§I’STEMATIC STUDIES w sam 1>sEUDOsc0R1>IONs (ARACHNIDA: Psmmosconrxorum) FRG4 TE SOUTHEASTERN UNITED STATES bv Jmes Everett pausen, B„S., M.A• ‘ Thesis submitted to the Graduate Faculty of the Virginia Fclytechnic Institute in cendidacy for the degree of D0cTOR OF PHILDSOPEY in ‘ Znelcgy gf APPROVED: E 0%man,E . rry . t , 4 . ‘^ @4/ • • r r. A r W. oa a August, 1967 Blaeksburg, Virginia ERRATA Page 47. Tap line to be included as the bottom line of page 46. Page 61. Delete "comb·like" and insert "pectinate". Page 184. "have been discussed above" should read "have been dlscussed on page l29". Page 219 and 2nd page of abstract. Delete "eco1ogy" and insert "habitat". 2TABLEOF CONTENTS Lam I. ACKNUWIEDGMBNTS............................................ 7 II. INTR¤UCTION............................................... 8 III. REVIEW O THE LITERATUR!................................... 13 IV. MATERIALS AND METHDDS...................................... 18 V. CEARACTERS OF SYSTEMAIIC IMTOEIAN¤E........................ 25 S•taa................................................... 26 Carapace................................................ 26 Chalicerae.............................................. 28 Coxal Area.............................................. 29 P•dipa1p................................................ 30 Leg•.................................................... 34 Abdumen................................................. 35 VI. SYSTEMATIC TFEATMENT....................................... 42 Order Pseudoscorpiuuida................................. 42 Suborder Heterosphyronida............................ 48 Family Tr1d•¤¤h:houi1da•.......................... 50 53 gp gg;gggg..................... ............. 54 Family Chthouildae................................ 57 Ge¤us.........4.................. 61 ........ 63 .. 70 72 3 Bass 5. ap. Il-••...................... 80 Genua.......................... 87 Subgenua............... 90 Subgenua.. 2*1 5. (15.) 555951;;.......................... 9:* 5. (5.) 99 Genua 108 5. ggludia............................... 109 Genua Chthoniua................................ 115 g. (5.) 116 Suborder Diploaphyronida................................... 128 Superfamily Garypoidea. ... .......................... 130 Superfamily Neobiaioidea................................ 131 Family Neobiaiidea................................... 133 Subfmily Neobiaiinae............................. 135 Genua Micz·obiaiu¤1.............................. 136 _tgi_.......................... 138 Genua Neobigig................................ 146 §. ca;olinenag.............................. 148 Ij. _t;_;_en__u_e_.................................... 155 _1}I_. _§1_g_1_I_;_; ap. 11.............................. 158 Subfamily Ideobiaiinae............................ 163 Genua Microcgeagria............................ 163 gg. at1ant:ica................................ 165 4 Bass Qi. 172 _l;_i_. _1g_g9_............................... 177 Suborder Monosphyronida.............................. 184 Superfamily Cheliferoidea......................... 185 Family Chemetidae.Subfam:I.1yCharuat:inae....................... 187 Genua 188 Q. _pg_rv_u_g_.................V............ 188 Genua Qllinicheruam..................... 192 _I_. c1;stingg_•.;s......................... 192 Family Cheliferidae. .. ......................... 195 Subfamily Chaliferinae...................... 196 Genua 199 Q. cogiogua........................... 200 VII. DISCUSSION................................................ 204 VIII. SlMMARY................................................... 218 DC. LITERATURE CITED.......................................... 220 ‘ X. VITA...................................................... 225 5 PLATE8 z2z.ss.a¤.2· 2.22.2 zam I. g_;i¤_9_g_q_........_.......................... 227 11. 5. gg£.¤¤sa............................................ 229 III. sa¤da1·ao¤i............................. 231 - IV. 5. BBIIÜBIBOIIÜ-••••••••••••••••••••••••••••••••••••••••• 233 V. Q@}tho¤iu• moastum................................. 235 VI. Q. m¤astus......................•..................... 237 VII. Q. m¤a¤t¤•............................................ 239 •p. VIII. gchthouiua bazharac ¤........................... 241 IX. Q. barbarae sp. :1..................................... 243 X. Q. barbaraa sp. ¤..................................... 245 XI. Klagtochtbouiua (5.) cm¤_!g21.......................... 247 XII. 5. cx·osbg}_............................................ 249 XIII. 5. mu1tig1¤o•u•...................................... 251 XIV. 5. m¤11:igi¤o•u•...................................... 253 XV. Kswuchthenius gg1udis................................. 255 XVI. 5. ga1udia............................................ 257 XVII. Chthmxius (5.) tat:acha1atus.......................... 259 XVIII. Q. t¤tracha1¤tu¤...................................... 261 XIX. Microbistxm co¤£usxm.................................. 263 XX. 5. con£u¤u¤1........................................... 265 ° XXI. 5. ccufuaum........................................... 267 MI!. Naobiaixm cato1i¤¤¤¤a................................. 269 6 E1.§£EPP.• ELELS 2,*32 H11!. 271 xxxv. gg. 273 mv. 213 gg. ev. ¤........................................ 277 XXVI!. gg. _;_h_9_}_g}_ sp. ¤................„....................„ 279 XXVIII. Microcxaagit é£1•:1£i¢•......„„.....„.„..„..„.... 281 mx. g. _;__¤g_¤_g_g..............................;.....4....... 263 xxx. gg. @1;............................................. ass xxx:. gg. ggg;............................................... 281 mx. gg. ggg.......................1.............. ......... 269 xxxux. xsexggmeuemas ............ 291 mw. :111¤1¤x¤m6• azecmcme.......................1....... 293 U2?. Dactzlochüifar 295 xxxv:. g. 297 7 I. ACKNOWLEDMENTS 8 II, INTRODUCTION The pseudoscorpions are chelicerste arthropode belonging to the Class Arachnida, Order Pseudoacorpionida, They have a body size ranging frm 0.8 ~». to 6.0 mm., and the body is divided into two tagmata, the prosoma and opisthosoms. Unlike that of the scorpion, the opisthosoa is entire and not divisible into a mesosoa and metssoma. These animals are superficially similar to diminutive scorpions, however, in the manner in which the pedipalps are carried by the animal. In addition to scorpions and peeudoacorpions, the Class Arachnida embraces auch well known forms ss spiders, ticke, mites, harvestmen, and other less well known animals: the amblypygids, uropygids, palpigrades, slpugids, and ricinuleids. In the United States, some species are widespread, while others are restricted in distribution to more limited regions auch as the southern, western, mnd southwestern United States. The habitats of the pseudoscorpions are quite diverse and include leaf meld, rotting tree trunks, the flsking bsrk of trees, rodent and bird nests, and cavee. One species, Chelifer cancroidee, is associated with huan habitations. Because of their inconspicuus habitata and the difficulty of collection, the biolegy and systematics of these animals have not been extensively studied. Pseudoscerpions are found in all areas of the world, except in the arctic and antarctic region. Savory (1964) has constructed a table, using Cha¤berlin‘s nonograph on the order an his surce, which gives a 9 summary of the world•wide distribution of the families. This table indicatea that sme famllies are localized, some cosmopolitan, but the greater number are troplcopolitan. This degree of localization of distribution does not preclude the possibility of species of some genera of tropicopolitsn families being found outside the tropics. For example, the heteresphyronid family Tridenchthoniidae and the monosphyronid family Atemnidae are considered circmtropical in their distribution but a few species of each family have been found in the United States, particularly some atemnids in subtropical Florida and soe tridenchthoniid species in the southeastern states. The implication is that ost groups have s wide geographlcal range (Hoff, 1949). The possible means of dispersal uployed by pseudoscorpions are tnadequately known. Chamberlin (1939) han discueeed several weye in which dispersal could occur. Firatly, small forms may be carried by air current: as are spider: and other small arthropods. Secondly, many forms could be transported by insects, mmmals, and birds. Thirdly, com forms may be transported by comerce, as indicated by the distribtion of Chelifer csncroides and by the amber of pseudoscorpions taken from incoming ehipments at quarantine stations in Boston and New York. Because of the esse wdth hich pseudoscorpions can be transported, Chamberlin (1939) doubts the possibility of true endemism. Hoff (1949) supports Chaberl1n's suggestions ou the dispersal of pseudoscorpions, but points out that the successful estsblishent of these man-intrnduced species is t known. 10 Pseudoscorpions have been found attached to the appendages and bodies of large flying insects. This phoretic habit of pseudoacorpions is not ectoparasitism and serves only as a dispersal mechnaism (Chamberlin, l93la). Vachon (giga, Ssvory, 1964) has not accepted the role of phoresy as a purely mechanical one for dispersal, but has proposed an alternate explanation, He was able to show that phoresy is not prevalent at all times, but occurs st frequcnt intervsls throughout the year. During one of these periods, Vachon collected several pseudoscorpions from the lege of harvestmen and all but one were females. This preponderance of feales led Vachon to conclude that phoresy has no