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Frederick A. Coyle.
The Mygalomorph Spider Genus Atypoides (Araneae: Antrodiaetidae).
Psyche 75(2):157-194, 1968.

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THE MYGALOMORPH SPIDER GENUS ATYPOIDES
(ARANEAE : ANTRODIAETIDAE) *
BY FREDERICK A. COYLE
Biological Laboratories, Harvard University INTRODUCTION
The genus At~poidcs belongs to the mygalomorph spider family Antrodiaetidae, the remainder of which I am presently revising, and is closely related to the much larger genus Antrodiaetus. Atypoides was established by 0. P.-Cambridge ( 1883) with his description of Atypoides riversi. Since then two additional species have been discovered. The genus is solely Nearctic and has a markedly dis- junct continental distribution, with two species in northern Cali- fornia and southern Oregon and one in southern Illinois and Missouri.
Like the other species of antrodiaetids, those of Atypoides live in tubular silk-lined burrows in the gound and nocturnally capture prey which come within reach of the burrow entrance. In the present revision Atypoides riversi is redescribed, the two new species are described, and the geographic variation of each species is analyzed. Information on the ecology, life history, and behavior of these species will be included in a future paper treating the entire family.
Acknowledgements.
I am indebted to Dr. H. W. Levi of the
Museum of Comparative Zoology for encouragement and advice and to Dr. W. J. Gertsch of the American Museum of Natural History for his suggestion that I study this genus and for the large loan of A.M.N.H. material, much of which he himself has collected. Mr. J. A. Beatty, Mr. Patrick Craig, the California Academy of Sciences, and the University of Kansas have each loaned specimens. I am grateful to Prof. Varley of the University Museum of Oxford University, England, for the loan of the six syntypes of Atypoides riversi. Dr. W. H. Bossert of Harvard wrote the computer pro- grams employed in the analysis of variation. My wife, Judy, con- structed the distribution maps and helped cheerily with several other portions of the research. A National Science Foundation Graduate Fellowship and grants from the Evolutionary Biology Committee of Harvard University supported my field work during the summers of 1966 and 1967.
Public Health Service Research Grant AI-01944 to Dr. Levi helped defray some expenses. Publication is supported "Manuscript received by the editor June 12, 1968 157




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by a grant from the Evolutionary Biology Committee of the Depart- ment of Biology, Harvard University.
METHODS
Statistical methods. An analysis of variation of quantitative char- acters was performed on the three species of Awides with two aims in mind: to discover quantitative characters of value in dis- tinguishing between species of antrodiaetids, and to investigate in a preliminary manner the geographic variation within each species. There is greater need for such studies in mvgalomorph spider tax- onomy than in many araneomorph groups because of the lack of diagnostically useful complex external genitalia in the former. How- ever two major difficulties confront anyone attempting to study variation in mygalomorph species : the difficulty of collecting samples of adequate size, and the more difficult problem of achieving age homogeneity within samples of females. Antrodiaetid females (and females of other mygalomorph species) live and continue to molt fo1- one to several years after becoming sexually mature. No external structure or characteristic gross difference in seminal receptacle form has been found which indicates when a female has reached sexual maturity or what instar an adult female may be. In the present study a female specimen was included in a popu- lation sample only if it had a longer carapace than the smallest reproductively active female (with an abdomen swollen with eggs or with brood in her burrow) from the entire sample of specimens of that species. An exception was made for A. riversi where the smallest of a large number of reproductively active females collected in the coastal population was considerably larger than the single reproductively active Siesran population sample female. The low size limit for each of these two major population samples was there- fore determined by the smallest reproductively active female within it. The non-reproductively active females included in a sample rep- resent first adult instar females collected in the summer just before or after their initial mating, later adult instar females without broods, and probably an occasional immature female. It is likely that the small number of reproductively active females (one each) collected in the Sierra11 samples of A. riversi and in Atyftoides gutschi result in samples somewhat biased toward the upper end of the actual adult female body size range. Tiventjy-one measurements and 13 meristic characters were re- corded for 60 males and 159 females of the three species. (The
abbreviations and definitions for these measurements and meristic



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19681 Code - Spider Genus Atypoides I59
characters are given in the appendix at the end of this paper.) These data were analyzed with the aid of a 709.1 IBM computer at the Harvard Computer Center. The computer program, written in Fortran 11, calculated the mean and standard deviation of each measurement, meristic character, and each of 44 different ratios formed from these for each population sample of each sex and for certain groupings of samples plus individual specimens into infra- specific or species units. The program then compared these samples and groupings pairwise in all desired combinations giving for each character for each con~parison a value of the distinctness of the two samples. This value, called the "distance", is equal to the difference between the mean of each of the two samples divided by the sum of their standard deviations. This enabled me to quickly select those characters of greatest diagnostic value, those characters which show the most marked geographic variation, and those infraspecific samples that were most divergent.
As will be evident in the species diagnoses and Tables I and 11, many of the measurements, counts and ratios are diagnostically use- ful in this genus.
Several measurements and counts which are not diagnostically useful are included in the tables for their descriptive value or because they are diagnostically important in Antrodiaetus, a genus in which the species are often morphologically much more similar to one another than is the case in Atypoidr~~ The analysis
of geographic variation is considered preliminary because of the small sample sizes and the lack of more samples from important parts of each species' distributional range. Each of the localities from which a population sample of A. rivers' or A. gertschi was obtained is labeled on Map I and identi- fied in the locality records by a capital letter. These letters will be used throughout the text when referring to a particular sample or locality. The population sample sizes are indicated in the Dice-Leraas diagrams and the species sample sizes are given in Tables I and 11. Measurements. All measurements were performed by myself with the same binocular stereomicroscope and eyepiece micrometer scale. A series of three specimens was remeasured for each char- acter five times during the course of the study and indicated that the measurements were accurate to one micrometer unit for each of the four different powers of magnification used. One micrometer unit had the following value for the following characters: 0.0753 mm for CL; 0.0377 mm for CW, SL, SW, and all leg and pedipalp segment lengths; 0.0182 mm for PTT; and 0.0092 mm for CAT and all eye measurements.




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I 60 Psyche [June
Illustrations.
Drawings were made with the aid of a squared grid reticle placed in an eyepiece of a binocular stereomicroscope. Illustrations of female structures are made from reproductively active specimens or specimens much larger than the smallest repro- ductively active one within that species sample. Descriptions.
Each description is a composite of all the adult material at hand with close attention given to the type specimen in order to point out any characters for which it is peculiar. Colors are described in strong illumination under the low power of the stereomi- croscope from a series of specimens which have been dead in 80% ethanol for six months to two years. Color changes in Atypoides appear to be minimal within this time period. Records. Only material that has been personally examined is included within the records section. Numbers of immature specimens are not recorded; a lack of or $ symbols means that only im- matures were collected at that particular locality. The following abbreviations are used for the names of the more frequently cited collectors : FAC - F. A. Coyle, JWG - J. W. Gertsch, WJG - W. J. Gertsch, WI - W. Ivie, VR - V. Roth. Morphological terminology. Female genitalia: In Atypoides the sperm storage diverticulum opens into the uterus just as the latter opens exteriorly at the epigastric furrow (Figs. 78, 79). The epigas- tric furrow is between a large anterior lip and a smaller posterior lip. On the posterior face of the anterior lip is a more heavily sclerotized area which frames and extends inward through the transverse slit-like opening into the sperm storage diverticulum. This thickened cuticle also forms the floor of a rather large chamber which will be called the bursa copulatrix. This thickened floor often forms paired shallow depressions or ill-defined pockets which possibly function in the positioning of the male palpal sclerites. Four trans- parent ducts leading from the bases of the semmal receptacles open into the very weakly sclerotized roof of the bursa copulatrix. The four seminal receptacles are arranged transversely in a single row, and are thick-walled and heavily sclerotized except distally where they are capped by a rounded thin-walled transparent bulb. The proximal narrow part of the heavily sclerotized portion will be called the stalk, and the distal expanded part will be termed the bowl EXPLANATION OF PLATE 7
Figs. 1-9. Carapace and chelicerae (lateral and dorsal views). 1-3. A. riversi. 1-2. 9 from C. 3. 8 lectotype. 4-6. A. hadros. 4-5. $ paratype, from type locality. 6. $ holotype. 7-9. A. gertschi. 7-8. 9 paratype, from 0. 9. $ paratype, from 0.




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(Fig. 82). JIdc pdpu~: 111 Jfy#oid~s the cmductor is composed of two sclerites ( Fig. 57 ) , an inner cond~cior scirritc, which tapers and mntah~ the ~jnh/us distally, a d an otitcr cofidwior scIcrit~, which lies outside and cradles the inner coductor derite. ~lfacru- ~ m - : A 1a:gc thick sera, articdated in a well-dedoped socket, 1 sld call a wm-rtisrfa. This is equivalent 10 the "spine" of most qider taxonomkts. *An ~n~iforui macroseta is one wl~ich tapers absuptl!, at its tem~ind end and is therefow rigid for its entire length. +An ~fimufitr macroseta is one which tapers gradually and is therefo~e 'cry sIe11dw distally and easily bent. The hir most lateral n1ac1-metae on the ventral surface of the tibia in Figure 71 are enci- form, the two median ones attenuate. 8fiimrrrtj-: The abbrtviations AL3 P31, and PL will be used to designate the following spinnerets re-pectivel>+ : anterior lateral, posterior median, and posterior lateral. SYSTEMATICS
A tyfioidcs 0. P.-Cambridge
Aiypoidcs 0. P.-Cambridge, 1883, Pmc. 2001. Sw. Lond., pa 354. [Type species by monotypy: A. riwrrsi 0. PXarnbridge, op. fit., p. 355, PI. 36, fig. 2.1
- Simon, 1884, Bull. SOC. 2001. Fr., 9: 3IJ1 314, 316. - Simony 1890, Actes Soc. Linn. Bord., 44: 307, 309. -Simon, 1892, Histoire Naturelie des Araignees, pp, 193-195, fig, 138. -Smith, 1908, Ann, Ent, SOC. Amer., I(+) : 210, 211, 231. - Cornstock, 1912, The Spider Book, pp. 230, 249, 250; 09. cd., rev. ed., 1940, pp. 2371 23L -Gerte.ch, 1949, American Spiders, pp. 127, 130,
1311 265. - BonneG 195& Bibliographia
Araneorum, 2: 811.
DESCKIFTIUN. Cwafinct: Pars cephdica elevated above pars thoracica. Thoracic groove longitudinal. Eyes: Situated on slightly raised area (more prominent in males) which is steeply inclined an- teriorly. ALE largest, forming transverse or slightly prmurved row with AXIE. Lareral limits of eye group formed by FLE. PME
widely separated, close to their respective PLE. Stwmmz - h&um:
4 pairs of sigilla; ant. pair large, just behind labium, often indis- tinct; post. pair larger than second or third pair. Cheiicerae: Robust with strong raste1lar spines in female. Single row of large teeth (mm-oteeth) on nwsal side of dosed fang; much sm&r teeth (mi- croteeth} on ectal side of post, porrion of macrotooth row. Large apophysis projects anteriorly and dorsally from adult male chtlicera; dense brush of large setae dorsaEy near distal end of apophysis. Pedi#al#~: Coxd endite quite mall. Patella of male much shorter than tibia; tibia swollen, Ouxer conductor scle~ite of male pdpus wide, serrated on external surfacey and curved in cross section so that it partially mvelopm inner conductor dcrite. Distd portion of inner conductor scierite heavily sclerocized, narrow, and envelops



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19681 Coyle - Spider Genus A typoides 163 distal portion of embolus. Legs: Legs I and IV longer than legs' I1 or 111. Row of 9-26 trichobothria on dorsal surface of metatarsus IV of female. Abdomen : 1-3 sclerotized areas ( tergites) anteriorly on abdominal dorsum. Spinnerets: 3 pairs. AL unsegmented and reduced. PM unsegmented with functional spigots distally. PL 3-segmented with functional spigots on second and third articles. Seminal receptacles: 4 receptacles; distinct sclerotized stalks. DIAGNOSIS.
Atypoides may be distinguished from the genus An- trodiaetus by each of the following two characters: I) three pairs of spinnerets (AL may be extremely reduced and difficult to spot in some specimens of A. g-ertschi). 2) Male with cheliceral apophy- sis. The much wider outer conductor sclerite of the male palpus of Atypoides allows separation of this genus from all known North American species of Antrodiaetus.
Atypoides may be distinguished from AZiaty~us by each of the following characters : I ) AL spinnerets unsegmented. 2) Male with cheliceral apophysis. 3) Patella of male pedipalp not greatly don- gated. 4) Thoracic groove longitudinal, 5 ) Burrow entrance struc- ture consists of a collapsible collar or a stiffened turret. 6) At
least 8 or g trichobothria on the dorsal surface of metatarsus IV of the female. Also the structure of the male palpus in each of these two genera is distinct.
DISCUSSION. Atypoides is closely related to Antrodiaetus. Both of these genera are much less closely related to Aliat~~pus. One of the two dia,gnostic structures separating the former two genera - AL spinnerets - is a degenerating character, and by itself would be a poor excuse for maintaining At~poides as a separate genus. On the other hand, the cheliceral apophysis is well developed in all Atypoides males; it does not seem to be in the process of degenerating. How-
ever, it is possible that in some member (or members) of an earlier Atypoides-Antrodiuetus stock a cheliceral apophysis could have been quickly lost and its function of holding the female during mating taken over by the first legs. A consideration of the evolution of Atypoides will await the revision and similar consideration of the other antrodiaetid genera. At present it seems desirable to maintain the generic status of Atypoides, since it can be rather easily char- acterized and distinguished from Anirodiaetus, Incorrect pZuce7nent.
Atypoides culifornica Banks = Aliat~pus
californicus (Banks). Banks (1896) described this species fsom an immature male specimen collected at Black Mountain, Santa Clara Co,, California. . Smith . ( 1908) discovered the adult male of this



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species and appropriately described a new genus (Aliatypus) for its placement.
Key to the Species of Atypoidef
Males
I. Width of sternum equal to or slightly less than its length (not including labium) (Fig. 37). Tibia I with prominent macro- setae clustered near proximal end on prolateral surface and retrolaieral aspect of ventral surface; often I or 2 macrosetae near middle on ventral &-face; no macrosetae at distal end of ventral surface (Figs. 71-73). Eastern U. S. (Illinois and ........................................................................ Missouri) hadros
Sternum (not including labium) clearly longer than wide (Figs. 35> 39). Tibia, I with macrosetae present in ill-defined ventral and prolateral rows which extend most of its length; usually I to 3 macrosetae present ventrally at distal end (Figs. 68, 69, ........ 74, 76, 77). Western U. S. (Oregon and California) 2
2. Palpal tibia not cylindrical, diameter in lateral view much greater at 113 the distance from ~roximal to distal end than nea,r distal end (Figs. 10-1 3). Metatarsus I with I to 4 macro- setae on ventral susface, usually clustered at distal end (Figs. 68-70). AL spinnerets 113 or more as long as PM (Fig. 31). ....................
Abdomen purplish grey or purplish brown
riversi
Palpal tibia swollen in cylindrical form of nearly equal diameter for most of its length (Fig. 15). Metatarsus I with ill-defined ventral longitudinal row,s of macrosetae extending nearly its entire length (Figs. 75-77). AL spinneret 115 or less (may be extremely reduced and difficult to spot) as long as PM (Fig. 30). Abdomen pale grayish yellow or pale gray ............ .......................................................................................... gertschi Females
I. Sternum (not including labium) as wide as long (Fig. 38). Fewer than 13 enciform macrosetae on metatarsus I. Fewer than 7 microteeth per chelicera (Fig. 43). Wide sets,-less area on ectal side of anterior portion of cheliceral macrotooth row, Eastern U. S. (Illinois and Missouri) ........................ hadrox Sternum (not including labium) clearly longer than wide (Figs. 36, 40). More than 13 enciform macrosetae on metatarsus I. Usually more tha.n 7 microteeth per chelicera (Figs. 4.1, 42). Seta-less area on ectal side of anterior portion of cheliceral macrotooth row very narrow or absent. Western U. S. (Ore- gon and California) .............................................................. 2



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19681 Coyle - Spider Genus A typoides 165 2, AL spinnerets 113 or more as long as PM spinnerets (Fig. 31). Abdomen purplish gray or purplish brown, Seminal receptacles hea,vily sclerotized with well developed bowl (Figs. 80-86). ............................................ IVA4L/IA,fL = I -03-1 -20
riversi
AL spinnerets 116 or less (may be extremely reduced and difficult to spot) as long as PhI spinnerets (Fig. 30). Abdomen pale grayish yellow or pale gray. Seminal receptacles less heavily sclerotized with bowl weakly developed to absent (Figs. goL ...............................
94.) IVhIL/IML = 1.34-1.45 gerischi
Aiypoides r-iversi 0. P.-Cambridge
Figures 1-3, 10-13, 16, 17, 22-29, 31, 33, 34, 39, 40, 42, 44, 45, 47-49, 53-56, 68-70, 80-86. Map I.
Atypoides riversii 0. P.-Cambridge, 1883, Proc 2001. SOC. Lond., p. 355, pl. 36, fig. 2. [Three male and three female syntypes from Berkeley, California; collected by J. J. Rivers; in the University Museum, Oxford University, England; all examined. (One of these males is here des- ignated the lectotype and the other five specimens paralectotypes and are so labeled.)] -Roewer, 1942, Katalog der Araneae, 1: 189. Atypo:dcs riversi: Simon, 1884, Bull. SOC. 2001, Fr., 9: 316. -Simon, 1892, Histoire Naturelle des Araignees, 1: 195, fig. 138. - Smith, 1908, Ann. Ent. SOC. Amer., l(4) : 209, 210, 229, PI. 15, figs. 7, 8, 9, PI. 19, fig. 2, p1. 20, figs. 3, 4. -Cornstock, 1912, The Spider Book, p. 250; o$. c..t., rev. ed., 1940, p. 238. - Gertsch, 1949, American Spiders, p. 131, - Bonnet, 1958, Bibliographia Araneorum, 2: 811. COMATENTS ON ORIGINAL DESCRIPTION. In his description and illustrations of A. riversi, 0. P.-Cambridge (1883) failed to in- dicate the presence of a conductor on the male palpus. This misled subsequent authors (Smith, I 908, and Comstock, I g I 2, I 940) , who used the lack of a conductor as a diagnostic character for the genus. What may have misled Cambridge is that in the male syntypes the embolus has slid distally within the inner conductor sclerite, and the inner conductor sclerite has also slid distally and rotated within the outer conductor sclerite to extend far beyond it (Figs. 54, 55). A few of the A. riverri males which I have collected have similarly "flexed" palpi. One should be aware of the possibility of such flexion when identifying male antrodiaetids.
The lectotype and paralectotypes have changed color considerably in the 85 years since their description. The greenish hues have dis- appeared so that the colors are dull light oranges and orange browns. DESCRIPTION.
See Tables I, 11, and I11 which contain measure- ments, meristic characters, and diagnostic ratios for a sample of the species and for the lectotype.




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19681 CoyZe -- Spider Genus Atypoides 167 fifale. Carapace:
(Figs. 3, I 7). Elongate oval. Thoracic groove narrow for entire length; center of thoracic groove clearly less than 2/3 distance from ant. to post. border of carapace. Setae present only near edge of carapace and around eye group, Pars cephalica weaklv elevated; highest at ocular prot~lberance. Height of clypeus at ALE about equal to vertical diam. of ALE. Diam. of AME 314. to 2 times their distance apart. Sternum: (Fig. 39). Elongate. 3 post. pairs of sigilla circular to oval. Chelicerae: (Figs. 3, 47-49). Apophysis relatively slender, strongly curved downward near distal end, Apophyseal setae all attenuate, post. ones stronger than ant. ones. Pedipalps: (Figs. 10-13, 53-56). Greatest diam. of tibia is about 113 the distance from proximal to distal end; tapers to much smaller diam. distally. Outer conductor sclerite of palpus heavily sclerotized, relatively short and wide, serrations strong; tip very broad and nearly straight across. Inner conductor sclerite extends far beyond end of outer conductor sclerite in unflexed condition; distally curved and grad~~ally tapering to rather sharp tip. Leg I: (Figs. 68-70). Tibia usually with a prolateral row and an ill-defined double or single row of macrosetae on the retrolateral aspect of the ventral surface extending most of its length; I to 3 (very rarely 0) macrosetae ventrally at very distal end. Tibia nearly straight and cylindrical, cuxing slightly downward at distal end. Metatarsus with I to 3 macrosetae ventrally at very distal end, rarely with one on prolateral surface 113 of way from proximal to distal end. Sfeta-
tarsus long and slender, tapering slightly toward distal end, nearly straight in lateral view, slightly bowed in ventral view with prolateral surface concave. Abdou~en: Large, median, longitudinal, tri-nodal dark pattern.
Large tergite in central node. No other tergites. Spin- nerets: ( Fig. 3 I ) . AL only weakly sclerotized terminally with spigot at tip, or scIerotized terminally without spigot. AL 1/3 to 112 as long as P&I. PA1 slightly expanded to unexpanded distally; maxi- mum diam. from 114 to slightly over 1/3 length. CoZoration: Pars thoracica yellowish gray to pale greenish brown. Pars cephalica pale greenish brown posteriorly, becoming brown to reddish brown an- EXPLANATION OF PLATE 8
Figs. 10-15. Male pedipalp (retrolateral view of left pedipalp), 10-13. A. riwer~i. 10. from D. 11. Ie~totype~ 12. from I. 13. from G. 14. A, hadrox, holotype.
15. A. gertschi, paratype, from 0. [2.0 mm scale line for 10-151 Figs. 16-21. Eyes (dorsal view with lateral border of carapace hori- zontal). 16-17. A. riverxi. 16. 9 from C. 17. 8 paralectotype, from C. 18-19. A. hadros. 18. 9 paratype, from type locality. 19. 8 holotype. 20-21. A. gut~chi. 20. 9 paratype, from 0. 21. 8 holotype. r0.5 mm scale line for 16-21]




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Psyche
x A. gertschi
A. riversi
Map 1.
Distribution of Atypoide~ species.




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19681 Coyle - Sfiider Genus Atypoides 169 teriorly. Sternum slightly paler yellnwi~h gray than pars thosacica) labium and palpal coxae slightly darkei- than pass thoracica. Cheli- cerae reddish brown to dark brown. Palpal femur and patella and dorsal surface of leg I light greenish brown to reddish brown; palpal tibia much lighter. Rest of legs dorsally similar to pars thosacica) ventrally lighter. Abdomen dull purplish brown with many small scattered light spots. Tri-nodal abdominal pattern a darker olive brown ; tergite lighter yellowish gray.
Female. Carafiace: (Figs. I, 2, 16). Somewhat elongate; in- creases slightly in width from post. to near ant. end where it suddenly narrows, Shape and position of thoracic groove and distribution of setae similar to that in male, Pars ce~halica usually strongly elevated) highest point just behind eye group. Height of clypeus I to 2 times vertical diameter of ALE. Diam. of ATLIE 3/4 to I 1/2 times their distance apart. Sternum: (Fig. 40). Elongate. Sigilla shape and distribution as in male. Sternal setae concentrated around edges of sternum; many stout sharply pointed ones. Chelicerae: (Figs. I) 2, 42). Dorsal protuberance prominent, post. slope steep. Microteeth positioned beside last g to last 3 macroteeth and usually extend beyond last macrotooth. Only a very narrow seta,-less area on ectal side of ant. half of macrotooth row. Abdomen: A single tergite corres- ponding to tergite of central node on male abdomen. Spinnerets: (Figs. 31) 33) 34). AL with or without spigot at tip) AL 113 to 112 as long as PM. PM slightly expanded to unexpanded distally, maximum diam. 113 to 1/2 length. Genitalia: (Figs. 80-86). Sem- inal receptacles with relatively long, very heavily sclerotized, straight or slightly curved stalk; bowl strongly developed. Ant. portion of bursa copulatrix a relatively heavily sclerotized pocket which in dorsal view appears as a transverse bipartite darkened area at bases of seminal receptacles. Coloration: Pars thoracica yellowish gray to pale greenish brown. Pars cephalica darker; light brown to greenish brown. Sternum slightly lighter than pass thoracica; labium and pal- pa1 coxae slightly darker than pars cephalica. Palpal segments and legs dorsally similar to pars thoracica (except tarsi of palpi and of legs I considerably darker)) lighter ventrally. Abdomen dull pur- plish brown) frequently darker than male abdomen) and without dark tri-nodal pattern of male ; tergite yellowish gray. DIAGNOSIS.
Both sexes can be readily distinguished from the other species by their darker abdominal coloration. Males. The structure of the palpus (Figs. 53-56) and the shapes and macrosetae distribu- tion patterns of the tibia and metatarsus of leg I (Figs. 68-70) are distinctive for this species. IML/CL and IFL/IML (Table I)



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allow separation from A. hadros because of the relatively long meta- tarsus I of A. riversi. The cheliceral apo~hysis is relatively thinner than in A. hadros and most specimens of A. gertschi (see CAT and CAT/CL, Table I). Fen~ales. The form of the seminal receptacles and bursa copu1atl-ix aids in distinguishing this species from the others (Figs. 80-86). IVML/IML is smalles for A. riversi than for the others (Table 11) because of the relatively short metatarsus I of A. hadros and the relatively long metatarsus IV of A. gertschi. GEOGRAPHIC VARIATION. Males. Comparisons of coastal popula- tion samples (L4, I?, C, D, E and F) with one another and com- parisons of Siersan population samples (G, H and I) with one an- other usualIy yielded smaller "distances" than coastal sample vs. Sierran sample comparisons for the same character. Sierran males are generally smaller than coastal males, but these is considel-able overlap (Fig. 22). Many other measurements and meristic char- actess are correlated with body size and showed patterns of geographic variation similar to that of CL, but a few (IFL, ITarL, PFL and PTL) showed less or no overlap between Sierran and coastal samples (Figs. 23, 24). These appendage segments are relatively longer in the coastal samples (Fig. 23). Ratios formed of all other leg seg- ment lengths over CL result in very wide overlap between Sierran and coastal samples. The ~alpal tibia is relatively more swollen in Sierran males than in coastal males (Fig. 24). PedipaIp figures IO- I 3 illustrate this variation.
EXPLANATION OF PLATE 9
Figs. 22-29. Geographic variation in A. river~i. (All measurements in mm. For scatter diagrams solid black dots represent Sierran specimens, open dots coastal specimens) and X's the two Sierran foothilIs specimens.) 22-24. Males. 22. Modified Dice-Leraas diagram of CL variation. (Hori- zontal line represents the observed range) vertical line the mean, open rectangle the standard deviation, solid black rectangle the 95 percent confi- dence interval for the mean, number to right of range line the number of specimens in the sample, and letter in left column the sample-locality.) 23. Scatter diagram of IFL plotted
against CL. 24. Scatter diagram of PTT
plotted against PTL.
25-29. Females. 25. Mod. Dice-Leraas diagram of CL variation. 26. Histogram of CMT.
(One unit of vertical scale equals a
single chelicera.)
27. Mod. Dice-Leraas diagram of CMTIICTR varia- tion. Two dots above sample G symbol represent the two Sierran foothills specimens. Top two symbols represent variation in entire coastal sample and entire Sierran sample. 28. Scatter diagram of SW plotted against CL. 29. Mod. Dice-Leraas diagram of ITLICL variation.



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A. riversi
1.5 A. riversi
0
A. riversi o
8
2.5 3.0
PTL
I
0
A. riversi
0
. 0 0
0 0
A. riversi
-9
+5
- l1
- l1
-'=
-I2 29
-a
0.50 0.60
ITL
'c L
A. riversi
A
B
c =-- m rn
0 m u m
E aLm
F a h
10' I5 20 25 34
CMT
26
cwsta1
- 63
siermn - 20
A
-9
B
+5
c
- l1
D
- l1
E
-'=
F
G
A. riversi
I + ' =




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Males from D and F usually have thinner, straighter cheliceral apophyses which bend nearer the distal end (Fig. 48) than the
apophyses of males from the type locality C (Fig. 47). Some Siersan males (Fig. 49) are closer in this character to C males than are the D and F males.
The outer conductor sclerite of the palpus of Sierran males is usually relatively shorter and wider than that of coastal males. Also, the base of the embolus (expanded portion with the looping sperm reservoir) is relatively smaller in Sierran specimens. Figures 53 and 56 are palpi representing the two extremes of geographic variation in these characters.
Most males, including all from C, G and H, have only a single enciform macroseta ventrally at the distal end of metatarsus I as in Fig. 68. The rest have either one or two additional macrosetae as in Fig. 70, and three of these specimens - all in sample I - possess a single enciform macroseta on the prolateral surface one-third of the way from the proximal end. Sierran males usually have fewer tibia I macrosetae than do the coastal males. The AL spinnerets of all coastal males possess a spigot originating from a weakly sclerotized terminus (Fig. 33). None of the Sierran males have a spigot or such a terminus on their AL spinnerets (Fig. 34). The ratio of AL length to PM length is usually I :2 1/2 to I :2 for coastal males and I :3 to I :2 1/2 for Sierran males. Females.
As was true for the males, distances were greater in comparisons of samples of coastal females with Sierran samples than in intra-coastal or intra-Sierran comparisons. Also, CL averages somewhat less in
Sierran population samples than in the coastal samples (Fig. 25). Many other measurements and meristic char- acters showed a similar pattern of geographic variation; a very few of these, such as CMT (Fig. 26) have small gaps between some samples. The ratio with the most marked geographic variation (with the largest distances in the paired comparisons) is CMT/ICTR (Fig. 27). The relative width of the sternum is greater for most EXPLANATION OF PLATE 10
Figs. 30-34.
Spinnerets of females (ventral view). 30. A. gertschi, para- type from 0.
31. A. riversi, from C. 32. A. hadros, paratype from type locality. 33-34. AL spinneret of A. river& 33. from C. 34. from G. [2.0 mm scale line for 30-32; 0.3 mm scale line for 33-34] Figs. 35-40. Sternum, labium, and palpal coxa (ventral view). 35-36. A. gertsch't 35. $ holotype. 36. Q paratype, from 0. 37-38. A. hadros. 37. $ paratype, from type locality. 38. Q paratype, from type locality. 39-40. A. riversi. 39. $ lcctotype. 40. ? from (7. [2.0 mm scale line for 3 5 -401




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0.3 rnrn
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174 Psyche [June
Sierran specimens than for the coastal specimens (Fig. 28). Likewise, metatarsus IV is longer relative to metatarsus I than in most coastal specimens, but the overlap is greater.
It is worth noting that the only
two mature specimens collected low (2000 to 2500 ft.) in the western foothills of the Sierras are less different from the coastal populations than are most of the higher (6500 to 8000 ft.) Sierran specimens in some characters (see Figs. 27, 28).
The only instance of striking
intra-coastal geographic variation is the relatively long tibia I (and femur I) of sample A (Fig. 29).
The ends of the AL spinnerets are sclerotized and lack spigots (Fig. 34) on all (20) Sierran females except for the two specimens collected lower in the western foothills; these each possess a single spigot on each AL spinneret. All (66) coastal females but g have a single spigot on each AL spinneret (Fig. 33) ; 8 of these g have one "normal" AL spinneret and one like the higher Sierran females, the last (from 5) lacks spigots on both. The AL to PM spinneret length ratio and the shape of the PM spinnerets vary geographically as in the male samples.
The coastal popula,tions appear to exhibit considerable intrapopu- lation as well as geographic variation in seminal receptacle form, so that all samples are connected by overlap. Many specimens examined resembled Figure 8 I.
Some have less prominent bowls (Fig. 83 ) , or are more stout (Fig. go), or have a larger bowl diameter to stalk diameter ratio (Fig. 82). The Sierran specimens usually have rela- tively longer stalks and relatively narrower bowls (Figs. 84-86) than most coastal specimens, but there is some overlap. DISCUSSION.
The patterns of geographic variation of a number of characters indicate a considerable reduction of gene flow between the coastal population and the Sierran population. At present the Central Valley must be a formidable barrier to such gene flow, and there appears to be no contact between these populations in the more favorable montane environment to the north of the valley. It
is, of course, necessary to assume that the Sierran population was once a part of (geographically continuous with and freely interbreed- ing with) the coastal population, possibly as recently as the more humid periods of the Pleistocene.
The important problem is whether or not sufficient isolating mech- anisms have evolved since their separation to make these two pop- ulations different species. A comparison of the morphological distinctness of these two populations with the differences between any pairing of the very distinct and not too closely related species of



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19681 Coy& Spider Genus Atypoides I75
Atypoides would be misleading. The genus Antrodiaetus contains two groups of quite closely related species, some sympatric and some allopatric - once Antrodiaetus is revised a comparison with the differences between its species will be more informative. Samples
from lower elevations on the western slope of the Sierra Nevada Mtns. should be studied to see more clearly what variation pattern accompanies such a wide elevational (and climatic) range. A more thorough search should be made in the foothills around the northern and eastern edge of the Central Valley for additional populations. Until these approaches shed light on this problem, I feel unjustified in describing the Sierran population as a good species. DISTRIBUTION.
A coastal population in northern California west of the Central Valley as far south as the Monterey Peninsula. A Sierran population, apparently geographically isolated from the coastal population, in the Sierra Nevada Mtns. and western foothills (Map I). There is no reason to doubt Rivers' ( 1891 ) and Smith's ( I 908) records of A. riversi from the Monterey Peninsula,. Simon 1890) lists a record of A. riversi from the San Bernardino Mtns. and cites George Marx for the locality data. Smith's ( 1908) mention of a report of A. riversi from the San Bernardino Mtns. probably refers to Simon's paper. The validity of Marx's record is rather doubtful.
RECORDS. Coastal population : CALIFORNIA. Alameda Co. : Berke- ley [sample-locality C], 3
d1, 3 (? (J. J. Rivers) ; Oct. 1919, 8
(Dietrich) ; 9 9 (several dates and collectors). - Castro Valley, 6 Apr. 1941, 4 $ (W. M. Pearce). -Oakland, 26 Feb. 1954 (R. 0. Schuster, B. Adelson). -'Humboldt Co.: 18 mi. W. of Willow Creek, 21 Aug. 1959, 2 9. - Miranda [sample-locality 4, 3 June 1936, 9 Q (C. R. Csosby). - Marin Co.: San Geronimo, 27 Sept. 1963, ? (WI). -Mendocino Co.: Christine, 30 May 1936, Q. - Piercy, 23 July 1953 (WJG, JWG). - Santa Clara Co.: Black Mtn. (R. W. Doane).
- Stanford Univ. campus, Feb. 1921 (J.
C. Chamberlin). -Stevens Creek, 20 Apr. 1941, 2 9 (W. M. Pearce) . - Santa Cruz Co. : Ben Lomond [sample-locality El, 25 June 1952, 3 Q (WJG); 20 July 1953, 3 ? (WJG) ; 3 Apr. 1960, 2 Q (WJG, WI, Schrammel) ; 28 March 1921, ? (0. E. Sette). - Felton, I 6 Aug. 1959, $ (VR, WJG) . - 7 mi. N.W. of Santa Cruz on Empire Grade Rd., 23 June 1965, 9 (H. B. Leech). - 2 to 5 mi. W. of Feltan on Ice Cream Grade Rd. [sample-locality F], 1000-1700 ft., 14 Aug. 1967, 2 8, 11 $ (FAC). -1.6 mi. and 4.0 mi. N. of Big Basin St. Pk. headqtrs. on Big Basin Highway sample-locality Dl, 1500 and 1750 ft., 15 Aug. 1967, 5 d, 12 9



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(FAC) . - Solano Co. : Green Valley Creek [sample-locality B] , 19 June 1953, 5 ? (E. I. Schlinger). -Sonoma Co.: Guerneville, 22 July 1953, 3 ? WJG, JWG).
Sierran population : CALIFORNIA. Alpine Co. : Ebbets Pass [sample-locality HI, 8730 ft., 19 Sept. 1963, 8 (WJG). - 5 mi. W. of Ebbets Pass on Rt. 4 [sample-locality HI, 7500 ft., 7-8 Aug. 1967, d , 9 (FAC). - Amador Co.: Pine Grove, 2500 ft., 7 July 1958, 9 (WJG, VR). -Calavaras Co.: Avery, 8 July 1958 (WJG, VR). -El Dorado Co.: Meyers, 6500 ft., 5 Sept. 1958, $ (E. I. Schlinger). - 3 mi. N. of Placerville, 2000 ft., 8 Sept. 1959, ? (WJG, VR). - Bayview Campground (U.S.F.S.) at Emerald Bay of L. Tahoe [sample-locality I], 6500 ft., 6-7 Aug. 1967, 9 d, 8 ? (FAC). - Mariposa Co.: Yosemite Natl. Pk., about 20 mi. E. of Crane Flat on Rt. 120 [sample-locality GI, 8000 ft., 8 Aug. 1967, 3 d, 8 ? (FAC).
Atypoides gertschi new species
Figures 7-9, 15, 20, 21, 30, 35, 36, 41, 50-52, 58-60, 61-67, 74-77, 90-94. Map 1.
TYPE SPECIMENS. Holotype male from Manzanita Lake, 5890 ft. (1790 m), Lassen Volcanic National Park, Shasta Co., California, 3 August 1967 (F. A. Coyle) in the Museum of Comparative Zoology. Four males and 12 females from type locality with same data (except 2 of the females collected 5 July 1952 by W. J. Gertsch) designated as paratypes and deposited in the M.C.Z. and A.M.N.H. This species is named in honor of Dr. W. J. Gertsch for his valuable contributions to the systematics of the atypoid mygalomorphs and in appreciation of his help. DESCRIPTION.
See Tables I, I1 and I11 which contain measure- ments, meristic characters, and diagnostic ratios for a sample of the species and for the holotype.
EXPLANATION OF PLATE 11
Figs 41-43.
Cheliceral teeth of females (ventral view of left chelicera). 41. A. gertschi, paratype, from 0.
42. A. riversi, from D.
43. A. hadros,
paratype, from type locality. [1.0 mm scale line for 41-43.] Figs. 44-45. Left palpal tarsus of A. riversi (retrolateral aspect of dorsal surface). 44. Q, from D. 45. penultimate 8, from F. [2.0 mm scale line for 44-45.]
Figs. 46-52.
Chelicerae of males (ectal view of dissected left chelicera without fang). 46. A. hadros, holotype. 47-49. A. riversi. 47. lectotype. 48. from F. 49. from I. 50-52. A. gertschi. 50. from P. 51. paratype, from 0. 52. from M. [2.0 mm scale line for 46-52]



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2.0 rnm




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Male. Carapace: (Figs. 9, 21 ) . Elongate oval. Thoracic groove narrow for entire length; center of thoracic groove less than 2/3 distance from ant. to post. border of carapace. Setae scattered over entire pars thoracica, densest at lateral edges. Setae rare to absent at base of pars cephalica, becoming numerous higher up. Pars cepha- lica weakly elevated, highest at eye group. Height of clypeus at ALE I to I 1/2 times vertical diam. of ALE. Diam. of AME 2/3 to slightly greater than their distance apart. Sternum: (Fig. 35). Elongate. 3 post. pairs of sigilla oval. Chelicerae: (Figs. 9, 50-52). Apophysis moderately to very thick; curving downward distally. All apophyseal setae attenuate, post. ones stronger than ant. ones. Pedi- palps: (Figs. 15, 58-60). Tibia swollen in cylindrical form, of nearly equal diam. fo1- most of its length. Outer conductor sclerite of pal- pus long, translucent, serrations small ; tip rounded to ~ointed. Inner conductor sclerite extends slightly to well beyond edge of outer con- ductor sclerite; narrowed and slightly curved distally, rather blunt at tip.
Leg I: (Figs. 74-77). Tibia with 3 ill-defined longitudinal rows of macrosetae (I prolateral and 2 ventral) running most of its length. Tibia nearly straight and cylindrical, curving downward slightly at distal end. Metatarsus with 2 ill-defined longitudinal lateroventral rows of macrosetae extending most of its length. Meta- tarsus nearly straight, decreasing gradually in diam. toward distal end. Abdomen: 2 large tergites, second slightly larger than first; often a bilateral pair of tiny sclerotized spots post. to second tergite. Spinnerets: (Fig. 30). AL greatly reduced ; lacking spigots ; not more than 1/5 as long as PM, often much less. PM elongate and cylindrical ; diam. 1/4 to 1/3 length. Coloration : Pars thoracica pale yellow to grayish orange. Pars cephalica light brown to brown. Sternum slightly lighter than pars thoracica; labium and palpal coxae slightly lighter than pars cephalica. Chelicerae light brown to dark reddish brown. Palpal femur and patella grayish orange to dark orange brown, tibia and tarsus slightly lighter. Leg I dorsally nearly as dark as palpal femur; rest of legs lighter dorsally, slightly darker than pars thoracica. Abdomen light grayish yellow to light gray; tergites darker yellowish gray. Holotype nearer to darker end of color range.
EXPLANATION OF PLATE 12
Figs. 53-60. Palpus (prolateral view of left palpus). 53-56. A. rive&. 53. from D. 54-55. Lectotype, in "flexed" condition. 54. ventral view. 55. prolateral view. 56. from G. 57. A. hadros, holotype (0.c.s. -outer con- ductor sclerite, ks. - inner conductor sclerite, emb. - embolus). 58-60. A. qcrtschi. 58. holotype. 59. from N. 60. from M.



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180 Psyche [June
Female. Carapace: (Figs. 7, 8, 20). Lateral border rounded in dorsal view, widest at or behind center; somewhat constricted at ant. portion of pars cephalica. Shape and ~osition of thoracic groove and distribution of setae similar to that in male. Pars ccphalica moderate- ly elevated, highest either at eye group or slightly behind. Height of clypeus at ALE I to 2 times vertical diarn. of ALE. Dim. of AME I
to 2 times their distance apart. Stcram: (Fig, 36). Elongate. Shape and position of sigilla as in male. Sternal setae all slender. Chelicerae: (Figs. 7, 8, 41 ). Dorsal protuberance broadly rounded in side view. Microteeth positioned beside last 7 to last 3 macro- teeth; often extend beyond last macrotooth. No seta-less area on ectal side of ant. half of macrotooth row. Abdomen: Single tergite corresponding to second tergite on male. Group of prominent setae ant. to tergite, sometimes with small patches of sclerotized cuticle at their bases, corresponding with first tergite on male. Spinnerets: (Fig. 30). AL greatly reduced; lacking spigots; not more than 1/7 as long a PM, often much less, PM as in males. Gen'ttalia: (Figs. 90-94). Seminal receptacles relatively weakly sclerotized; stalks strongly sinuous to stouter and non-sinuous; bowl very weakly developed, its border with bulb ill-defined. Coloration: Pars cephalica pale yellow to light grayish orange; pars thoracica the same or slightly paler. Sternum same as pars cephalica; labium arid palpal come slightly darker. Chelicerae pale grayish orange to orange brown. Pedipalps and legs dorsally similar to pars cephaiica, ventrally slightly lighter, Abdomen similar to male; tergite usually lighter than male tergites. DIAGNOSIS.
Both sexes of A. gertschi can be separated from the other species by the greatly reduced AL spinnerets which are 1/5 or less as long as the PSI spinnerets and by the setae scattered over the pars thoracica. Also, the relatively small size of the eye group and the AME allow separation from the other species by OQW/CL and CL/AMD (Tables I and 11). Males. The structure of the palpus (Figs. 58-60) and the shapes and macrosetae distribution pat- terns of the tibia and metatarsus of leg I (Figs. 74-77) are distinctive EXPLANATION OF PLATE 13
Figs. 61-67. Geographic variation in A, 9ert~chi. (All measurements in mm. For scatter diagrams solid black dots represent P specimens, X's represent M specimens.) 61-63. Males. 61. Mod. Dice-Leraas diagram of CL variation.
62, Scatter diagram of ITarL plotted against CL. 63. Scatter diagram of IML plotted against CL. 64-67. Females. 64. Histogram of ICTR.
(One unit of vertical scale represents a single specimen.) 65. Histo- gram of CT.
(One unit of veriicat scale represents a single chelicera.) 66. Mod. Dice-Leraas diagram of CT/ICTR variation. 67. Scatter diagram of IVTL plotted against CL.




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M - 2
A. gertschi
N --t--4
0 ______I______5
P

A. gertschi
A. gertschi
M d
A. gertschi o 0 o
0 0
0 0 0
P
6 8 10 12
ICTR 64
A. gertschi
M




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I 82 Psyche [June
for this species. The shape of the palpal tibia. (Fig. 15) is an excellent distinguishing character and is quantified by the ratios PTT/CL and PTT/PTL (Table I). Because of the relatvely long metatarsus IV and short tarsus IV of A. gertschi, separation from the other two species is aided by IVML/IML and IVML/IVTarL (Table I) and from A. riversi by IVTarL/CL (Table I). Because of the large body size of A. gertschi, IVML, PFL, and even CL will help in identification.
GEOGRAPHIC VARIATION. Males. M is the most divergent popula- tion sample and P the next most divergent in most of the characters studied. CL (Fig. 61 ) and almost all other measurements and meristic characters correlated with body size were much smaller in the two M specimens than in the other males. The greatest distance between sample M and the other samples occurs with the ratio ITarL/CL (Fig. 62) and other ratios expressing the relatively long tarsus I in these two specimens. Sample P has strikingly smaller values of IML/CL than the other males of A. gertschi (Fig. 63). Most of the other ratios yield smaller distances between populations than the measurements and meristic characters. CAT varies considerably within samples N, 0 and P, but averages thicker in P, and thinner in M, than in N and 0, so that there is a north to south cline of increasing CAT diameter. Figure 52 is a cheliceral apophysis from M near the lower end of the range of variation, Figure 51 is an apophysis from 0 near the middle of the CAT range, and Figure 50 is an apophysis from P near the maximum CAT. The geographic variation pattern of the ratio CAT/CL is less clinal and shows much overlap.
The tip of the outer conductor sclerite is rounded in the holotype (Fig. 58)
and in all other males except two of sample N (from 6700 ft. near Sand Flat Rd.) with considerably more pointed tips (Fig. 59) and both males from M with strongly pointed tips (Fig. 60). The distance which the inner conductor sclerite extends beyond the end of the outer conductor sclerite varies considerably within most samples, but is longest in the M males. This latter variation may not be genetic, but rather due to palpal flexion prior to death. Intrapopulation variation in the number of enciform macrosetae on the tibia (range for all specimens i,s I 3-1 8) and metatarsus (range for all specimens is 9-24) of leg I is large, and all population samples are connected by wide overlap.
Figure 75 shows a metatarsus I
with relatively few enciform macrosetae and Figure 74 shows a tibia with a larger number than the holotype (Fig. 76).



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19681 Coyle - Spider Genus Atypoides 183 Most of the sample P specimens, both from M, and a few other individuals have AL spinnerets which are reduced to nearly imper- ceptable swellings marked by two or three crooked setae. Other specimens have AL spinnerets intermediate in size 'between these and those in samples N and 0 which are usually 1/6 to 1/5 as long as the PM spinnerets.
Females.
As in the male sample comparisons, M is the most divergent and P the next most divergent sample of females. Wide overlap exists among all four population samples in almost every character. For measurements and meristic characters, the greatest interpopulation "distances" show up in ICTR (Fig. 64), IVCTP, IVCTR, and CT (Fig. 65) in each of which sample M is divergent. The only ratios with strong geographic variation are CT/ICTR (Fig. 66), which produces a gap between sample M and the others, and IVTL/CL (Fig. 67) and IVML/CL both of which separate sample P somewhat from the other samples. The setation of the carapace and chelicerae of females (and males) from M and from Ney Springs, Siskiyou Co., Calif., is sparcer than on most specimens from other localities. The dorsal protuberance of the chelicerae in side view is slightly higher and less broadly rounded in most females of sample M. The AL spinnerets of several specimens from P and one from M are greatly reduced as described above for the males. Sample 0 has the highest mean AL spinneret length, N and M the next highest.
The seminal receptacles exhibit wide va,riation in shape. Samples N and 0 and a small sample from Crater Lake Natl. Park have widely overlapping ranges, with stalks usually quite sinuous (Figs. 90-92) . Population sample P differs markedly from these samples, with nearly non-sinuous to slightly sinuous and short stalks (Fig. 94). M also differs, usually with nearly non-sinuous and stout stalks ( Fig. 93
DISCUSSION.
These patterns of geographic variation may be the result of reduced interpopulation gene flow at the periphery of the species range and consequent divergence of certain peripheral popula- tions (M and P). Population M appears to be near the northwest- ern periphery of the species range and is much lower in elevation and in a somewhat different habitat than the other three population samples. Population P is near the southern edge of the species range. Larger sample sizes (particularly of males) and geographically in- termediate samples are needed to clarify the status of these two pop- ulations. The possibility that isolating mechanisms have evolved



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between M and the other populations studied should be emphasized, but on the basis of the present incomplete data I feel unjustified in assigning a subspecific or specific status to the population represented by sample M.
DISTRIBUTION. Cascade Mtn. Range and foothills from southern Oregon south and east into the northern end of the Sierra Nevada Mtns. of California (Map I).
RECORDS. CALIFORNIA.
Shasta Co.: 8 mi. S. Dunsmuir, I I July
1954 (R. 0. Schuster, E. E. Gilbert). -Hatchet Creek, 10 mi. W. of Burney, I Aug. 193 (WJG, JWG) . - Lassen Vol. Natl. Pk., Manzanita Lake [sample-locality 01, 5890 ft., 3-4 Aug. 1967, 5 d , 10 Q (FAC) ; 5 July 1952, 2 $ (WJG) . - Sierra Co. : Yuba Pass [sample-locality PI, 6700 ft., 5 Aug. 1967, 5 8, 14 $ (FAC). - 3 mi. W. of Yuba Pass, 6200 ft., 5 Aug. 1967 (FAC). -Sis- kiyou Co.: Bartle, 21 July 1941, d (W. M. Peai-ce). - Ney Springs, 5 mi. W. of Mt. Shasta City, 12 Sept. 1959, 2 $ (WJG, VR). - 5 mi. E. of McCloud on Rt. 89 along McCloud Cr., I Aug. 1967 (FAC). - 5 to 14 mi. E. of Mt. Shasta City on Rt. AIO [sample-locality v, 1-2 Aug. 1967 ( McBride Springs, 4950 ft., 9; 5700 ft., 9; Sand Flat Rd., 6700 ft., 2 rf, 4 ?; Panther Meadows Campgrd., 7440 ft., 2 d\ I I 9) , (FAC) . - Tehmna Co.: Deer Creek, 6 July 1952, 2 c? (WJG). OREGON. Jackson Co.: Ashland, Lithia Park, 2000 ft., [sample-locality MI, 31 Aug. 1959, 2 d, 9 Q (WJG, VR); 31 July 1967, Q (FAC). -KZanzath Co.: Crater L. Natl. Pk., Annie Springs Campground, 6000 ft., 24 Aug. 1956, c? (R. W. Fredrickson) ; 30 July 1967, Q (FAC). Atypoides hadros new species
Figures 4-6, 14, 18, 19, 32, 37, 38, 43, 46, 57, 71-73, 78, 79, 87-89. Map I.
TYPE SPECIMENS.
Holotype male from Ferne Clyffe State Park, Johnson Co., Illinois, 13 September 1966 (F. A. Coyle) in the Museum of Comparative Zoology.
One male and 4 females from
type locality with same date designated as paratypes and deposited EXPLANATION OF PLATE 14
Figs. 68-77.
Male leg I tibia and metatarsus (left leg). 68-70. A. riversi, from D. 68. ventral view. 69. prolateral view. 70. ventral view of distal end of metatarsus; another specimen.
71-72. A. hadros, holotype. 71. ven-
tral view.
72. prolateral view. 73. A. hadros, from Pine Hills, Union Co., 111.; ventral view of tibia. 74-77. A. gertschi. 74. paratype, from 0, ventral view of tibia.
75. from M, ventral view of metatarsus. 76. holotype, ventral view. 77. holotype, prolateral view.



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186 Psyche [June
in the M.C.Z. and the A.M.N.H. Hadros is a Greek adjective meaning "stout" or "thick".
DESCRIPTION.
See Tables I, 11, and I11 which contain measure- ments, meristic characters and diagnostic ratios for a sample of the species and for the holotype.
Male. Carapace: (Figs. 6, 19). Quite broad. Post. part of thoracic groove expanded laterally; center of thoracic groove about 2/3 distance from ant. to post. border of carapace. Setae present only near edge of carapace and around eye group. Pars cephalica moderately elevated; highest anteriorly at eye group. Height of clypeus at ALE much less than vertical diam. of ALE. Diam. of AME slightly greater than their distance apart. Sternum: (Fig. 37). Nearly as wide as long. 3 post. pairs of sigilla oval, 2 posterior-most pairs usually drawn out and tapered toward edge of sternum, post. pair most drawn out. CheZicerae: (Figs. 9, 46). Apophysis thick and short, weakly curved downward distally. Apophyseal setae strong, post. ones non-attenuate and stiff. Pedipalps: ( Figs. 14, 57). Tibia with greatest diam. in lateral view about 1/3 the distance from proximal to distal end, tapering to much smaller diam. distally. Outer conductor sclerite of palpus heavily sclerotized, serrations strong, end concave with one side extending beyond other. Inner conductor sclerite extends well beyond end of outer conductor sclerite in unflexed condition; distally tapering, only slightly curved and relatively thick. Leg I: (Figs. 71-73). Tibia with group of mostly enciform macrosetae at proximal end on prolateral surface, a smaller group (sometimes only I ) of macrosetae proximally on retrolateral aspect of ventral surface and I (rarely o or 2) macroseta near mid- dle ventrally. Tibia nearly cylindrical, curving downward slightly at distal end ; swollen slightly at proximal groups of macrosetae. EXPLANATION OF PLATE 15
Figs. 78-79. Female genital region, A. hadros, paratype, from type locality. 78. Semidiagrammatic long. section through a median seminal receptacle, ant. and post. lips slightly pulled apart. 79. Posterior ventral
view of genital region. (a.1. - anterior lip, p.1. -posterior lip, b.cop. - bursa copulatrix, o.b.cop. - opening of the b.cop., ut. - uterus, sem.r. - seminal receptacle) [0.5 mm scale line for 791 Figs. 80-94. Seminal receptacles (dorsal view). 80-86. A. rivers;. 80. from D. 81. from C. 82. from San Geronimo, Marin Co., Calif. 83. from Guerneville, Sonoma Co., 'Calif. 84. from Placerville, El Dorado Co., Calif. 85. from I. 86. from G. 87-89. A. hadros. 87-88. paratypes, from type locality. 89. from Little Grand Canyon, Jackson Co., 111. 90-94. A. gertschi. 90. paratype, from 0. 91. from Crater L. Natl. Pk., Klamath Co., Ore. 92. from N.
93. from M. 94. from P. [0.5 mm scale line for 80-941



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o b cop. P 1.
0.5 rnm




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188 Psyche [June
Aletatarsus usually with 4 macrosetae - 3 ventral, and I prolateral at distal end - positioned as in Figure 71. Metatarsus not greatly elongate, bowed so that prolateral surface is concave; proximal half swollen in lateral view. Abdomen: 2 large tergites (second larger than first) and usually a bilateral pair of small sclerotized patches post. to second tergite. Spinnerets: (Fig. 32). AL without spigots, almost 1/2 as long as PM. PM slightly expanded distally, maximum diam. 1/3 to 1/2 its length. Coloration: Pars thoracica pale yellow to tan.
Pars cephalica light brown to brown. Sternum similar to pars thoracica; labium and palpal coxa similar to pars cephalica. Chelicerae reddish brown to very dark reddish brown. Pedipalps pale grayish orange to orange brown dorsally. Leg I orange to bright orange brown, darkest dorsally. Rest of legs duller, dorsally similar to pars thoracica, ventrally lighter. Abdomen very light grayish yellow, tergites darker yellowish gray. Holotype at lighter end of range.
Female. Cara/*ace: (Figs. 4, 5, 18). Quite broad. Shape and position of thoracic groove and distribution of setae similar to that in male. Pars cephalica very strongly elevated, highest just behind eve group. Height of clypeus at ALE 1/2 to I times the vertical diam. of ALE. Diam. of AME slightly less or equal to their distance apart. Sternum: (Fig. 38). As wide as long. Shape and position of sigilla similar to that in male. Numerous stout, sharp sternal setae as well as long slender ones. Chelicerae: (Figs. 4, 5, 43). Very robust. Dorsal protuberance broadly rounded in side view. Micro- teeth positioned beside last 2 or last macrotooth and extend post. to last macrotooth. Wide sets,-less area on ectal side of ant. half of macrotooth row. Abdomen: Single large tergite corresponding to second tergite on male. Ant. to this is bilateral pair of prominent setae sometimes with tiny sclerotized patches at bases corresponding to first tergite of male. Spinnerets: (Fig. 32). Slightly thicker than in males, but length ratios similar. AL without spigots. PM
slightly expanded distally, maximum diam. about 1/2 length. Geni- talia: (Figs. 78, 79, 87-89). Seminal receptacles with short, rather stout, straight to slightly curved stalk, wide prominent bowl. Colorn- tion: Pars cephalica light brown; pars thoracica slightly paler. Ster- num similar to pars cephalica; labium and palpal coxae darker brown. Chelicerae brown. Pedipalps and legs dorsally similar to pars thoracica (except tarsi of pedipalps and of leg I darker), lighter ventrally. Abdomen and tergite as in male.
DIAGNOSIS.
Both sexes of A. hadros can be easily distinguished from the other species by the very wide sternum (Figs. 37, 38),



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19681 Coyle - Spider Genus Atypoides 189 which difference is expressed by SL/SW for the females in Table 11. The slight expansion of the posterior portion of the thoracic groove and the more posterior placement of the thoracic groove aid in iden- tification. Males. The alp us structure (Fig. 57) and the shapes and macrosetae distribution patterns of the tibia and the metatarsus of leg I (Figs. 71-73) are distinctive for this species. The small value of IML for A. hadros separates it from the other species (Table I). The shortness of both tibia and metatarsus of leg IV allows separation from the other species by IVTL/CL and IVML/ CL (Table I). The stiff non-attenuate setae forming the posterior half of the cheliceral apophysis and the form of the apophysis (Fig. 46) helps to identify this species. Females. The wide seta-less area on the ectal side of the anterior half of the cheliceral macrotooth row is not found in the other species.
The small number of IMS and
CAIT and the related ratio IVMCR/IMS (Table 11) allow easy separation from the other species. FCT number (Table 11) aids in rJi tingu-'shinq A. hydros from A. gertschi. Metatarsus I is relatively short (see IML/CL and IFL/IML, Table 11) and also aids in
identification.
VARIATION. Males. The two specimens from Ferne Clyffe St. Pk. are much smaller than the average size of the 15 males from Pine Hills. The characters which show the most geographic variation are CAT/CL and PTL/CL. The cheliceral apophysis is thicker relative to body size in all Pine Hills males, and the palpal tibia is relatively longer in the Ferne Clyffe St. Pk. males. Considerable intrapopulation variation exists in macrosetae number and placement on leg I; the group on the retrolateral aspect of the ventral surface at the proximal end of the tibia contains from 2 to 7 macrosetae (I to 4 enciform macrosetae) and the prolateral group contains from 4 to I 5 macrosetae (4 to 14 enciform macrosetae) . Figure 73 illus-
trates a tibia with high numbers in each group of macrosetae. The left metatarsus I of 12 of the 15 males has 4 macrosetae positioned as in Figure 71. In two specimens the distal prolateral macroseta is absent, and in one only this macroseta and the most proximal one are present.
Females. The seminal receptacles of the three females are il- lustrated to show the variation in the prominence of the bowl. Fig- ure 89 has large bowls which are distinct from the stalks, Figure 88 has slightly less prominent bowls, and Figure 87 has smaller bowls less clearly distinct from the stalks.
DISTRIBUTION.
Southern Illinoi,~ and eastern Missouri (Map I ) .



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190 Psyche [June
RECORDS. ILLINOIS. Jackson Co.: Little Grand Canyon, about 5 mi. W. of Pomona, 12 Oct. 1967, Q (J. A. Beatty) . - Johnson Co.: Ferne Clyffe St. Pk., 13-14 Sept. 1966, 2 d\ 59 (FAC). - Union Co.: Pine Hills, about 3 mi. N.E. of Aldridge, 7-14 Oct. 1967, d ; 14-20 Oct. 1967, g ; 27 Oct. - 3 Nov. 1967, 3 C? (J. Nelson). MISSOURI. Dent Co.: Montauk St. Pk., 12 Sept. 1966 (FAC).
LITERATURE CITED
BANKS, N.
1896.
New Californian spiders. Jour. New York Ent. Soc. 4(4): 88-91. COMSTOCK, J.H.
1912. The Spider Book. New York. 721 pp. 1940. op. cit., rev. ed., Ithaca, N. Y. 729 pp. PICKARD-CAMBRIDGE, 0.
1883. On some new genera and species of spiders. Proc. 2001. Soc. Lond. pp. 352-365.
RIVERS, J. J.
1891. Description of the nest of the Californian turret building spider, with some references to allied species. Zoe 2: 318-320. SIMON, E.
1890. Liste des espkces de la famille des Aviculariides qui habitent 1'AmGrique du Nord. Actes Soc. Linn. Bord. 44: 307-339. SMITH, C. P.
1908. A preliminary study of the Araneae Theraphosae of California. Ann. Ent. Soc. Amer. l(4) : 207-249.
APPENDIX
Abbreviations and definitions of measurements and meristic characters used in this study.
CL Maximum length of carapace (between lines tangent to anterior- most and posterior-most parts of carapace) along line parallel to median longitudinal axis, with lateral border of carapace hori- zontal.
CW Maximum width of carapace along line perpendicular to median longitudinal axis.
IFL Length of femur I taken as length of straight line connecting the proximal and distal points of articulation. All leg and pedipalp segment length measurements were made in side view along retrolateral surface of appendages after removing them from spider. ITL
Length of tibia I taken as length of straight line connecting proximal and distal points of articulation.
IML Length of metatarsus I taken as length of straight line connecting proximal point of articulation with distal-most point of segment. ITarL Length of tarsus I taken as length of straight line connecting most proximal exposed point of tarsus with distal-most point of dorsal surface.
IVFL, IVTL, IVML, IVTarL Leg IV segment lengths measured in same manner as corresponding leg I segments.




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19681
PFL
PTL
PTT
S L
sw
OQW
ALS
ALD
AMS
AMD
CAT
CT
CMT
PCT
ICTP
ICTR
Coyle - Spider Genus Aiypoides
191
Length of palpal femur measured same as IFL. Length of palpal tibia measured same as ITL. Maximum diameter of proximal half of ~alpal tibia perpendicular to line defined by PTL.
Maximum length of sternum on line parallel to median longitudinal axis. Anterior border of sternum is its pointed anterior extension lateral to labium.
Maximum width of sternum perpendicular to line defined by SL. Maximum width of eye group (ocular quadrangle) on line perpen- dicular to median longitudinal axis of carapace. All eye measure- ments are made in dorsal view with lateral border of carapace horizontal.
Minimum distance between anterior lateral eyes. Maximum diameter of left anterior lateral eye. Minimum distance between pupils (light colored saucer-shaped cen- tral area of eye) of anterior median eyes. Transverse diameter of left anterior median eye pupil. Cheliceral apophysis thickness taken as diameter at midpoint of apophysis in lateral view.
Number of cheliceral macroteeth (per chelicera), the large teeth forming main tooth row on chelicera (Fig. 41). Number of cheliceral microteeth (per chelicera) , the much smaller teeth scattered on ectal side of macrotooth row (Fig. 41). Number of teeth on claw of female pedipalp. All claw tooth counts include even the reduced proximal teeth. Counts must be made carefully under high magnification in strong light. Number of teeth on prolateral claw of left tarsus I. Number of teeth on retrolateral claw of left tarsus I. IVCTP Number of teeth on prolateral claw of left tarsus IV. IVCTR Number of teeth on retrolateral claw of left tarsus IV. PTSP
Number of enciform macrosetae on prolateral surface of tarsus of female pedipalp.
PTSR Number of enciform macrosetae on retrolateral surface of tarsus of female pedipalp.
IMS Number of enciform macrosetae on metatarsus of left leg I of fe- male.
IVMT Number of trichobothia in row on dorsal surface of metatarsus IV. IVMCR Number of setae in large comb on retrolateral surface at distal end of metatarsus IV.
EGS Number of epiandrous gland spigots. These are located just an- terior to genital opening on abdomen of adult males.



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Psyche
Table I. Measurements, meristic characters, and diagnostic ratios for adult males of each species of Atypoides. Figures represent range, mean, and standard deviation in mm. Range and mode given fcr meristic characters.
riversi gertschi hadros
N
CL
cw
I FL
ITL
IML
ITarL
IVFL
IVTL
IVML
IVTarL
OQW
ALS
ALD
AMS
AMD
PFL
PTL
P IT
riversi gertschi hadroi
CAT
ICTP
ICTR
1VCTP
IVCTR
EGS
IML
-
CL
IVTL
-
CL
IVML
-
CL
IVTarL
CL
OQ W
-
CL
CL
-
AMU
PTT
-
CL
CAT
-
CL
I FL
-
1ML
IML
IVML
-
IVTarL
PTT
-
P I'L




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CoyIe - Spider Genus Atypoides
Table 11. Measurements, meristic characters, and diagnostic ratios for adult females of each species of Atyfoides. Figures represent
range, mean, and standard deviation in mm. Range and mode given for meristic characters. Number of '(reproductively active" females in parentheses after N.
riversi gcrfichi hadros
riversi gcrtschi hadros
N
CL
cw
IFL
ITL
IML
ITarL
IVFL
1VTL
IVML
IVTarL
SL
sw
OQW
ALS
ALD
AMS
AMD
CT
CMT
PCT
ICTP
1CTR
IVCTP
1VCTR
PTSP
PTSR
IMS
IVMT
IVMCR
IML
-
CL '
ITarL
-
CL
IVML
-.
CL
OQW
A
CL
CL
-
AMD
I FL
-
IML
IVML
-
1ML
SL
-
sw
1VMCR
-
IMS




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194 Psyche [June
Table 111. Measurements and meristic character values for the type speci- mens of the species of Atypoides. Measurements in mm. riversi gertschi hadros
(lecto-
(holo- (holo-
type) type) type)
riversi
gertschi hadros
(lecto- (holo- (holo-
type) type) type)
CL 5.5
CW 4.29
I FL 5.35
ITL 3.58
IML 5.01
ITarL 3.05
IVFL 5.05
IVTL 3.46
IVML 5.24
IVTarL 3.65
0,QW 1.24
ALS 0.60
ALD
AMS
AMD
PFL
PTL
PTT
CAT
ICTP
ICTR
IVCTP
IVCTR
EGS




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Volume 75 table of contents