later. Evidence that an increased TC is not strictlytroglomorphic comes from: 1) cave species with TC= 3-5-4-5 (T. kokoweef, shoshone, brevistyla, andthe cavernicole specimen of T. bifureata); and 2)epigean species with TC = 3-5-5-5 (T. fendi and theepigean population of T. longistyla). A TC of3-5-5-5 is there<strong>for</strong>e regarded as a synapomorphy <strong>for</strong>the mulaiki subgroup (with a reversal in T. bi/obataand T. homi), with further increases in TC beingrelated to cave adaptation.20) Stylar apophysis (SA). The SA is absent inbifureata (Fig. 14) but appears to be represented inthe other Cali<strong>for</strong>nian species (except possibly deserticola)as a weakly developed laterobasal carina(Figs. 28, 41). The SA is absent in the two speciesof the brevistyla subgroup (Figs. 54, 75), but is welldeveloped in the remaining species of the mulaikigroup, where it is typically produced into atooth-like process (Figs. 78, 84, 87, 96, 107, 117,121, 136, 161, 164, 168, 176, 180, 187,201). It isnot clear whether the SA is a synapomorphy <strong>for</strong> thekokoweef group plus the mulaiki group (with a reversalin the brevistyla subgroup) or whether theSA is independently derived (autapomorphic) in thetwo clades. Given the structural differences betweenthe two types of SA, the latter possibility seemsmore plausible.21) Stylar apophysis shape. The simplest <strong>for</strong>mof the SA is found in T. kokoweef and T. shoshone(Figs. 28, 41). A more prominent SA, which bearsa tooth (reduced to a tubercle in T. hardeni, Fig.88), seems to be synapomorphic <strong>for</strong> the mulaikisubgroup. In the mulaiki infragroup (Figs. 96, 107)and T. longistyla (Fig. 78) the SA consists of a basaltooth and a lateral carina. The absence of a carinain the remaining species may be derived. Long,attenuated SA teeth are a synapomorphy <strong>for</strong> thespinoperea infragroup (Figs. 168, 176, 180, 187);here the unique SA in homi, consisting of arectangular lateral carina, appears to beautapomorphic (Fig. 201). A triangular, apicallyscrolled SA (which clips onto the PSL) is asynapomorphy <strong>for</strong> the reddelli infragroup (Figs.121, 136-147).22) Stylus shape. The stylus is compressed inthe bifureata group (Fig. 14), the kokoweef group(Figs. 26, 42, 45), and the longistyla infragroup(Figs. 78, 84). If this compressed state is plesiomorphic,then the tubular stylus in the brevistylasubgroup (Figs. 53, 54, 74, 75), the mulaikiinfragroup (Figs. 86, 87, 95, 96, 107, 108), thebi/obata infragroup (Figs. 116, 117), T. fendi (Fig.187) and T. homi (Fig. 201) could be consideredderived, as could the apically spatulate stylus in thereddelli subgroup (Figs. 121, 141, 147, 161, 164,168, 176, 180).23) Parastylar lobe (PSL) shape. The PSL ofBanksula are claw-like: curved and tapering to apoint (Fig. 5). Similar PSL are found in many speciesof Texel/a, including T. desertieola (Fig. 45),jungi (Fig. 67) and most species of the reddelli subgroup(Figs. 120, 136, 161, 163, 169, 177, 180)and suggest plesiomorphy. However, in T. bifureata(Fig. 15), the remaining two species of thekokoweef group (Figs. 27, 41), and T. brevistyla(Fig. 53) the PSL are attenuated and ribbon-like,suggesting that this is the plesiomorphic condition.The PSL in the longistyla and mulaiki infragroupsare lobe-like (a robust, rectangular lobe with folds;Figs. 77, 83, 86, 94, 108) and appear derived. Thethree additional <strong>for</strong>ms of PSL in Texella [spiral inbrevistyla (Fig. 53), hooked claw in bi/obata (Fig.116), and vermi<strong>for</strong>m infendi and homi (Figs. 187,198)] are regarded as autapomorphies.24) Mesal megaspines of palpal tibia. In Banksulaand most species of Texella the palpal tibia hasthree mesal megaspines (Fig. 10). The presence ofonly two megaspines in the longistyla and mulaikiinfragroups (Fig. 97) seems to be synapomorphic.The presence of two megaspines in desertieola andshoshone is regarded as a parallelism (autapomorphic<strong>for</strong> the two species).25) Ventral plate (VP) setal length. The lateralsetae of the VP are short in Banksula and of moderatelength in all species of Texella (Figs. 15, 42-47,74-79, 81, 86, 94, 106, 120, 136) except the brevidentaand spinoperea infragroups, where they aredistinctly longer (Figs. 160-165, 169, 177, 181,186, 200).26) Apical spine (AS) tip. The apical spine issimple or bifurcate (Figs. 15, 29, 47, 63, 76, 79,82,87, 123, 148, 149, 151) [or, rarely, trifurcate inT. mulaiki (Fig. 102) and some specimens of T.reyesi (Fig. 150)] in all species except the brevidentaand spinoperea infragroups, where it is polyfurcateto plumose and considered a synapomorphy(Figs. 160-165, 170, 172, 179, 191, 198-200).27) Genital opercular (GO) spines. The apicalmargin of the GO is unmodified in most species. Inthe spinoperea infragroup the apical margin of thefemale GO bears a pair of spines (Figs. 173, 183) [2pairs in T. diplospina (Fig. 171) and a pair of blunttubercles in T. fendi (Fig. 194)] which are consideredsynapomorphic. A few additional species havea pair of apical tubercles: males and females of T.jungi (Figs. 64, 71) and T. shoshone (Fig. 38) andsome males of T. spinoperea (Fig. 181). The presenceof these modifications in the more distantly164
elated species IS probably best interpreted as aparallelism.28) Stylar apophysis (SA) ongm. The SA ofmost species originates laterally on the stylus (Figs.77, 84, 88, 96, 107, 116, 120, 136, 164). In thespinoperea infragroup (except T. homi, Fig. 201)the SA is clearly in a more ventral position (Figs.169, 177, 180, 187), which is considered derived.29) Ventral plate setae number. The lowestnumber of < 15 setae per prong is found in Banksulaand the species of the bifureata and kokoweefgroups (Figs. 5, 6, 15, 27, 41, 42-47); the highest(> 25), apparently derived, in the brevidenta andspinoperea infragroups (Figs. 160-164, 169, 177,181, 188,200).The character trans<strong>for</strong>mations were studied withthe help of MacClade (Maddison and Maddison,1987). The preferred tree (Table 1) has a length of96 steps and a character index of 0.50.Some ambiguities exist within the mulaiki speciesgroup. The relationships within the mulaiki subgroupare uncertain as two characters, 23 (PSLshape) and 24 (palpal megaspines), cluster the mulaikiand longistyla infragroups whereas a third (22,S shape) clusters the mulaiki and bilobata infragroups.This problem may be resolved as additionaland fresh material of T. bilobata becomes availableto permit more detailed examination of that unusualspecies. The relationships of the three species in themulaiki infragroup are likewise unresolved. However,the <strong>for</strong>m of the SA varies among the threespecies (see discussion under the infragroup) and themost strongly developed SA tooth in T. mulaikimay be plesiomorphic. Finally, the spinop'erea infragroup,with the exception of the synapomorphies<strong>for</strong> T. fendi and homi (10, reduced TrIV spur; 15,absence of POP; 22, tubular S; 23, attenuated PSL)is unresolved. However, the high number (11) andbiserial arrangement (12) of the AT in T. diplospinasuggests plesiomorphy.BIOGEOGRAPHYThe species of Texella are strongly allopatric intheir distribution and closely resemble the westernNearctic phalangodid genera recently studied. Thebifureata and kokoweef groups are Cali<strong>for</strong>nian andoccur at the extreme parts of the state, both spatiallyand ecologically (Map 1). Markedly disjunct is themulaiki group (Maps 2, 5) which occurs in SE NewMexico and adjacent Texas (longistyla infragroup)and central Texas (all remaining species). With oneexception, all species are geographically isolated, asare all higher level clusters, from infragroup tospecies group. This high degree of disjunction immediatelysuggests a vicariant model of speciation.The presumed barriers, and their relative appearance,are immediately evident (Map 3). Given ourphylogenetic interpretation, the initial barrierswould have been in what is now Cali<strong>for</strong>nia: the firstbarrier separating Texella from Banksula (currentlyrestricted to caves of the central Sierran foothills;see map in Briggs and Ubick, 1981); the secondseparating T. bifureata from the kokoweef group; thethird separating the Cali<strong>for</strong>nian species from the mulaikigroup. All subsequent barriers would havebeen in Texas, the fourth isolating the brevistylasubgroup and the fifth separating the mulaiki andreddelli subgroups.Of some interest, but of uncertain significance, isthe fact that the relatively plesiomorphic elements(Banksula and T. bifureata) occur on presumed exoticterranes, whereas the relatively apomorphicelements (all remaining Texella) occur on the NorthAmerican Plate (Silberling et aI., 1984).However, some dispersal is nonetheless requiredto arrive at the present distribution, as indicated bythe single instance of sympatry: that of T. mulaikiwith diplospina and renkesae. Interestingly, the<strong>for</strong>mer species is the most troglomorphic, whereasthe others (especially T. diplospina) are the leasttroglomorphic of all cavernicolous Texella. Giventhe presumably lower dispersal potential of troglobites,the simplest hypothesis requires the dispersalof the two species of the spinoperea infragroup intothe range of T. mulaiki.TROGLOMORPHYOver two thirds of the species of Texella occur incaves and, as expected, show at least some morphologicalmodification to that environment. The severaltroglomorphic characters identified in Texella(most of which are given in Table 2) appear torepresent four basic types:1) Appendage elongation. The most apparenttroglomorphy is leg elongation. To remove the effectsof body size on leg length the ratio of leg IIlength (the longest leg) to the scute length (whichexhibits less intraspecific variation than the totalbody length) was used. The lowest LII/SL valuesare found in epigean species (2.3-3.1, except <strong>for</strong> 3.4in T. desertieola); the longest in the mulaiki infragroup(LII/SL = 9.9-15.3), T. reyesi (4.3-8.7), andT. welbourni (4.6), the species here consideredtroglobites. Intermediate leg lengths occur in allremaining (cavernicolous) species (3.2-4.3) except<strong>for</strong> T. brevistyla and diplospina which have shorter165
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PREFACEThe present volume is the se
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TABLE OF CONTENTSHOLSINGER, JOHN R.
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the state of Coahuila in northern M
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Fig. 2.-Anesia welboumi, new specie
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\\. \ - -' ..........---~\ I// --..
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Mexiweckelia hardeni, new speciesFi
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2 sets of 1 or 2 setae each; dactyl
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AFig. 7.-Holsingerius smaragdinus,
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have as many setae on the inner pla
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Antenna 1 about 33 % length of body
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Of biogeographic interest for the h
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Bowman, T .E. 1992. Two subterranea
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A note by Scott Harden that accompa
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~~..~ ~~ I))\\ "-,'.=bFig. 3.-Speoc
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unarmed, except in pleopod 2, which
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Cokendolpher, LC., and l.R. Reddell
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lobes. The larger setae vary greatl
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the relationships of the order and
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zomids their absence is considered
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some species could be either split,
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have small pores over the surface o
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inhabits tropical deciduous forest
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huitvnolotitlensis from A. stygius
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8-10, figs. 5-7; Rowland, 1973c:136
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(in row) and one pair large posteri
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(0.34); tarsus 0.64 (0.58); total 4
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Abdomen: Tergite I with two pairs a
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setae near posterior margin. stemit
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setae, and ten ventral setae. Stemi
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1,980 m elev., 26 Dec. 1986 (T. Tre
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asitarsal-tarsal proportions: 15:4:
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and one pair setae at base of proce
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Cephalothorax: Propeltidium 1.66 mm
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Male adult unknown.Immature paratyp
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IV: trochanter 1.20 (1.10); femur 3
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Cokendolpher, 1.C. 1981. The order
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Gertsch, W.J. 1992. Distribution pa
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same families and genera, but also
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species from the United States and
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the posterior median pair short and
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great many species of North America
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Key to the Eyed Females1. Eight eye
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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
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Figs. 37-48.-Ventral and retrolater
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Cicurina blanco, new speciesFigs. 7
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Type-data.-Female holotype from ins
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Description.-Female holotype: Lengt
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Description.-Female holotype: Lengt
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lengths: first femur 2 rom, fourth
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Cicurina pablo, new speciesFigs. 10
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Description.-Female holotype: Lengt
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canal in nearly vertical posItion;
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Cicurina vespera, new speciesFigs.
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- Page 125 and 126: Cicurina caverna, new speciesFigs.
- Page 127 and 128: 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
- Page 133 and 134: 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: pairs) are found in six species: T.
- 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
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Figs. 136-139.-Texella reyesi, new
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Figs. 144-147.-Texella reyesi, new
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Figs. 152-155.-Texella reyesi, new
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1989 (W. Elliott, J. Reddell, and M
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Table 3.-Continued.# locality sex S
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mesoapical; patella, 2 mesal; tibia
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Figs. 162-165.-Texella gmbbsi, new
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Figs. 166-169.-Texella diplospina,
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Figs. 174-177.-Texella renkesae, ma
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Figs. 178-18\.-Teulla spinoperca, n
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Distribution.-Known only from Fayet
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Figs. 190-193.-Texellafendi, new sp
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CLASSIFICAnONTexellabifurcata group
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Chandler, D.S. 1992. The Pselaphida
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Key to Species1. Abdominal segments
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stemite VI slightly impressed at ba
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vertexal carinae, and the laterally
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is associated with rotten woods (Ch
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small ventral carina near base, pro
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Grigarick, A.A., and R.O. Schuster.
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Cicurifla (Cicurella) holsiflgeri G