. Starkville, œ t t o n field nr Craig Springs, IM, X-8-1979, W. H. Cross, (NCMH) ; IM, X-15-1979; 2M, XII-18-1979; IM, XII-31-1979, (NCMH); State College, 2M, VI-16-1959, Shoford, (FSAC). Pearl R. Co., llmi. E. Bogalusa, floodplain, IF, 1-21-1965, N. B. Causey, (FSAC); Pearl R., SR. 26, 2M, III-2-1969, B. G. Mbrizot, (FSAC). Pike Co., Percy Quinn S. P., 4.5mi. NW. Magnolia, 4M 5F U , VI-28-1959, (FSAC). Panola Co., 6mi. SW. Como, cotton field, IM, IX-13-1979, W. H. Cross, (NCMH). Webster Co., Mathiston, IM, IV-4-1960, J. Brooks, (FSAC). Wilkinson Co., circa 1.0 air mile SE. Fort Adams, IM, III-30-1974, D. A. Rossman, (FSAC). MCSSOUEd: Franklin Co., 3mi. ESE. Sullivan, camp Cave, IM, V- 286 2-1982, J. E. Gardner, (CPWC). St. Louis Co., IM 4F, III-19-1953, W. Harman, (FSAC). NEW JERSEY: *Camden Co., Delaware R., III-27-1915, (Chamberlin, 1947). *Westmont, IM, X-I-1916, (Chamberlin, 1947). Soiærset Co., Middlebush, IM 4F, III-27-1962, Whitehead, (RIHC). NEW YORK: Albany Co., l.Omi. from Altamount, IF, VIII-25-1936, (FSAC). Genesee CO., Bergen Swamp, IM U , VI-3-1957, R. L. Hoffinan, (RIHC). Greene CO., along SR. 23A nr Jarrett, IF, VI-15-1975, R. M. Shelley, (NCMH) ; along North Lake Rd., 3.0mi. N. SR. 23A at Haines Falls, IM IF, VI-15-1975, R. M. Shelley, (NCMH) ; Diamond Notch nr Lanesville, 3M, VIII-19-1978, R. M. Shelley & T. L. McCabe, (NCMH) ; *Catskill, several M. & F., Khi^t, (Chamberlin, 1947). Monroe Co., Menden Ponds, 2M, X-7-1954, W. B. Muchmore, (RIHC). Onondage Co., Jully Forest, Syracuse, 4M 13F IIT, VIII-8-1960, L. C. Stegen, (FSAC). Ringwood Co., IM IF, VII-19-1950, R. E. Crabill, (RIHC). Tcarpkins Co., 2M, IV-29-1974, T. L. McCabe, (NCMH); East Lansing, IM 2F, X-22-1963, EMR, (RIHC); Ithaca Fish Hatchery, IM, IV-21-1955, D. J. Pirone, (RIHC); Ithaca Six Mile Ck., IM IF, VIII-18-1957, D. J. Pirone, (RIHC) ; 2M IF, V-14-1959, R. Hutt, (RIHC) ; *Ithaca, IV-27- 1936, several specimens, J. W. Rehn, (Chaniberlin, 1947). Ulster C o . , 5mi. W. New Paltz, IF U , V-11-1958, K. K. Asplund, (RIHC) ; nr Poenicia, IM, VIII-18-1978, R. M. Shelley, (NCMH). NORTH CAROLINA: Ashe Co., 2.5mi. E. Jefferson, along SR. 88- 16, 3M, VII-9-1977, J. B. Funderburg, (NCMH). Avery CO., Linville, Grandfather Mt., 5000', 7M 2F, X-14-1939, N. B. Causey, (FSAC). Beaufort Co., Goose Ck. S. P., 2M IF, X-17-1979, R. M. Shelley & P. T. Hertl., (NCMH). Bertie Co., 7.2mi. NW. Windsor, SR. 305, 3mi. N. US Rt. 13, U , IX-12-1983, R. M. Shelley, (NCMH). Brunswick Co., ravine on Cape Fear R., 16.2mi. N. Bolivia, 4M, X-16-1979, R. M. Shelley & P. T. Hertl., (NCMH) ; 4.8mi. NNW. Maco, ravine along Bryant Mill Ck. off Cape Fear R., nr CR. 1421, IF, VII-17-1978, D. L. Stephan, (NCMH). Carteret Co., 5.8mi. N. Cape Carteret, CR. 1004 at Hadnot Court, IM, XI-14-1982, D. L. Stephan, (NCMH). Craven Co., 8mi. WNW. New Bern, US Rt. 70 at Batchlor's Ck., 3F, VI-8-1976, R. M. Shelley & M. Filka, (NCMH). Cumberland Co., Fayetteville, Clark Park, 2M, IX-29-1975, R. M. Shelley & J. C. Clamp, (NCMH). Currituck Co., Imi. S. Sligo, IM IF, IV-4-1947, R. L. Hoffinan & KLeinpeter, (RIHC). Dare CO., along SR. 1206, Nags Head Woods, 1.3itii. NW. Kitty Hawk, IM, X-17-1979, R. M. Shelley & P. T. Hertl., (NCMH); Buxton, 7M 2F U , XI-19-1980, D. L. Stephan, (NCMH). Eurham Co., North Carolina State University Forest, 14.Imi. N. Durham, CR. 1614, 1.3mi. W. CR. 1613, U , V-30-1972, R. M. Shelley, (NCMH); N. Durham along Eno R., W. US Rt. 501, IM, V-10-1975, R. M. Shelley, (NCMH). Edgecomb Co., Taraboro, 3J, IX-13-1973, R. M. Shelley, (NCMH). Franklin Co., 3.4mi. W. Youngsville, CR. 1100(1147), 0.2mi. E. CR. 1137, IM IF, X-10-1972, R. M. Shelley, (NCMH). Gaston Co., 8.5mi. SW. Gastonia, CR. 1104, 0.3mi. S. CR. 1115, IM, IV-29-1976, R. M. Shelley, (NCMH); 4.8mi. NE. Gastonia, CR. 2201, 1.5mi. N. Rt. 7, IF, VII-7-1976, M. Filka & W. Thomas, (NCMH) ; 4.5mi. S. Bessermer City, CR. 1125, jet CR. 1106, 2J, X-15-1976, M. Filka & G. Wicker, (NCMH). Gatzes Co., 2.3mi. NW. Eure, 1200, O.lmi. W. jet. US Rt. 13, U , IX- 16-1971, R. M. Shelley, (NCMH). Granville Co., 6.6mi. E. Creedmore, 287 CR. 1705, W. CR. 1706, IM IF, X-10-1972, R. M. Shelley, (NCMH). Greene Co., 13 at CR. 1328, Imi. N. Snow Hill, IM, X-19- 1979, R. M. Shelley & P. T. Hertl., (NCMH). Halifax Co., 3.Uni. ENE. Hollister, CR. 1323, O.Smi. N. CR. 1002, Medoc Mt., 2^' 6F, R. M. Shelley, (NCMH). Jackson Co., Coyle Farm, l.Smi. SW. Webster, 2200', IM, IV-30-1970, F. A. Cqyle, (NCMH) ; IM, 1-4-1970, (NCMH). Jones Co., Hunter's C k . , at SR. 58, 1.5mi. N. CR. 1102, IM, III-27-1978, R. E. Ashton, (NCMH); roadside table on SR. 17, Imi. S. CR. 1114, 3mi. N. Maysville, 8M IF, X-16-1979, R. M. Shelley & P. T. Hertl., (NCMH). Johnston Co., 12.4mi. E. Clayton, CR. 2117, 0.4mi. SE. SR. 42, 2M IF, X-27-1971, R. M. Shelley, (NCMH); 10.6mi. WSW. Clayton, SR. 50, 0.3mi. S. Wake Co. line, IM, X-3-1972, R. M. Shelley, (NCMH) ; 11.Imi. SW. Smithfield, CR. 1330, 0.2mi. W. SR. 210, IM, XI- 14-1973, R. M. Shelley, (NCMH). Lee Co., 10.8mi. NE. Sanford, US Rt. 1, 0.3mi. N. CR. 1434, U, IX-14-1972, R. M. Shelley, (NCMH) ; 3.8mi. NE. Sanford, CR. 1508, O.lmi. E. CR. 1002, 3J, IX-14-1972, R. M. Shelley, (NCMH). Mitchell Co., Altapass, 3M 3F, V-20-1956, Keeton, Lurd, & R. L. Hoffman, (RIHC). Montgomery Co., 12itii. NW. Troy, SR. 109, 1.2mi. S. CR. 1300, U , VII-19-1972, R. M. Shelley, (NCMH). N a ^ Co., 2mi. SW. Spring Hope, US Rt. 64 at Tan R., IM, XI-4-1973, R. M. Shelley, (NCMH). New Hanover Co., 8.5mi. S. Wilmington, US Rt. 421, 1.2mi. S. SR. 132, 5M, IV-10-1975, R. M. Shelley, (NCMH). Northairpton CO., 5.2mi. WSW. Gaston, CR. 1220, 1.2mi. S. SR. 46, IM 3F, V-4-1973, R. M. Shelley, (NCMH). Onslow Co., 6mi. N. Swansboro, CR. 1434, Webb Ck., 2.Imi. NW. CR. 1449, IM IF, X-15-1977, R. M. Shelley & J. C. Clamp, (NCMH). Orange Co., Duke Forest, 4.9mi. NE. Chapel Hill, CR. 1731, 0.6mi. W. CR. 1734, IM IF, VII-7-1972, R. M. Shelley, (NCMH) ; 5.6mi. S. Hillsborough, SR. 86, 0.3mi. S. CR. 1723, 2M IF, X-26-1974, R. M. Shelley, (NCMH). Perquimans Co., along CR. 114, 0.3mi. vp Goodwin C k . , 2F, XII-30-1979, J. Perry, (NCMH) ; Imi. E. Winfall, SR. 37, 2M 2F, IV-5-1947, R. L. Hoffinan & KLeinpeter, (RIHC). Pitt Co., Imi. W. SR. 903, SR. 43, 1.3mi. W. Greenville, lOM 13F, X-19-1979, R. M. Shelley & P. T. Hertl., (NCMH). Randolph Co., S.Omi. SW. Asheboro, Cedar Rock Mt. at radio tower, 1.3mi. from CR. 1143, 1.4mi. jet CR. 1142, U , III-22-1972, R. M. Shelley, (NCMH) ; 12.0mi. WSW. Seagrove, 1105 at IMiarrie R., 3M, IV-24-1973, R. M. Shelley, (NCMH). Sampson Co., nr Roseboro, CR. 242, 0.9mi. N. CR. 1002, IM IF, III-24-1975, M. R. Ctooper, (NCMH) ; SR. 1107 at SR. 1100, ravine on Wild Cat Ck., 3mi. SSE. Harrells, 14M 2F, R. M. Shelley & P. T. Hertl., (NCMH); SR. 1103 at Black R., 7.8mi. SSE. Harrells, 5M, X-15-1979, R. M. Shelley & P. T. Hertl., (NCMH) ; CR. 701, O.lmi. E. CR. 1724, 4mi. E. Newton Grove, IM IF, X-19-1979, R. M. Shelley & P. T. Hertl., (NCMH) ; Clinton Six Run Ck., IM IF, II- 18-1948, (RIHC). Stokes Co., CR. 2009, 0.2mi. E. CR. 2010, Hanging Rock S. P., U , V-30-1973, R. M. Shelley, (NCMH). Vance Co., 4mi. N. Henderson, SR. 39, Imi. S. CR. 1308, 3M 5F, V-16-1973, R. M. Shelley, (NCMH). Wake Co., 8mi. NW. Ralei^, Ifinstead Park, CR. 1650, 0.5mi. SE. jet CR. 1647, U , IV-19-1972, R. M. Shelley, (NCMH) ; Ifiistead Park, 3M, III-27-1973, R. M. Shelley, (NCMH) ; U , VIII-2- 1972, R. M. Shelley, (NCMH) ; 4J, IX-20-1972, R. M. Shelley, (NCMH) ; 4.4mi. NW. Raleicfi, Umstead Park, IM 4F, X-11-1971, R. M. Shelley, (NCMH) ; 2M 15F, R. M. Shelley, (NCMH) ; Ifiistead Park, CR. 1649, 288 l.Omi. from CR. 1664, 2M 3F, V-2-1972, R, M. Shelley, (NCMH); Yates Pond, IM IF, VI-29-1972, C. Liébrandt, (NCMH); Ralei^, CR. 909, Northclift Dr., 2M IF, III-9-1974, R. M. Shelley, (NCMH); 4.25mi. S. Cary, O.lmi. NW. SR. 1300, l.Omi. from jet SR. 1009, IF, VI-16- 1975, J. C. Claitp, (NCMH) ; Bayleaf, 2M IF, X-31-1975, D. S. Lee, (NCMH). Watauga Co., Blowing Rock, Gofonth Rd., O.Smi. N. US Rt. 321, IM, X-16-1971, R. M. Shelley, (NCMH); 3mi. NNE. Boone, CR. 1326, 0.3mi. E. CR. 1325, IM, IV-25-1974, R. M. Shelley, (NCMH) ; Valle Ctucis, SR. 194, 0.6mi. S. CR. 1113, IM, X-11-1975, J. C. Clanp, (NCMH) ; US Rt. 421, 4mi. N. Vilas, 3M 2F, VII-11-1962, R. L. Hoffinan (RIHC). Wilkes Co., 8.5mi. SE. Wilksboro, CR. 2425, 0.2mi. NW. CR. 2428, 2M, VII-15-1976, M. Filka & F. Scott, (NCMH). OHIO: Adams Co., Cedar Forks (3ave, IM, 1-26-1980, M. Flynn & H. H. Hobbs III, (CPWC) ; Stout Run, IF, V-26-1967, (OSEM). Ashland Co., Londonville, 5M 2F U , VIII-1951, L. Gray, (FSAC). Ashtabula Co., Windsor, IM IF, VI-21-1955, W. B. Muchmore, (RIHC); T537, 0.7mi. S. SR. 322, 2J, VII-5-1984, C. P. Withrow, (CPWC); 0.4mi. E. SR. 84, CR. 22, 3J, VII-4-1984, C. P. Withrow, (CPWC). Bulter Co., Oxford, 12M 4F, X-1927, R. A. Hefner, (CPWC). Chairpaign CO., Kiser Lake S. P., IM, IV-27-1969, J. T. Wehner, (EMNH). Clermont Co., Cincinnati Nature Center, IF, IV-14-1979, JEM, (C3WC). Ctolurribiana Co., Zepemick Wildlife Area, 2F, VII-4-1984, C. P. Withrow, (CPWC). Coshocton Co., CR. T219, 0.4mi. off SR. 643, 2J, VI-25-1984, C. P. Withrow, (CPWC). Fairfield Co., Bamébey Center, 2M 2F, IV-27-1984, C. P. Withrow (USNM), neotype of Polydesmus olaucescens; SR. 664, O.lmi. N. SR. 312, IM, IX-12-1981, C. P. Withrow, (CPWC). Geauga Co., 0.4mi. W. Bundysburg, IM 2F 6J, VII-5-1984, C. P. Withrow, (CPWC) ; Holden Arboretum, imi. SE. T80 on CR. 5, IM, VII-5-1984, C. P. Withrow, (CPWC). *Gallia Co., Vinton, VI-19/23-1901, Morse, (Morse, 1904). Greene Co., Clifton Gorge, IM, IX-5-1967, G. A. COovert, (EMNH) ; jet US Rt. 75 & CR. 80, U , VI-27-1984, C. P. Withrow, (CPWC). Hamilton Co., Cincinnati, 2M 2F, VII-8-1957, W. B. Muchmore, (RIHC). Hi^iland Co., Dancing Cave, 2M, 11-27-1982, H. H. Hobbs III, (NCMH) ; 2mi. E. Hillsboro, IM, X-16-1960, (OSEM). Hocking Co., Neotonna, IM, IV-25-1946, Walker, Green, & Bailey, (RIHC). Holmes Co., CR. 58, 1.2mi. S. SR. 62, 3F U , VII-3-1984, C. P. Withrow, (CPWC). Jefferson Co., SR. 152, 3.0mi. E. SR. 213, 3F 2J, VII-4-1984, C. P. Withrow, (CPWC). Knox Co., Knox Wood Nature Preserve, 2M 3F U , VII-3-1984, C. P. Withrow, (CPWC). Lake CO., CR. 217, 0.3mi. S. Grand R., U , VII-5-1984, C. P. Withrow, (CPWC) ; *Painesville, IX-1918, E. R. Kalmbach, (Chamberlin, 1920b). Licking C o . , Burlirgton Township, SR. 657, Imi. SE. SR. 62, IF, V-I-1965, F. J. Moore, (OSEM). Medina CO., CR. T126, 0.3mi. E. T197, 2M IF 5J, VII-6-1984, C. P. Withrow, (CPWC). *Montgomery Co., several mi. N. Dayton, summer 1963-1964, J. M. Ramsey, (Ramsey, 1966); Dayton, IM, IV-3-1969, G. A. Coovert, (EMNH) ; IF, G. A. Coovert, (EMNH) ; Imi. E. Vandalia, IV-8-1969, G. A. Coovert, (EMNH). Ottawa Co., CR. 135, 0.2mi. S. T218, 3F, C. P. Withrow, (CPWC). Pike Co., Pike S. F . , IF, IV-12-1969, G. A. Coovert, (EMNH) ; Frost Cave, IM, VI-24-1980, M. Flynn & H. H. Hobbs III, (CPWC). Summit Co., Virginia Kendall S. P., 2F, VII-6-1984, C. P. Withrow, (CPWC). Tuscarawas Co., CR. 82, 0.5mi. N. Zoar, 2J, VII-3-1984, C. P. Withrow, (CPWC). Warren Co., 289 Findley Reserve, IF, VI-25-1969, G. A. Coovert, (CMNH) ; IF, VI-30- 1969, G. A. coovert, (EMNH) ; Upper Findley Farm, IF, VI-26-1969, G. A. Coovert, (EMNH). Washington Co., Marietta, IM, 1929, R. A. Hefner, (CPWC); Coal Run, IM, V-27-1943, H. E. Story, (PMNH); IF, VI-8-1943, H. E. Story, (FMNH). Wayne Co., WAA Cabin Wooster, IM, IX-3-1963, W. A. Shear, (WASC) ; Wooster Funk Hollow, 2M IF U , XI- 1961, W. A. Shear, (WASC). PENNSYIVANIA: Bedford Co, 3mi. S. Bedford Valley, 12M 5F, V- 25-1958, R. Hiiÿiton & I. Huber, (RIHC) ; New Paris, sink no. 4, IF, V-18-1963, R. E. Graham, (FSAC) ; New Paris Sink, 2M, 1961, N. D. Richmond, (RIHC) ; circa 3mi. S. Bedford Valley, IM, V-25-1958, R. Hilton & I. Huber, (RIHC). Berks Co., Shafer Cave, (loomis, 1939) ; Morgantown, IM IF, III-27-1971, W. B. Muchmore, (WASC). Bradford Co., 4.5mi. S. Pfonroeton, US Rt. 220, 3M, VI-8-1958, Hilton, (RIHC), neotype of Polvdesmus pensvlvanicus. Chester Co., Phoenixville, IM 3F, X-15-1960, M. S. Wilson, (FSAC). *Dela:varc Co., Cobbs Ck., III-16-1912, 1 specimen, H. L. Mathis Jr., (Chamberlin, 1947). *McKean Co., Port Alleghany, VIII-1904, 4 specimens, T. D. Keim & H. W. Fowler, (Chamberlin, 1947). *Perry Co., Cove Mt., IX- 25-1914, IF, W. Stone, (Chamberlin, 1947). *Philadelphia Co., Holmesburg, VI-25-1910 & III-23-1913, H. W. Fowler, (Chamberlin, 1947) ; *Fairmont Park, III-25-1911, 2 specimens, H. L. Mathis, Jr., (Chamberlin, 1947). Seward Co., sawdust pile, 3J, VII-24-1959, W. Suter, (FSAC). *Bristol, IF, IV-27-1913 W. Stone & Edgehill; IV-16- 1893, several M & F, P. Nell, & York Furance, V-1906, W. Stone & H. W. Fowler, (Chanberlin, 1947). SOUTH CAROUNA; Richland Co., Pocahontas, 2M IF U , Spring 1963, Simpkins, (RIHC). TENNESSEE: Arxlerson Co., Hole Cave, IM, II-5-1965, J. Payne, (RIHC) ; 4mi. N. Clinton, 2M, (WASC). Davidson Co., non-cave, Imi. S. city limits, 200 yds. W. US Rt. 431, 2F, VI-8-1957, T. C. Barr, (RIHC). Haywood Co., Brownsville, 25M lOF, X-28-1961, M. Lea, (FSAC). Hunpreys Co., US Rt. 140, 3.2mi. W. SR. 13, 2M 5F, VII-5- 1983, C. P. Withrow, (CPWC). Obion Co., Reelfoot Lake, 2M, IV-28- 1956, F. J. Etges, (FSAC). Putnam Co., Cookeville, Well Cave, IF, XI-19-1960, (FSAC). Rutherford Co., US Rt. 24, 3.9mi. E. Rt. 64, 2F U , VII-7-1983, C. P. Withrow, (CPWC). Sevier Co., GSMNP, Cades Cove, Giant Poplar Trail, 2000', U , IX-13-1953, H. S. Dybas & S. A., (FMNH). TEXAS: *Harris Co., (Causey, 1952b). Jasper Co., Jasper H i ^ School, 3M 2F, III-5-1962, Jasper Sci. Club, (FSAC); 4M 4F, III-1/6- 1962, J. Rand, (FSAC); Jasper, 3M 3F, 1-15-1962, (FSAC); IM IF, I- 26-1962, Science Club, (FSAC). Tyler Co., Woodville, IM, II-8-1962, R. Timbrook, (FSAC). VERMONT: Bennington Co., Big Equinox Summit, 2M IF, X-18-1968, W. A. Shear, (WASC). *Washington Co., Barre, IF, VI-10-1950, E. Beck, (Chamberlin, 1951b). Windham Co., Westminster, IM 3F 3J, VI-24 to VII-16-1960, M. S. Wilson, (FSAC) ; IM 2F, VIII-8-1960, M. S. Wilson, (FSAC) ; 9M 4F 3J, IX-12-1960, M. S. Wilson, (FSAC). VIRGINIA: Albemarle Co., Charlottesville, IM 2F, V-22-1947, R. L. Hoffman (RIHC) ; IF, V-3-1960, (FSAC) ; Sugar Hollow, 1948, 2M, (RIHC); Saddle Hollow, IM IF, III-1948, (RIHC); Scottsville, lOM IF, 290 V-10-1947, R. L. Hoffman (RIHC). Alle^iany Co., Lowmoor, IF, VI-1- 1947, R. L. Hofj&nan, (RIHC) ; Stulls Cave, Richpatch, 2M, VI-16-1947, (RIHC) ; McGraw's Gap, 3mi. NW. Clifton Forge, IM 2F, IV-13-1947, R. L. Hoffinan, (RIHC). Augusta Co., Huirpback Mt . , IM, VI-17-1947, (RIHC) ; 4.Imi. b y rd., SW. of Firetower turnoff at Reddish Knob, IM 2F, IV-13-1947, Hilton, Jones, & Littleford, (RIHC). Botetourt Co., "Sugarland", SW. side of j^ple Orchard M t . , IM, V-27-1962, (RIHC) ; W. side of i ^ l e Orchard M t . , circa 3500*, IM, VII-8-1972, R. L. Hoffman, (RIHC). Buchanan Co., Grimleysville, IM 3F, VI-30-1951, R. L. Hoffman & Newman, (RIHC); 2.7mi. SW. Vansant, IM, VII-1-1951, R. L. Hoffman & Newman, (RIHC). Caroline Co., Milford, 2M 2F, IV-17- 1968, D. Pugh, (RIHC). Chesterfield Co., Pocahontas S. P., IM, VI- 18-1971, R. E. Woodruff, (FSAC), neotype of Polvdesmus serratus. Clarke Co., SE. Millwood, US Rt. 50, 1.2mi. SE. jet SR. 225, 2M 2F, VI-12-1957, Hilton & Barry, (RIHC); 1.7mi. N. Ashby nr Paris, V-10- 1958, Hilton, Gerd, & Wohler, (RIHC). Craig Co., top of Pott's Mt., E. of Point C k . , 3M IF, III-13-1962, R. L. Hoffinan, (RIHC); Skyline Drive, St Mary's Rock, Shenandoah N. P., 4M 4F U , VIII-1959, E. Bullington, (FSAC). Dinwiddle Co., along 613 at Butterwood Ck., 7mi. NW. McKsnney, IF, VI-28-1978, R. M. Shelley & W. B. Jones, (NCMH). Fairfax Co., Merrifield, IM, V-21-1953, W. Harman, (FSAC). Frederich Co., 5.4mi. S. Hayfield, SR. 600, 4M 2F, V-10-1958, N. & V., (RIHC). *Giles CO., Spruce Run Cave, several specimens, (Chamberlin, 1947). Grayson Co., Mt. Rogers, circa 4500', 4M 2F, R. L. Hoffman, (RIHC) ; Mt. Rogers, VPI E>^)edition, IX-7-1957, W. T. Keeton, (RIHC) ; Mt. Rogers, IM, V-19-1975, D. W. Ogle, (RIHC). Greene Co., Imi. W. Lydin, IF, IV-10-1959, R. L. Hoffinan, (RIHC) ; Shenandoah N. P., Skydrive Drive milepost 71.5, 4M, VII-13-1959, R. L. Hoffman & Brownell, (RIHC). Hanover Co., Cairp Hanover, 14mi. SE. Mechanicville, 4F 8J, VII-1963, Hall, (RIHC). Henrico Co., Richmond, downtown, IM, X-18-1978, R. E. Ashton, (NCMH). Hi^aland CO., Monterey, IM, VII-3-1948, R. L. Hoffinan & Peters, (RIHC) ; Sounding Knob, 4390', 4mi. S. Monterey, IM IF, IX-17-1974, R. L. Hoffinan et fils., (RIHC) ; ravine along Ramsey's Draft jet US Rt. 250, 8mi. SE. McDowell, IM, VII-3-1976, J. E. Cooper, (NCMH). loudoun Co., 0.7mi. S. Aldie, IF, IV-19-1959, Highton, (RIHC). Louisa Co., 4.5mi. S. Louisa, Courthouse Bluff on S. Anna R., IV-9-1959, R. L. Hoffman, (RIHC) ; 4mi. NW. Louisa Courthouse, IF, VIII-12-1947, R. L. Hoffman, (RIHC). Madison Co., Shenandoah N. P., Hemlock Springs, milepost 39, Skyline Drive, 15M 12F, VII-13-1957, R. L. Hoffnan & Brownell, (RIHC). Nansemond Co., Magnolia, IM 3F, XI-9-1952, Roger Rageot, (RIHC). Norfolk Co., Ihorogood Estates., IM IF 3J, X-25-1958, H. F. Loomis, (RIHC). Page & Madison Cos., Hawkbill Mt., 3300-4000', 6M 5F, VII-3-1957, R. L. Hoffman & Brownell, (RIHC). Page Co., Shenandoah N. P., milepost 45.7, Hawkbill Gap, 9M 3F, VII-13-1957, R. L. Hoffman & Brownell, (RIHC) ; Thornton Gap on west side, IM, III-29-1947, R. L. Hoffiman, (RIHC) ; Shenandoah N. P., n r Luruz, VII- 5-1933, Gertsch, (AMNH). Princess Anne Co., North landing, 2M IF, V- 8-1949, Gertsch, (RIHC). Pulaski Co., lOmi. NE. Pulaski, 2M 7F, IV- 7-1963, D. Staples, (RIHC); Snowville, IM, VI-1963, D. Staples, (RIHC). *Rodkbridge Co., Natural Bridge, L. M. Underwood, (Bollman, 1893). Russell Co., along Laurel Bed lake, SW. Saltville, Clinch 291 Mt., IM, IV-26-1S75, D. Ogle, (RIHC); W. side of Clinch Mt., 2500', circa 2mi. S. Fugate Hill, 2M 2F, VII-31-1965, R. L. Hoffman & Brownell, (RIHC). Scott Co., Richwood, along stream, SR. 72, circa 3.5itii. NE. Dungannon, IM, VII-31-1965, Ri L. Hoffman & Brownell, (RIHC). Smyth Co., Big Walker Mt., W. Hungry Mother S. P., IM IF, IX-9-1956, R. L. Hoffiian, (RIHC). Tazewell Co., Beartown Mt., Burkes Garden, 4M IF, VI-29-1947, R. L. Hoffman & Peters, (RIHC) ; Garden Mt., 3800', county line, IM IF, X-16-1966, A. Q. field trip, (RIHC). Warren Co., Elizabeth Furnace, nr Front Royal, IF, VII-22-1966, L. Kii^t, (RIHC). % t h e CO., Gullion Fork Wildlife Management Area, E. of Blacklick, IM, VII-28-1962, R. L. Hoffman, (RIHC) ; Reed Ck., at Carter Wayside, Fort Chiswell, 2M IF, X-4-1969, R. L. Hoffman, (RIHC). WEST VIRGINIA; Barbour Co., US Rt. 119, 1.6mi. N. Belington, 5M 4F, VIII-7-1977, C. P. Withrow, (CPWC). Cabell Co., SR. 10, imi. S. jet alternate SR. 10, 7M 4F, V-20-1983, C. P. Withrow, (CPWC). Doddridge Co., SR. 18, l.7mi. S. Market, 3M 5F, C. P. Withrow, (CPWC). Grant Co., 3.2mi. E. Hopesville, off SR. 28 across bridge, 2M 7F, VIII-7-1977, C. P. Withrow, (CPWC). Greenbrier Co., Kate's Mt. IM 4F, IV-20-1968, W. A. Shear, (WASC); Kate's Mt., Greenbrier S. F . , 114, IV-20-1971, W. A. Shear, (WASC) ; North Anthony C k . , 5mi. M Œ . Neola, IM IF, X-1-1972, W. A. Shear: & W. Ash, (WASC). Hancock Co., Thomlinson Run S. P., O.Smi. inside park, 4M 2F, VIII-24-1977, C. P. Withrow, (CPWC). Harc^ Co., SR. 55, 4.6mi. E. jet SRs. 55 and 259 at Wardensville, 2M IF, IV-29-1961, R. Hilton, (RIHC). Marion Co., CR. 31/14, Valley Falls S. P., 8M 3F, VIII-7-1977, C. P. Withrow, (CEWC). Marshall CO., US Rt. 50, 1.2mi. S. Limestone, 3M 2F, VIII-24-1977, C. P. Withrow, (CEWC). Mason Co., SR. 7, Tenmile Ck., 3.7mi. from SR. 62, 5M, VI-20-1977, C. P. Withrow, (CPWC). Mercer Co., Speedway Roadside Park, SR. 20, IF, IV-26-1966, D. Gillenwater, (WASC) ; Brusii Ck. Falls, IM, V-9-1970, W. A. Shear, (WASC). Monongalia Co., Henry Clay Furance, circa 12mi. E. Morgantown, 3M IF, VI-26-1971, W. A. Shear, (WASC); 2.Imi. E. Wetzel Co. line, SR. 7, 2M 4F, V-18-1978, C. P. Withrow, (CPWC). Monroe Co., top of mt. btwn Waiteville & Gap Mills, IM, VII-14-1958, Hilton, (RIHC); 2mi. N. Ballard, SR. 12, 5M lOF, IX-14-1962, R. L. Hoffinan, (RIHC). Morgan Co., 0.7mi. into Cacapon S. P., IM, VIII-23- 1977, C. P. Withrow, (CEWC). Nicholas Co., SR. 39, circa Imi. W. Swiss, 2F, IX-2-1965, R. L. Hoffman & Brownell, (RIHC). Ohio Co., 3.5mi. S. West Liberty St. College, SR. 88, 3M 3F, V-18-1978, C. P. Withrow, (CEWC). Pendleton Co., SR. 14, off US Rt. 33 at White's Run Caitpground, 7M 2F 3J, VIII-7-1977, C. P. Withrow, (CEWC) ; Franklin, nr mouth of small cave, 2M 3F, III-25-1962, Whitehead, (RIHC). Pleasants Co., 2.4mi. S. St. Marys, SR. 2, 4M IF, VIII-24-1977, C. P. Withrow, (CEWC). Pocahontas Co., Hills Ck. Fall Scenic Area, IM, V-18-1968, W. A. Shear, (WASC); US Rt. 33, E. of Elkins, at Handley Federal Fish Hatchery, 2M 7F, VIII-6-1977, C. P. Withrow, (CEWC). E^reston C o., Terra Alta, IM IF, VI-26-1971, W. A. Shear, (WASC) ; US Rt. 50, Cathedral S. P., 3M 7F 5J, VIII-14-1977, C. P. Withrow, (CEWC). Ralei^ Co., Grandview S. P., NE. of Beckley, IM IF, IX-14- 1962, R. L. Hoffman, (RIHC). Randolph CO., 7mi. S. Whitmer, SR. 14, 20M 13F, X-1-1977, C. P. Withrov? & G. Wamsley, (CPWC) ; US Rt. 33, 292 0.9mi. W. Wymer, 6M 6F, IX-30-1977, C. P. Withrow & G. Wamsley, (CPWC) ; SR. 14, 7mi. S. Whitmer, 12M lOF 3J, X-1-1977, C. P. Withrow & G. Wamsley, (CEWC) ; l-2itii. W. Bear Heaven Picnic Area on rd to Bickles Kiob, IF, VI-12-1957, Hilton & Barry, (RIHC). Ralei^ Co., Grandview Park, NE. of Beckley, IM IF, IX-14-1962, R. L. Hoffman, (RIHC). Taylor Co., US Rt. 250, 0.9mi. S. Pruntytown, 9M 4F 9J, VIII-14-1977, C. P. Withrow, (CPWC). Tucker Co., US Rt. 119, 1.6mi. N. Belirgton, VIII-7-1977, 5M 4F, C. P. Withrow, (CEWC) ; Femow Es^jerimental Forest, 2M, X-17-1965, J. E. & M. R. Cooper, (NCMH) ; 0.2mi. N. Lanesville, trail, 13M 12F, X-1-1977, C. P. Withrow & G. Wamsley (CPWC). SR. 20, 0.9mi. S. Kanavha Head, IM, VIII-15-1977, C. P. Withrow, (CPWC). Webster Co., North Fork of Cranberry R., 5mi. SW. Three Forks, IM, VI-18-1963, R. L. Hoffman, (RIHC) ; off SR. 20, l.Omi. into Holly R. S. P., 2M IF, VIII-15-1977, C. P. Withrow, (CPWC). WISCONSIN: Clark Co., Worden Township, IF, VI-21-1946, B. Patterson, (EMNH). Dane Co., Madison, Arboretum, IF, IX-25-1954, F. A. Iwen, (FSAC). Fond du Lac Co., 6mi. N. Fond du Lac, 4J, VIII-17- 1953, H. S. Dybas, (EMNH). Pseudopolvdesmus oaludicola VIRGINIA: Princess Anne CO., Sand Bridge, 5mi. S. Virginia Beach, IM, V-8-1949, L. M. Carter, H. I. KLeirçjeter & R. L. Hoffman, (USNM), holoi^^ of Pseudopolvdesmus paludicolus; Northfolk, E. side Northfolk Reservoir, IM, V-18-1963, P. Hall, (RIHC). Pseudopolvdesmus caddo LOUISIANA: *Allen Par., (Loomis, 1959). Caddo Par., 5mi. NW. Shreveport, 2M IF, IV-13-1936, L. Hubricht, (USNM), holotype of Polvdesmus caddo. Iberville Par., 13.3mi. N. Bayou Sorrel on Maringouin Levee, 2M, X-30-1965, R. E. Tancty, (FSAC). Lablanc Par., Kinder, US Rt. 190, 7M 5F, XII-19-1958, Loomis, (USNM & RIH), holotype of Pseudopolvdesmus bidens. Natchitoches Par., Kisatchie, U , X-30-1954, W. T. Harman, (FSAC). Madison Par., IM IF, VII-10- 1966, R. E. Tandy, (FSAC). St. Landry Par., 14.2mi. SE. Lebeau, US Rt. 71, IM, XI-6-1965, R. E. Tandy, (FSAC). TEXAS: Brazoria Co., Old Ocean, 2M IF, XII-23-1962, R. O. Albert, (FSAC). Refugio C o., Tivoli, 3M 3F, Loomis, (FSAC). Pseudopolvdesmus minor ARKANSAS: Crai^ead Co., Jonesboro, 2F, III-17-1962, N. B. Causey, (FSAC). Desha CO., McGehee, 4M 8F, 1-7-1954, N. B. Causey, (FSAC). Faulkner Co., Conway, IM IF, XII-24-1956, M. A. Jackson, (ESAC); *1.5mi. W. Conway, IM IF, XII-24-1953, M. A. Jackson, (Loomis, 1959). Mississippi Co., Blytheville, 26J, 1-20-1955, Mrs. T. E. Rowlett, (FSAC). Poinsett Co., IM 3F, IV-17-1954, G. Doly, 293 (ESAC). Pulaski Co., Little Rock, (USNM), holotype of Polvdesmus m inor. INDIANA; *Benton Co., Boswell, 1 spec., D. M. Mbttier, (Bollman, 1893). Posey Co., Wabash R., West Half Moon Lake, IM, V- 14-1958, rfcCoy, Wilson & Chandler, (FSAC). Tippecanoe Co., Lafayette, IM, IV-12-1956, (FSAC). ILLINOIS: cook CO., Chicago, 6J, V-31-1943, H. S. D^ioas, (EMNH) ; imi. E. Thornton, 4J, X-29-1955, H. S. I^oas, (EMNH) ; IM IF, (FSAC) ; Illinois Collection Nat. Hist. m s . , (AC3195), 3M IF, (FSAC). KENTUCKY: Henderson Co., IM IF, IV-24-1957, R. E. Woodruff, (FSAC). IDÜISIANA: Orleans Par., New Orleans, nr. Mississippi R., btwn Broadway and State Streets, 2M IF, IV-17-1936, L. Hubricht, (USNM, A2668), holotype of Polvdesmus neoterus. MISSOURI: Callaway Co., 17M 4F 25J, X-21-1966, (FSAC); Missouri R . , flood plain, IM 6F U , 111-16-65, W. W. Dowdy, (FSAC) ; flood plain, IM, W. W. Dowdy, (FSAC). Cole Co., Jefferson R., bottom, IM, X-17-1964, W. W. Dowc^, (FSAC); flood plains, 3M IF U , VI-13-1965, (FSAC). De Kalb Co., Ami'll, 2M IF, VI-2-1962, K. Dyer, (FSAC); IM, VIII-1-1966, (FSAC). Johnson Co., Robins, IM, IV-23- 1959, J. N. Brooks, (FSAC) ; Bristle Ridge, IM, IV-26-1959, J. N. Brooks, (FSAC). St. louis Co., Buder Park, imi. SE. Valley Park, IM IF, III-15-1936, L. Hubricht, (USNM), holotype of Polvdesmus euthetus. No Data: IM, (FSAC). 294 OTHER SPECIMENS EXAMINED Cerastelactivs cavemicola (Sinclair, 1912). THAILAND; Gua Tanan, near Biserat, Patani R., IM, topqparatype, X-20-1910, (RIHC). Eutrichodesmus demanaei Silvestxi, 1910. INDOCHINA: Tonkin, Phu-Ly, IM, topoparatype, VII-2-1968, Paluo, (RIHC). Mastigonodesmus viqnai Strasser, 1974. ITALY: Sardinia; Carbonia, Riu Cannas, Grotta dei Fior, IM, topotype, XII-8-1967, A. Serra & S. Puddu (CA). Ecanerchodus fPrionomatis) minutissimus Murakami, 1976. JAPAN: Kagoshima Pref., Kawanabe-gun, at Kamiyamada of Kawanabe-cho, tuff cave Gongen-ana, IM, topotype, V-6-1973, (CPWC). E. sulcatus Miyosi, Y. 1954. JAPAN: Aichi Pref., Ja-Ana, Ishimaki- Mura, IM, X-16-1967, Y. Murakami. (CPWC). E. aculeatus Miyosi, 1954. JAPAN: Kitayama-Kyo, IM, X-17-1967, Y. Murakami, (CEWC). E. acuticluvis Murakami, 1970. JAPAN: Tokushima Etef., Kurogo, Kamo-cho, Anan-shi, I^-no-iwaya Cave, 2M2F, topotype, XII- 10-1967, Y. Murakami, (CPWC). E. masatakai Murakami, 1970. JAPAN: Tokushima ETef., Takano, Kisawa-mura, Haia-gun, Togen-daiichi-do Cave, IM, topotype, VIII-8-1969, M. Kiuchi (CPWC). E. bidens Takakuwa, 1954. JAPAN: Yamanashi Pref., Saiko-komori-ana cave, IM, IX-5-1969, S. I. Uneno & K. Kato, (NCMH). Archipolvdesmus maroccanus Attems, 1898. MOROCCO: nr Tetuan, IM, XII-1949, Fesselman, (RIHC). Polvdesmus (Acanthtarsius^ endatulus bidentatus loksa, 1958. HUNGARY: nr Szakonyfalw, IM, V-13-1984, W. C. Welboum, (CPWC). Polvdesmus (Brachvdesmus) epomeanus Verhoeff, 1952. ITALY: Vaglia, IM 2J, V-25-1984, W. C. Welboum & R. Nannelli, (CPWC). Polvdesmus fBrachvdesmus) superus Latzel, 1884. Ohio: Hamilton Co., Miami, 5M 2F, 1918, R. A. Hefner, (CPWC). Polvdesmus fSpanobrachium^ collaris Kbdh, 1847. ITALY: Vallombrosa, IM, V-27-1984, W. C. Welboum, (CEWC). Polvdesmus (Polvdesmus^ inconstans Latzel, 1884. Ohio: Franklin Co., OSU greenhouse, 2M 2F, VI-18-1984, C. P. Withrow, (CEWC). Polvdesmus (Nomarchus) renschi Schubart, 1934. ITALY: 2km E. Tosi, IM 2F, V-25-1984, W. C. Welboum, (CPWC). Speodesmus edhinourus Loomis, 1939. Texas: Kerr Co., Kerrville Etassel Ranch Cave, IM, holotype, VI-17-1938, K. Dearolf, (MCZ). Other localities for Speodesmus edhinourus as determined by Elliott (1976) 295 Texas; Bandera Co., Station "C" Cave No. 1; Haby Swallow Cave; Fossil Cave; Haby Water Cave; Fog Fissure; Ifeese Cave; Garrison Hilltop Cave; Sutherland Hollow Cave; Medina Lake Sinkhole. Canal CO., Bad Weather Pit; Fair Hole; Dierk Cave; Honey Creek Cave; Bender's (Bartel's) ; Rittiman Cave; Voges Cave. Edwards Co., V ^ t t Cave; Three Bounce Cave; Deep Cave. Hay Co., Boyett's Cave; Morton's Cave; Donaldson Cave; Boggus Cave; Wonder Cave; Ezell's Cave; McCarty Cave. Kendall Co., Kohl Ranch Cave No. 1; Behr's Cave; Pfeiffer Crawlway Cave; 474 Cave; Victor Phillips Water Cave; Schneider Ranch Cave; Century Caverns (Cave-Without-A-Name) ; Cricket Cave; Skunk X Water Cave; Cascade Caverns. Kerr Co., Pinto Ranch Cave; Stowers Cave; Mingus Root Cave; Seven Room Cave; Smith Cave, lyfeson Co., Kothmann Cave. Kfâdina Co., Daveiport Cave; Sixty Minute Cave. Sutton Co., Felton Cave; Harrison Cave. Uvalde C o., Tairpke Cave; McNair Cave; Dripstone Cave; Story Cave; Cedar Brake Cave. Val Verde CO., Twin Tree Cave. Edwards Co., Jacofcy Cave; Devil's Sinkhole; Vance Cave. Menard Co., Powell's Cave; Nell Cave. Real Co., Pape (Bradford Cave) ; Skeleton Cave; Orell Crevice Cave; Tucker Hollow Cave; Section 6 Cave; Haby Cave (Cave of the Lakes) ; Emmett Wilson Cave. Speodesmus #1, new species, as determined b y Elliott (1976) Texas: Hays Co., Halifax Bat Cave; Travis Co., Pipelane Cave; Goat Cave; Beckett's Cave; Pennie's Caver Cave X. Speodesmus bicomourus. Causey, 1959.. Texas: Bec}:'s Ranch Cave, Beck Ranch, 6mi. W Round Rock, 3M 4P, XII-10-1955, W. Alister & D. Kyser. (AMNH). Other localities for Speodesmus bicomourus as determined b y Elliott (1976) Texas: Lairpasas Co., Jackson One-Bat Cave. Travis Co., Ireland's Cave; Beckett's Cave; Cave X; Airman's Cave; Cave Y; Lost Gold Cave; Broken Straw Cave; Bee Creek Cave; Bandit Cave; Tooth Cave; McNeil Bat Cave; Dead Dog Cave No. 1; Dead Dog Cave No. 2; Cotterell Cave; Jack's Joint; Kretschmarr Cave; Lunsford Cave; Schulze Cave. Williamson Co., Great Mid Cave; Four Comers Cave; Circle Cave; Man-With-A-Spear Cave; Bone Cave; Steam Cave; Inner Space Cave (laubach Cave) ; Three Mile Cave; Williams Cave; Bat Well; Cobb Cavern. Speodesmus #2, new species, as determined by Elliott (1976). Texas: Bexar Co., Government Canyon Bat Cave; Helotes Hilltop Cave; Adam Wilson's Cave. Medina Co., Surprise Cave; Goat Cave. Speodesmus #3, new species, as determined by Elliott (1976). 296 Texas; Comal Co., Lewis Cave. Speodesmus #4, new species, as determined by Elliott (1976). Texas: Val Verde Co., Fern Cave. Scytonotus berorothi Chamberlin, 1911. Washington: Kitsap Co., Bremerton, IM, holotype, 1910, E. Bergroth, (USNM). S. columbianus Chamberlin, 1920a. CANADA: British Columbia: "Columbia Valley", IM, holotype, IX-26-1883, J. B. Tyrrell, (MCZ). S. qranulatus (Say, 1821). Ohio: Fairfield C o . , B a m e b e y Center, 2M 3F, IV-12-1985, C. P. Withrow, (CPWC). S. orthodox Chamberlin, 1925. Utah: Cache Co., Logan Canyon, Bear lake, IM, holotype, 1924, R. V. Chamberlin, (USNM). S. simplex Chaniberlin, 1941. OREGON: Douglas Co., John Day Creek, IM, holotype, XI-18-1939, J. C. Chamberlin, (USNM). S. michauxi Hoffman, 1962. Tennessee: Unicoi Co., Unicoi Mt., 2M IF, VI-16-1983, C. P. Withrow, (CPWC). S. virqinicus (Loomis, 1943). Virginia: ] % g e & Rappahannock Cos, Thornton Gap, 5M, IV-3-1949, R. L. Hoffman, (RIH). Utadesmus henriensis (Chamberlin, 1930) Utah: Henry Mountains, Mount Ellen, IM, holotype, IX-1929, R. V. Chamberlin, (USNM), U. hoffi Chamberlin and Hoffman, 1950. New Mexico: Santze Fe- Sandoval Cos. nr Albuquerque, Sandia Mountains, IM, holotype, X-22-1948, C. C. Hoff, (USNM). Bidentoqon helferorum Buckett & Gardner. 1968. California: Mendocino Co., Imi N. Mendocino, IM, holotype, VII-21-1964, J. S. Buckett, M. R. Gardner & J. R. Heifer, (USNM). B. new species. California: Humboldt Co., Areata, HSU campus, IM IF, IV-1-I978, A. K. Johnson, (NCMH); 1935 "H" St., IM, X-4-I976, A. K. Johnson, (NCMH). Coronodesmus bitnberculatus (loomis, I960) California: Alameda Co., Altamount Pass, above Niles, IM, holotype, XII-I-I926, 0. F. cook, (USNM). Ç. vosemitensis Causey, 1954. California: Tuolumne Co., Yosemitze Natl. Park, Vernal Falls, IM, holotype, II-2-1952, Gorman, (AMNH). Ç. califomicus Chamberlin, I9I8. California: Santa Clara Co., Sacramento, Stanford University, IM, I-19I4, M. G. Childs, (MCZ). Marin Co., Muir Beach, 3M IF, XII-24-I980, A. K. Johnson, (NCMH) ; same, 4M IF, XII-24-I980, A. K. Johnson, (NCMH) ; IN Stinson Beach, along SR 1, IM 4F, XII-24-1980, A. K. Johnson, (NCMH). Ç. species. California: Contra Costa Co., Bolinger Caryon Rd., nr St. Marys Col., IM, XII-24-1974, A. K. Johnson, (NCMH) ; Richmond, 1895 Tulare Ave., 2M 2F, XII-25-I977, A. K. Johnson, (NCMH). 297 Speorthus txiaanbius Chamberlin, 1952. New Mexico; Eddy Co., Carsbad Caverns, IM, holotype, IV-24-1924, 0. G. Babcock & Bailey, (USNM). Idahodesmus dentatus new species Idaho: Latah Co., 4N-8E Harvard Banks Gulch, holotype, allot^^, and 2M IF paratypes, IX-16-1977, A. K. Johnson, (NCMH), same locality, IM, IX-16-1977, A. K. Johnson, (NCMH); 4N-1.5E. Harvard, USFS cpgd., IM, IX-17-1978, A. K. Johnson, (NCMH). Revision of the genus Pseudopolvdesmus Attems, 1898 and it relationships to North American genera in the family Polydesmidae leach, 1815 The revision of the genus Pseudopolvdesmus and it relationships to other North American polydesmids first required the examination of the hi^er taxa in the family. It was determined that the sister grotç) of the Polydesmidae is the family Doratodesmidae from Southeast Asia. A cladistic analysis, based on 65 characters, was performed. Four subfamilies are proposed for the Family Polydesmidae: Mastigonodesminae Attems, 1914, Scytonotinae new status, Polydesminae new status, and Epanerchodinae new status. The Mastigonodesminae consists of 2 genera and is limited to the western rim of the Mediterranean; the Scytonotinae consists of 7 genera and is primarily western Nearctic in distribution; Polydesminae, with approximately 11 genera and Holarctic; and Epanerchodinae with one large heteromorphic genus locate in eastern Asia, primarily Japan. The North American genera of the subfamily Scytonotinae are: Speodesmus loomis, 1939, consisting of 7 troglobitic species located in westcentral Texas; Scytonotus Koch, 1847, consisting of 11 species from eastern and northwestern North America; Utadesmus Chamberlin and Hoffman, 1950, with 2 species from New Mexico and Utah; Bidentoqon Buckett & Gardner, 1968, consisting of 2 species from central Ceilifomia; Coronodesmus new genus, with 3 species from coastal California; Speorthus Chamberlin, 1952, with 1 troglobitic species from western Texas and New Mexico; Idahodesmus new genus and n ew species, I. dentatus. from northern Idaho. The native fauna of the subfamily Polydesminae is represented by the genus Pseudopolvdesmus. A cladistic analysis based on 36 characters, divided the genus into two groups; canadensis, including the species pinetorum (Bollman, 1888), and two two-species conplexes tallulanus (Chamberlin, 1943^-erasus (Loomis, 1943) and canadensis (Newport, 1844)-collinus. Hoffman, 1974, group serratus, including serratus (Say, 1921), oaludicola Hoffman, 1950, minor (Bollman, 1888), and caddo Chamberlin, 1949. Twenty-one new synonyms in the genus Pseudopolvdesmus are recognized.