so we reduced the data matrix from 92 to 44characters.In determining the polarity of character states wehave generally followed the five "rules of thumb"used by Shear and Gruber (1983): "I) occurrence inoutgroups speaks <strong>for</strong> plesiomorphy, 2) less differentiated,more homonomously patterned meristic charactersare plesiomorphic, 3) states resembling thosein juveniles are plesiomorphic, 4) characters consistentlycorrelated with others known to be apomorphicare likely themselves to be apomorphic, and (5)correlations between morphological and chronologicalorecological positions (as in the <strong>for</strong>egoing) areto be used with caution. "PHYLOGENY OF THE PROTOSCHIZOMIDAEThe Uropygi and Schizomida (superorder Camarostomata)differ from other members of the Arachnidea(Tetrapulmonata, Megoperculata) by numeroussynapomorphies. These include: fusion of thepedipalp coxae, posterior narrowing of the intercoxalregion, reduction or loss of a postcerebralpharynx, presence of 2-3 transverse bridges with 1-2fenestra on the endosternite, great elongation of thepatellae of leg J, presence of two well-developedtrichobothria distodorsally on leg tibiae I, singletrichobothrium on tibiae II-IV each, presence of pretarsaldepressor muscles originating from the posteriorwalls of patellae II-IV, absence of postgenitaleversible appendages, presence of a pygidial exocrinegland, presence of a multi-jointed pygidial flagellum,absence of a distinct middle piece on thespermatozoan axonome, female grasping of maleabdomen during mating, and the presence of aprenymph and four nymphal instars (Weygoldt andPaulus, 1979; Shear et aI., 1987; van der Hammen,1989; Shultz, 1990; Shultz, pers. comm.). Severalother characters are known to separate Uropygi andSchizomida from other Arachnidea, but their polaritieshave not been established. These characters arefound in spermatogenesis and spermatozoa morphology(Alberti and Palacios-Vargas, 1987).In order to better understand the relationshipswithin the Protoschizomidae, we first investigatedTable I.-Characters and presumed polarities <strong>for</strong> cladisticanalyses of selected taxa of the superorder Camarostomata. 0 =plesiomorphic; 1,2,3 = apomorphic.O. Heart segments V and VI well developed (0), prosomal heartsegements absent (I).I. Prodorsum not divided into pro-, meso-, and metapeltidia(0), divided (I).2. Anterior process absent (0), present (I).3. Anterior prodorsum without seta (0), with setae (I).4. Median eyes present (0), absent (I).5. No setal pairs on prodorsum (0), with pairs of setae (I).6. Two pairs stigmata in segments VID and IX (0), one pair inVIII (I).7. Eight pairs intestinal diverticula in the abdomen (0), with sixpairs (I).8. Anterior abdominal neuromeres absent (0), eight abdominalneuromeres present (I).9. Propeltidium and tergites without distinct setal pattern (0),with (I).10. Tergites without microsetae (0), tergites I-II with pairs ofmicrosetae (I).II. Pointed process on coxa 2 absent (0), present (i).i2. Unable to jump, femur IV not enlarged (0); able to jump,femur IV and its associated muscles enlarged (i).i3. Male flagellum divided into many segments (0), undivided(I).14. Cheliceral fixed digit with two teeth (0), with three teeth (I),with more than three teeth (2).15. Cheliceral sermla with rounded knobs (0), with hyaline teeth(I).16. Cheliceral brush absent (0), present (I).17. Row of two setae on anterior process (0), one seta (I), pairof setae followed by single seta on anterior process (2).18. No seta at base of anterior process (0), pair of setae (I).19. More than two pairs dorsal propeltidial setae (0), twoanteriorly placed setal pairs present (i), two widely spacedsetal pairs present (2).20. Pedipalps sexually dimorphic (0), not dimorphic (i).2!. Male and female pedipalps of approximaly equal length ascompared to the body length (0), male pedipalps longer(I).22. Pedipalpal trochanter not produced (0), slightly produced(I).23. Basitarsal-tarsal spurs symmetrical (0), slightly asymmetrical(I), asymmetrical (2).24. Femur IV less than 4.8 times longer than deep (0), more (I).25. Trochanter IV about 1/3 length of femur IV (0), about 1/2length (1,2).26. Tergite III with two setae (0), with four setae (I).27. Male sternites with scattered or irregular rows of setae (0),with two distinct rows of setae (I).28. Sternite VI long (0), short (I).29. Eight dorsoventral muscles (0), seven (i).30. Female flagellum with segments and articles (0), withsegments only (I), without segments or annuli (2).31. Female flagellum with position 2 annulus (0), annulus absent(I).32. Flagellar setal patterns same in both sexes (0), patternsdifferent (I).33. Dm2 seta not present on female flagellum (0), present (1,2).34. Dm2 seta not present on male flagellum (0), present (I).35. Vm4 setae present on male flagellum (0), absent (I).36. Dm4 seta present on female flagellum (0), absent (I).37. Male flagellum not expanded distally (0), expanded (1,2).38. Male flagellum with distinct stalk (0), without stalk (I).39. Male flagellum over 3x long as wide (0), less than 3x (1,2).40. Male flagellum distally rounded (0), laterally compresseddistodorsally (I).41. Microsetae at base of flagellum lobes (0), microsetae onlobes (I), microsetae absent (2).42. Spermathecae with one pair lobes with pits (0), one pairlobes without pits (I), two or more pairs lobes without pits(2).43. Receptaculum margins smooth without pits (0), smooth withpits (I), lobed with pits (2), saw-toothed with pits (3).34
the relationships of the order and then those <strong>for</strong>members of the family. The character numbers inthe following discussion are the same as those appearingin Tables 1-2 and Fig. 2.Some authors consider the Schizomida to be asuborder of Uropygi, whereas others consider it anorder. We, like some previous authors, prefer torecognize the Schizomida and Uropygi as separateorders within the Camarostomata. Most authorscombining the two orders emphasize the existence ofplesiomorphies or characters of undetermined polarity.Some of the differences between Schizomida andUropygi are as great as the differences among theUropygi, Amblypygi, and Araneae. Shear et al.(1987) suggested that the pygidial exocrine glandwas an autapomorph <strong>for</strong> the Uropygi. This is not thecase, as such glands are also present in the Schizomida(Brignoli, 1973).Synapomorphies <strong>for</strong> the Uropygi are: pedipalpschelate, coxal gland orifices associated with leg IIIabsent, lateral tergocoxal muscle inserted on thepleural membrane adjacent to the coxa, presence ofommatoids on dorsal surface of anal somite, flagellumwith clear regions ventrally which are sensitiveto light, presence of five posterior abdominalneuromeres, and modification of cheliceral cleaningorgan.The Schizomida can be separated from theUropygi as well as other Arachnidea by the followingsynapomorphies: prosomal heart segments absent(character 0); prosoma divided into pro-,meso-, and metapeltidia (character 1); modificationof the anterior portion of the prosoma into a pointedprocess (character 2); presence of setae on and basalto the anterior process of the propeltidium (character3); median eyes absent (character 4); presence ofpairs of dorsal setae on the propeltidium (character5); stigmata lacking on segment IX (character 6);with six pairs of intestinal diverticula in the abdomen(character 7); with eight anterior abdominalneuromeres (character 8); male flagellum not dividedinto numerous segments (character 13); propeltidiumand tergites with distinct setal pattern(character 9); tergites I-II with pairs of microsetae(character 10); anterior distal tip of coxa II prolongedas a sharp process (character 11); ability tojump, femur IV and its associated muscles beingenlarged (character 12). Three additional charactersare known which appear to be synapomorphies <strong>for</strong>the Schizomida, but because they have only beendetermined (examined) in a single species of Hubbardiinaewe do not code them <strong>for</strong> the data matrix:absence of a posteriorly directed dorsal endosternitesuspensor muscle; absence of anterior transpatellarmuscle; and absence of the posterior patellotibialmuscle. Additional characters are known (Millot,1942) <strong>for</strong> which we have not determined the polarities.Specifically, uropygids have four pairs of prosomaticdiverticula, schizomids have one palr;Table 2.-Character states in selected members of thesuperorder Camarostomata (0 codes plesiomorphic condition;1,2,3 codes apomorphic conditions; ? codes unknown conditionor condition not relevant to region of cladogram being resolved).1, Uropygi; 2, Protoschizomus pachypalpus; 3, P. rowlandi; 4,P. occidentalis; 5, P. sprousei; 6, P. genschi (male unknown);7, P. purificacion (male unknown); 8, P. treacyae (maleunknown); 9, Agastoschizomus lucifer; 10, A. huitzmolotitlensis(female unknown); 11, A. srygius (male unknown); 12, A. patei;13, Megaschizominae; 14, Hubbardiinae.TaxaI 2 3 4 5 6 7 8 9 10 11 12 13 14Character0.0 1 I I 1 1 1 1 I I 1 1I. 0 1 I I I 1 I I I I I I2.0 I I I 1 I 1 1 I I I I3.0 I I 1 1 1 1 I 1 1 1 14.0 I I 1 1 I I 1 1 1 1 15.0 1 1 I I 1 1 1 1 I 1 16.0 1 1 1 1 1 1 1 I 1 I 17. 0 I I 1 1 I 1 I I I I I8.0 1 1 1 1 1 I 1 1 1 1 19.0 1 I 1 1 1 I I I 1 1 I10.0 1 1 I 1 I 1 1 1 I 1 111. 0 I I I I 1 1 I I I I I12.0 1 I 1 I 1 1 1 1 I 1 I13.0 1 I 1 I ? ? ? I I ? I I 114.0 0 0 0 0 0 0 0 0 0 0 0 1 2IS.? 0 0 0 0 0 0 0 0 0 0 0 1 116.0 0 0 0 0 0 0 0 0 0 0 0 1 117.? 0 0 0 0 0 0 0 1 1 1 1 2 0,318.0 1 1 1 1 1 1 1 1 1 1 1 0 019.? 1 I 1 0 0 0 0 0 0 0 0 0 0,220.0 1 I II? ? ? 1 ? ? 1 0 021.? 1 1 1 0 0 0 0 0 0 0 0 ? ?22.? 0 0 0 I 0 1 0 0 0 0 0 ? ?23.? 0 0 0 0 0 0 0 0 0 0 0 1 224.? 0 0 0 0 0 0 0 I I I I 0 025.0 I I I 1 1 I I 0 0 0 0 2 026.? I 1 1 0 0 0 0 0 0 0 0 ? ?27.? 0 0 0 0 ? ? ? II? I 0 028.? I I I 1 I 1 1 0 0 0 0 0 029.0 0 0 0 0 0 0 0 0 0 0 0 1 130.? 0 0 ? 0 0 0 0 0 ? 0 2 0 I31.? I I ? I 1 I II? 1 I 0 0,232.? I I ? I ?? 1 1 0 133.? 0 0 ? 0 0 0 0 ? 2 0 0 034.? 0 0 0 0 O? 1 I ? 1 0 035.? I I 0 0 0 0 0 0 0 0 0 ? ?36.? 0 0 ? I 0 I 0 0 ? 0 0 0 037.? 1 I 1 I ? ? ? 0 0 ? 0 0 0,238.? 1 I 1 I ? ? ? II? 1 0 039.? 0 0 0 0 ? ? ? II? 1 0 0,240.? I I 0 0 ? ? ? 0 0 ? 0 ? ?41.? 0 0 0 1 ? ? ? 0 2 ? 0 ? ?42.? 0 0 0 0 0 0 0 0 ? 0 0 I 243.? 1 1 1 0 0 0 0 0 ? 3 2 ? ?35
- Page 7 and 8: PREFACEThe present volume is the se
- Page 9: TABLE OF CONTENTSHOLSINGER, JOHN R.
- Page 12 and 13: the state of Coahuila in northern M
- Page 14 and 15: Fig. 2.-Anesia welboumi, new specie
- Page 16 and 17: \\. \ - -' ..........---~\ I// --..
- Page 18 and 19: Mexiweckelia hardeni, new speciesFi
- Page 20: 2 sets of 1 or 2 setae each; dactyl
- Page 23 and 24: AFig. 7.-Holsingerius smaragdinus,
- Page 25 and 26: have as many setae on the inner pla
- Page 28: Antenna 1 about 33 % length of body
- Page 31 and 32: Of biogeographic interest for the h
- Page 33 and 34: Bowman, T .E. 1992. Two subterranea
- Page 35 and 36: A note by Scott Harden that accompa
- Page 37 and 38: ~~..~ ~~ I))\\ "-,'.=bFig. 3.-Speoc
- Page 39 and 40: unarmed, except in pleopod 2, which
- Page 41 and 42: Cokendolpher, LC., and l.R. Reddell
- Page 43: lobes. The larger setae vary greatl
- Page 47 and 48: zomids their absence is considered
- Page 49 and 50: some species could be either split,
- Page 51 and 52: have small pores over the surface o
- Page 53 and 54: inhabits tropical deciduous forest
- Page 55 and 56: huitvnolotitlensis from A. stygius
- Page 57 and 58: 8-10, figs. 5-7; Rowland, 1973c:136
- Page 59 and 60: (in row) and one pair large posteri
- Page 61 and 62: (0.34); tarsus 0.64 (0.58); total 4
- Page 63 and 64: Abdomen: Tergite I with two pairs a
- Page 65 and 66: setae near posterior margin. stemit
- Page 67: setae, and ten ventral setae. Stemi
- Page 71 and 72: 1,980 m elev., 26 Dec. 1986 (T. Tre
- Page 73 and 74: asitarsal-tarsal proportions: 15:4:
- Page 75 and 76: and one pair setae at base of proce
- Page 77 and 78: Cephalothorax: Propeltidium 1.66 mm
- Page 79 and 80: Male adult unknown.Immature paratyp
- Page 81 and 82: IV: trochanter 1.20 (1.10); femur 3
- Page 83 and 84: Cokendolpher, 1.C. 1981. The order
- Page 85 and 86: Gertsch, W.J. 1992. Distribution pa
- Page 87 and 88: same families and genera, but also
- Page 89 and 90: species from the United States and
- Page 91 and 92: the posterior median pair short and
- Page 93 and 94: great many species of North America
- Page 95 and 96:
Key to the Eyed Females1. Eight eye
- Page 97 and 98:
39111012Figs. 1-12.-Ventral and dor
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1419 23Figs. 13-24.-Ventral and dor
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27 293334Figs. 25-36.-Ventral and d
- Page 103 and 104:
Figs. 37-48.-Ventral and retrolater
- Page 105 and 106:
Cicurina blanco, new speciesFigs. 7
- Page 107 and 108:
Type-data.-Female holotype from ins
- Page 109 and 110:
Description.-Female holotype: Lengt
- Page 111 and 112:
Description.-Female holotype: Lengt
- Page 113 and 114:
lengths: first femur 2 rom, fourth
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Cicurina pablo, new speciesFigs. 10
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Description.-Female holotype: Lengt
- Page 119 and 120:
canal in nearly vertical posItion;
- Page 121 and 122:
Cicurina vespera, new speciesFigs.
- Page 123 and 124:
procurved canal across sac; connect
- Page 125 and 126:
Cicurina caverna, new speciesFigs.
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with sac of similar size set in obl
- Page 129 and 130:
February 1964 (J. Reddell, D. McKen
- Page 131 and 132:
County: Diamond Cave, 16 August 196
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Roth, V.D. 1992. A new and first tr
- Page 135 and 136:
LITERATURE CITEDBarr, T.C. 1963. Ec
- Page 137 and 138:
Muchmore, W.B. 1992. Cavernicolous
- Page 139 and 140:
Species of Aphrastochthonius are kn
- Page 141 and 142:
FAMILY NEOBISIIDAE CHAMBERLINGenus
- Page 143 and 144:
trochanter 2.5 (2.6), femur 5.55 (5
- Page 145 and 146:
ottom of entrance pit, Ogle Cave (O
- Page 147 and 148:
tactile seta on tibia and basitarsu
- Page 149 and 150:
Missouri may be conspecific (unpubl
- Page 151 and 152:
Trichobothriotaxy of chela generall
- Page 153 and 154:
Female (figures given first for all
- Page 155 and 156:
transverse furrows; eyespots not ev
- Page 157 and 158:
Reddell and W. Russell); I female f
- Page 159 and 160:
and Acuminochernes, along with the
- Page 161 and 162:
Chamberlin, J.C. 1946. The genera a
- Page 163:
(0.36); chela (without pedicel) 2.0
- Page 167 and 168:
although not recently studied is no
- Page 169 and 170:
and stylar outgrowths, present in s
- Page 171 and 172:
in Phalangodes (et al.) is thick an
- Page 173 and 174:
pairs) are found in six species: T.
- Page 175 and 176:
elated species IS probably best int
- Page 177 and 178:
legs than expected (2.6-3.2). This
- Page 179 and 180:
~ 3.3 are synapomorphic); all trogl
- Page 181 and 182:
TAXONOMYTEXELLA Goodnight and Goodn
- Page 183 and 184:
14. BK absent (Figs. 177, 180). SA
- Page 185 and 186:
Figs. 8-11.-Texella bijUrcata (Brig
- Page 187 and 188:
Description.-Total body length, 1.5
- Page 189 and 190:
male examined closely has fewer set
- Page 191 and 192:
Figs. 26-29.-Texella kokoweej, new
- Page 193 and 194:
apical region which loses the apica
- Page 195 and 196:
Figs. 38-41.-Texella shoshone, new
- Page 197 and 198:
Distribution.-Known only from the t
- Page 199 and 200:
Figs. 52-55.-Texella brevistyla, ne
- Page 201 and 202:
Texellajungi, new speciesFigs. 60-7
- Page 203 and 204:
Figs. 66-69.-Texellajungi, new spec
- Page 205 and 206:
than S; SA with laterobasal carina
- Page 207 and 208:
~81//J/'/ ;'?/ ~~.........--~~I, II
- Page 209 and 210:
cylindrical, retina and cornea abse
- Page 211 and 212:
Figs. 93-96.-Texella cokendolpheri,
- Page 213 and 214:
SA with well developed prong and re
- Page 215 and 216:
Figs. 105-108.-Texelia mulaiki Good
- Page 217 and 218:
and McCarty Caves, which are known
- Page 219 and 220:
Color orange. Body of medium rugosi
- Page 221 and 222:
Notes.-The type locality was errone
- Page 223 and 224:
Figs. 128-131.-Taella reyesi, new s
- Page 225 and 226:
Figs. 136-139.-Texella reyesi, new
- Page 227 and 228:
Figs. 144-147.-Texella reyesi, new
- Page 229 and 230:
Figs. 152-155.-Texella reyesi, new
- Page 231 and 232:
1989 (W. Elliott, J. Reddell, and M
- Page 233 and 234:
Table 3.-Continued.# locality sex S
- Page 235 and 236:
mesoapical; patella, 2 mesal; tibia
- Page 237 and 238:
Figs. 162-165.-Texella gmbbsi, new
- Page 239 and 240:
Figs. 166-169.-Texella diplospina,
- Page 241 and 242:
Figs. 174-177.-Texella renkesae, ma
- Page 243 and 244:
Figs. 178-18\.-Teulla spinoperca, n
- Page 245 and 246:
Distribution.-Known only from Fayet
- Page 247 and 248:
Figs. 190-193.-Texellafendi, new sp
- Page 249 and 250:
CLASSIFICAnONTexellabifurcata group
- Page 251 and 252:
Chandler, D.S. 1992. The Pselaphida
- Page 253 and 254:
Key to Species1. Abdominal segments
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stemite VI slightly impressed at ba
- Page 257 and 258:
vertexal carinae, and the laterally
- Page 259 and 260:
is associated with rotten woods (Ch
- Page 261 and 262:
small ventral carina near base, pro
- Page 263:
Grigarick, A.A., and R.O. Schuster.
- Page 266 and 267:
Cicurifla (Cicurella) holsiflgeri G