A Cladistic Analysis of the Opilionid Superfamily

Home , Acuclavella, Caddidae, Ceratolasmatidae, Cosmetidae, Dicranolasmatidae, Dyspnoi

AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2844, pp. 1-29, figs. 1-41, tables 1-9, maps 1-3 June 10, 1986

A Cladistic Analysis of the Opilionid Superfamily Ischyropsalidoidea, with Descriptions of the New Family Ceratolasmatidae, the New Genus Acuclavella, and Four New Species

WILLIAM A. SHEAR'

ABSTRACT A cladistic analysis ofthe opilionid superfamily lasmatidae is described as new for the genera Cer- Ischyropsalidoidea revealed three groups of gen- atolasma, Hesperonemastoma, Crosbycus, and era, each probably monophyletic, and recognized Acuclavella, new genus. Acuclavella consists offour here as families. The family Ischyropsalididae Si- new species from Idaho and Washington: cosme- mon is redefined to include only the genus Ischy- toides, shoshone, merickeli, and quattuor. Cros- ropsalis, the family Sabaconidae Dresco is ex- bycus dasycnemus (Crosby) is redescribed and il- panded to include Sabacon (=Tomicomerus, new lustrated, and new distribution records are given. synonymy) and Taracus, and the family Cerato-

INTRODUCTION The opilionid family Ischyropsalididae has 1904), but nothing substantial was done re- been problematical for opilionologists for garding its composition or relationships until many years, with a piecemeal approach being Martens (1969) revised the type genus, dis- taken to the inclusion or exclusion ofvarious posing ofa vast plethora ofextraneous species taxa in the family. Originally proposed by names proposed by Roewer (1950). For the Simon (1879) to cover the genera Ischyrop- first time, Martens' work allowed some clear salis, Sabacon, and Taracus, the family was insights into the true nature ofthis taxon. For readily recognized (Hansen and Sorensen, example, he established that Ceratolasma tri-

I Research Associate, Department of Entomology, American Museum of Natural History; Professor of Biology, Hampden-Sydney College, Hampden-Sydney, Virginia 23943. Copyright © American Museum of Natural History 1986 ISSN 0003-0082 / Price $3.20 2 AMERICAN MUSEUM NOVITATES NO. 2844 cantha, originally described by Goodnight and gestion of a family for Crosbycus alone, he Goodnight (1942) as a trogulid (on the mis- saw in the future. taken assumption it was related to Ortholas- Work with Sabacon, Crosbycus, Cerato- ma, now known to be a nemastomatid, but lasma, and Hesperonemastoma had suggest- at that time considered a trogulid also), was ed to me that there might be a solution to the in fact related at some level to Ischyropsalis, evident difficulties in the taxonomy of this while Taracus and Sabacon were more dis- family, and when evidence became available tantly so. A nonconforming element of the a few years ago ofan exciting new genus from family was removed when Martens and Su- Idaho and Washington related to Ceratolas- zuki (1966) showed that Nipponopsalis was ma, I resolved to try to make some sense of in fact allied in some way to the trogulids. relationships in the Ischyropsalidoidea. Later, Gruber (1978) and Shear and Gruber (1983) placed Ruaxphilos petrunkevitchou Goodnight and Goodnight (1945) in the syn- ACKNOWLEDGMENTS onymy of Ortholasma bolivari, an ortholas- I thank, first of all, Frank Merickel of the matine nemastomatid. University ofIdaho for his generosity in pro- Dresco (1970) suggested a monotypic fam- viding specimens ofAcuclavella and notes on ily for Sabacon, and this proposal was ac- their habits. He was referred to me by my cepted by Martens (1983), who characteris- colleague in American opilionid taxonomy, tically provided much better evidence for the James C. Cokendolpher (Texas Tech Uni- idea in his revision of European Sabacon. versity). I had first been made aware of the However, Martens did not include Taracus. existence ofthis new genus by Rod Crawford Gruber (1970) showed that the North ofthe Burke Memorial Museum, Seattle, who American species described by several au- sent a single female specimen from Lewis thors in Nemastoma were not even members County, Washington. The scanning electron of Nemastomatidae but related to Ischyrop- micrographs illustrating this paper were tak- salis; he gave them the new (unfortunate) en at the Virginia-Maryland College of Vet- name Hesperonemastoma. In several publi- erinary Medicine, through the kindness of cations, Gruber (1970) and Martens (1976, Brent Opell. Additional specimens of ischy- 1978) presented evidence for including Cros- bycus in the ischyropsalidid assemblage; ropsalidoids were provided by Cokendol- Martens persistently suggested a monotypic pher, by Jochen Martens (Johannes-Guten- new family for it, but has never actually used berg-Universitat, Mainz), and by Nobuo the name "Crosbycidae." Tsurusaki, Hokkaido University, Japan. Jiur- In two important papers, Martens (1976) gen Gruber (Natural History Museum, Vi- and Martens et al. (1981) used, among other enna [VM]) very kindly turned over to me characters, details of the penes and oviposi- his voluminous notes on Crosbycus dasy- tors to produce a widely accepted cladogram cnemus, and commented extensively on an ofthe order Opiliones. It was not unexpected earlier draft of this paper, as did Martens, that their analysis led to a dismemberment Cokendolpher, and Norman Platnick of the of the old "tribe" Dyspnoi, in which Ischy- American Museum of Natural History. Ma- ropsalididae had been allied with Nemasto- terial for study came from my own collection matidae, Dicranolasmatidae, Nipponopsali- and that ofthe American Museum ofNatural didae, and Trogulidae. Martens raised the History (AMNH; Norman Platnick), the Field taxon to superfamily level and placed it close Museum of Natural History (FMNH; Henry to the superfamilies Phalangioidea and Cad- Dybas and John Kethley), the Burke Me- doidea. morial Museum of the University of Wash- However, as Gruber (1978) had pointed ington (BMM; Rod Crawford), the Museum out, the single family Ischyropsalididae is one ofComparative Zoology at Harvard Univer- of the most heterogeneous taxa in Opiliones. sity (MCZ; H. W. Levi), and the personal He decried the solution of monotypic fami- collection of James C. Cokendolpher (JCC). lies for each genus, which, considering the This research was aided by grants from the Sabaconidae of Dresco, and Martens' sug- Faculty Research Fund of Hampden-Sydney 1986 SHEAR: ISCHYROPSALIDOIDEA 3

College, and by equipment purchased under and inserting by means of a long tendon on a grant (J-3 1) from the Jeffress Trust. the base of the glans. The penis, as in many other ischyropsalidoids, phalangioids, and nemastomatoids, bends at the base of the FAMILIES OF glans and the base of the stylus when this ISCHYROPSALIDOIDEA muscle contracts. According to Martens this The superfamily Ischyropsalidoidea has arrangement of the penial muscle and "weitere Merkmale, die hier nicht diskutiert wer- been defined by Martens (1976) and by Mar- den" justify a monotypic family for Sabacon. tens et al. (1981) on the basis of characters ofthe genitalia and palpi. The ovipositor has The additional characters are not given. As secondarily lost its segmentation (retaining Martens pointed out, at least one genus in vestiges ofsegmentation in the musculature), each of the families Phalangiidae and Nem- astomatidae have a very similar arrange- and the palpal claw is vestigial or absent. In ment, and just as importantly, there are at this study, I add to these characters the pres- least two other apomorphies (dense distri- ence of unique sensory cones on the meta- bution of plumose setae on the tibia and tar- peltidial midline. The superfamily corre- sponds roughly in extent to the old "family," sus of the palp, with the tarsus curved back on the of a slot in the the history of which was reviewed to 1976 tibia; presence deep by Gruber (1978). According to the analysis carapace between the chelicerae) shared by and Taracus, and Taracus does not of Martens, the sister group of Ischyropsali- Sabacon doidea is Caddoidea + Phalangioidea. Unit- have the penis muscle as in Sabacon. I there- of the in Sabacon ing these three taxa are two synapomorphies: fore regard the form penis as the one muscle in the penis and plumose setae an autapomorphy only for genus. shows that Sa- (figs. 2-9) on the palpi. This latter character, My cladistic analysis (fig. 1) and taken are the previously thought to occur primarily, if not bacon Taracus, together, exclusively, in the Ischyropsalidoidea, has sister group of all the other Ischyropsalidoi- basis I have decided to been verified for a number of species of Pha- dea. On this recognize a which includes these langioidea (Tsurusaki, personal commun.; family Sabaconidae, two genera. Cokendolpher, personal commun.; unpub- lished data) and Caddoidea (see fig. 5). How- Similarly, Ischyropsalis (with its isolated in central and western is ever, such setae may be synapomorphic for position Europe) shown to be the sister ofthe Opiliones as a whole, since scanning electron group remaining genera. I choose to incorporate this situation microscopy ofthe major palpal setae of some Laniatores has revealed minute tubular ducts in the classification by limiting the family to the like those illustrated here. A more compre- Ischyropsalididae only genus Ischy- and a family Ceratolas- hensive survey of seta types in Opiliones is ropsalis, recognizing needed. matidae which includes four genera. The type some How many families of Ischyropsalidoidea genus was selected because there is still should there be? Dresco (1970) suggested two: ambiguity about the position of Crosbycus, and Ceratolasma is the next oldest name in Ischyropsalididae and a monotypic Saba- conidae. This idea has been criticized and the group. The revised classification is as fol- rejected by Shear (1975) and Gruber (1978), lows: but lately accepted by Martens (1976, 1978, Superfamily Ischyropsalidoidea Simon 1981), who has also called for the establish- Family Ischyropsalididae Simon ment of a monotypic family based on Cros- Ischyropsalis C. L. Koch bycus. Family Ceratolasmatidae, new While Dresco's arguments for a monotypic Ceratolasma Goodnight and Goodnight family based on Sabacon were not convinc- Acuclavella, new ing (Shear, 1975), Martens (1981) has drawn Crosbycus Roewer attention to the unusual penis muscle in Sa- Hesperonemastoma Gruber bacon. The muscle actually lies outside the Family Sabaconidae Dresco shaft ofthe penis, originating on the two arms Sabacon Simon of a fork extending from the base ofthe shaft Taracus Simon 4 AMERICAN MUSEUM NOVITATES NO. 2844

Ischyropsalis Acuclavella Ceratolasma Crosbycus Hesperonemastoma Sabacon Taracus

ISCHYROPSALID IDAE CERATOLASMATIDAE SABACON IDA' 111 12, 13, 14 910 1

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THE GENERA OF Acuclavella is the closest relative of Cera- ISCHYROPSALIDOIDEA tolasma, as shown by my cladistic analysis. At present the species of this genus are The larger genera within the superfamily grouped together on the basis ofthe cuticular Ischyropsalidoidea are well defined. How- microsculpture of large, rounded warts. The ever, the monotypic genera Ceratolasma, penis shows slight torsion in the stylus, but Crosbycus, and Tomicomerus present certain this may not be a consistent difference. The problems. Placing the species concerned in plesiomorphic characters mentioned above monotypic genera implies that they are the for Ceratolasma are also found in Acuclavella sister groups ofall the other species included (and Ischyropsalis). The question to be an- in the most closely related genus or group of swered here is whether or not Acuclavella and genera. Ceratolasma should be merged. Possible Ceratolasma tricantha as a species has at characters to unite them could be the orna- least two significant autapomorphies: trogu- mentation of the leg femora (also found in loid facies and the presence of a palpal/che- Hesperonemastoma), the dorsal ornamenta- liceral stridulatory apparatus. These are not tion, and the extended eye tubercle, but C. found in any other ischyropsalidid species. tricantha is, in fact, strikingly different from Interestingly, these characters coexist with a the species ofAcuclavella. In my opinion, the number ofsignificant plesiomorphic features: single species included in Ceratolasma is in- the simple penis, endites on all coxae, and a deed the sister group ofthe rest ofthe species, large sternum and labium (Gruber, 1978). and the information retrieval aspects of the 1986 SHEAR: ISCHYROPSALIDOIDEA 5

TABLE 1 Characters for Cladistic Analysis of Ischyropsalidoidea Characters Plesiomorphic Apomorphic 1. Metapeltidial sensory cones Absent Present 2. Ovipositor Segmented Segmentation lost 3. Palpal claw Large Reduced or absent 4. Cheliceral glands of males Absent Present 5. Genital operculum No suture With suture 6. Palpus Gracile Tibia and tarsus enlarged 7. Front of carapace Not indented Deep slot in midline 8. Sclerotization Heavy Reduced 9. Legs With setae and microtrichia With raised scalelike tubercles 10. Abdominal scutum Without spikes With spikes 11. Chelicerae Normal size Enlarged and spiny 12. Metapeltidial sensory cones 2 More than 2 13. Plumose setae of palpus Present Lost 14. Seminal receptacles 4 More than 4 15. Eye tubercle Rounded, low With median spike 16. Cheliceral glands of males Present Lost 17. Metapeltidial sensory cones Present Reduced or lost 18. Microsculpture of scutum Small acute denticles Anvil-shaped tubercles 19. Microsculpture of scutum Small acute denticles Large, round warts 20. Troguloid facies Absent Present 21. Stridulatory mechanism None Cheliceral/palpal 22. Spikes of abdominal scutum Erect, acute Recumbent, blunt 23. Macrosetae of legs Erect Recumbent, sinuous 24. Scutum Parvum Magnum 25. Palpal femora of males Without glands With glands 26. Muscles of penis In truncus Not in truncus 27. Metapeltidial sensory cones 2 1 classification are better served by retaining dasycnemus, but again, that species is ex- both names. traordinarily different from them, and again, Crosbycus dasycnemus is likewise difficult I argue that this information is best conveyed to place. Some clear affinities with Hespero- by maintaining the genus Crosbycus for this nemastoma are seen in the presence ofanvil- single species. shaped tubercles as cuticular macrosculpture The case of Tomicomerus, however, is dif- (fig. 17) and in the evident loss of the meta- ferent. The included species, T. bryanti, is peltidial sensory spines (fig. 30). Unlike Is- known at this time from a single mangled chyropsalis, Ceratolasma, andAcuclavella, the male specimen. In my 1975 revision, I con- palpi of Crosbycus and Hesperonemastoma sidered Tomicomerus intermediate between have plumose hairs (figs. 6, 8). Evidence might Taracus and Sabacon. A careful reexamina- be garnered concerning the relationships of tion of the data I collected at that time, to- C. dasycnemus if males were present, since gether with more information about the range the penes of Hesperonemastoma species are of species in Sabacon (Suzuki, 1974; Mar- highly apomorphic-but this species is al- tens, 1972, 1983; Cokendolpher, 1984), has most certainly parthenogenetic. Of the per- convinced me that there are no striking aut- haps 100 specimens in collections, all are fe- apomorphies in this species that could justify males, and the ovipositors lack seminal (as in the above cases) the retention of a receptacles (fig. 41). The microsculpture and monotypic genus by suggesting that T. bryan- setation of the legs of C. dasycnemus are ti is the sister group of all other Sabacon. unique (fig. 12). The species ofHesperonem- Accordingly, I now consider Tomicomerus a astoma are the closest relatives of Crosbycus synonym of Sabacon. 6 AMERICAN MUSEUM NOVITATES NO. 2844

EVOLUTION AND RELATIONSHIPS contrast to the Phalangioidea, Caddoidea, and Cyphophthalmi, where segmentation has DISCUSSION OF CHARACTERS been retained. The basal number of seminal In the following discussion, I have again receptacles for opilionids is not known, but taken up the principles set forth in Shear and is probably two or four (Martens et al., 1981). Gruber (1983) for the determination of po- In any case, by outgroup comparison, the larity in transformation series in opilionid presence in many species of Ischyropsalis of groups, namely: (1) occurrence in outgroups more than four is apomorphic (character 14). speaks for plesiomorphy, (2) less differen- Palpal Characters. The palpal claw of all tiated, more homonomously patterned char- ischyropsalidoids has been reduced to a small acters are plesiomorphic, (3) states resem- nubbin or entirely lost (character 3); this is bling those in juveniles are plesiomorphic, apomorphic by outgroup comparison, as well (4) characters known to be consistently cor- as by the presence of claws on the serially related with others known to be apomorphic homologous legs. The plumose setae of the are likely also to be apomorphic, and (5) cor- palpi (figs. 2-9) are plesiomorphic for the su- relations between morphological and ecolog- perfamily (synapomorphic for Ischyropsali- ical or distributional characters should be used doidea and its sister group), so their loss, cautiously. which has taken place twice (character 13) is I have accepted the arguments of Martens apomorphic. In Sabaconidae, a key synapo- and Martens et al. (1981) that morphy (because it makes the palpi even less (1976, 1981) like legs) is the very dense distribution of the immediate outgroup of Ischyropsalidoi- plumose setae, the swollen tibia and tarsus, dea is Phalangioidea and Caddoidea. and the tarsus reflexed on the mesal surface of the tibia. I regard this suite of adaptations LIST OF CHARACTERS as functionally linked, even though the function is not known (character 6). In the genus The numbering of the characters below is Hesperonemastoma, males have the second- the same as in table 1 and in figure 1. ary sexual character of a swollen palpal fe- Metapeltidial Sensory Cones. These unique mur, with a gland (character 25); this is a sense organs, the function of which can only synapomorphy for the genus found nowhere be guessed at (Juberthie et al., 1981), are found else in the superfamily, though there is a par- only in the Ischyropsalidoidea and constitute allel development in males of Ortholasma an unequivocal synapomorphy for the group (Nemastomatidae; Shear and Gruber, 1983). (character 1). In Crosbycus and Hesperonem- This character has not been used in the anal- astoma, these organs have been lost, or are ysis, but both Crosbycus and Hesperonemas- in the process of being lost (character 16). In toma have palpi nearly as long as the first Hesperonemastoma modestum, examination legs. under high magnification shows a pair of Chelicerae. The presence of cheliceral depressions on either side of the midline of glands (Martens and Schawaller, 1977) in the metapeltidium; these are probably the re- males (character 4) is apomorphic by out- duced sensory cones. There is no trace ofthese group comparison. In Hesperonemastoma, structures in Crosbycus dasycnemus (fig. 38). these glands have been lost (character 16), Two additional apomorphic modifications probably to be replaced functionally by the also occur; in Taracus species, the number palpal glands. Greatly enlarged, spiny chelic- ofcones is reduced to one (character 27), and erae have been developed twice in the su- in most Ischyropsalis species, multiple cones perfamily (Ischyropsalis and Taracus, char- occur (character 12). However, it would re- acter 11), as has happened sporadically in quire an ultrastructural examination to prove virtually every opilionid superfamily. I re- that the midline metapeltidial structures in gard this as parallelism here, since no other Ischyropsalis are all sensory cones. synapomorphies suggest a sister-group rela- Ovipositor. According to Martens et al. tionship between Ischyropsalis and Taracus. (1 98 1), the ovipositor ofischyropsalidoids is In particular, the pattern ofcheliceral muscle secondarily unsegmented (character 2), in origins on the carapace is quite different, since 1986 SHEAR: ISCHYROPSALIDOIDEA 7 0I§ 1X

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FIGS. 2-5. Scanning electron micrographs of parts of palpal tibiae. All bars = 10 gim. 2. Acuclavella merickeli (distal to lower right, x 975). 3. Ceratolasma tricantha (distal to upper right, x 975). 4. Ischy- ropsalis hellwigi (distal to lower right, x 430). 5. Acropsopilio chomulae (distal above, x 975). in Ischyropsalis the big muscles operating the should be regarded as apomorphic wherever basal article ofthe chelicera originate behind it occurs. Here, it is regarded as a synapo- the eye tubercle, whereas in Taracus these morphy for Sabaconidae (character 8), where muscles originate on either side ofit. In Cer- it is especially significant in affecting the car- atolasma, the chelicera is part of a unique apace and metapeltidium in many species cheliceral/palpal stridulating mechanism (Shear, 1975; Martens, 1983). Reduction of (character 21; see Gruber, 1969, 1978). sclerotization to a much lesser degree, and Genital Operculum. By outgroup compar- primarily in the abdomen, is found in some ison, a suture (partial or complete) setting off species of Ischyropsalis (Martens, 1969). An the genital operculum from the second ab- additional synapomorphy for Sabaconidae is dominal sternite is apomorphic (character 5). the slot in the frontal midline ofthe carapace Sclerotization and Ornamentation of Dor- extending in most species nearly to the eye sum. Outgroup comparison at a basic level tubercle (character 7). On the other hand, fu- in the cladogram of Martens et al. (1981) sion of the carapace and abdominal scutum forces one to the conclusion that heavy scler- (scutum magnum) is clearly an alternative otization of the body is an apomorphy only synapomorphy (by principle 4 above, only in at the level ofthe order Opiliones as a whole, Hesperonemastoma; character 24). Having and therefore a reduction in sclerotization serially arranged, paired spikes on the scutum 8 AMERICAN MUSEUM NOVITATES NO. 2844

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FIGS. 6-9. Scanning electron micrographs of parts of palpal tibiae. All x 990, bars = 10 ,um. 6. Crosbycus dasycnemus (distal above). 7. Taracus pallipes (distal to upper left). 8. Hesperonemastoma modestum (distal to lower left). 9. Sabacon occidentalis (distal to lower right). is synapomorphic for Ceratolasmatidae, by and becomes a part of the typical troguloid outgroup comparison (character 19); acute in facies, contributing to the camerostome Hesperonemastoma andAcuclavella, these are (character 20; see Gruber, 1978, and Shear blunt, somewhat recumbent cones in Cera- and Gruber, 1983, for a discussion of the tolasma (character 22). In the heavily scler- troguloid facies). otized species, small, acute thorns on the Legs. Legs ornamented with microtrichia sclerites appear to be the plesiomorphic form and ordinary micro- and macrosetae (figs. 14- of sculpture. In Acuclavella these become 16) are plesiomorphic by principle 2 above; quite large, rounded warts (character 19), and ornamentation of raised scalelike tubercles in Crosbycus and Hesperonemastoma appear subtending setal bases (figs. 10, 1 1, 13) is apo- as anvil-shaped tubercles (figs. 17, 38; char- morphic (character 9). Crosbycus has unique acter 18), a parallelism with Nemastomati- procumbent, curly hairs on the femora, pa- dae. tellae, and tibiae ofthe legs (fig. 12; character Eye Tubercle. Apomorphic by outgroup 23). Martens (1983) has illustrated a pattern comparison, Acuclavella and Ceratolasma ofraised scales on the femora ofsome species carry a forward-slanting spike on the eye tu- of Sabacon, but these raised scales do not bercle (character 15). In Ceratolasma this subtend setae and are clearly a different de- spike extends virtually horizontally forward velopment. 1986 SHEAR: ISCHYROPSALIDOIDEA 9

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FIGS. 10-13. Scanning electron micrographs ofparts ofsecond leg femora. All bars = 100(,um, except for figure 12, where bar = 10 ,um. 10. Acuclavella merickeli (distal to upper left, x 100). 11. Hespero- nemastoma modestum (distal to lower right, x 235). 12. Crosbycus dasycnemus (distal to upper left, x 225). 13. Ceratolasma tricantha (distal to lower right, x 105).

CLADISTIC ANALYSIS tion on families, the family Sabaconidae is accepted, but expanded to include Taracus. The relationships of the ischyropsalidoid Tomicomerus is regarded as a synonym of families and genera are illustrated in figure Sabacon. 1; the character numbers refer to those found Crosbycus and Hesperonemastoma remain in table 1. The base of the cladogram is, un- difficult elements. Each has a large number fortunately, only weakly resolved by regard- of autapomorphies, and in Crosbycus, these ing the suture of the genital operculum as are coupled with substantial numbers of synapomorphic for Ischyropsalididae and primitive characters. However, ifthey are in- Ceratolasmatidae. The suggestion ofMartens deed ischyropsalidoids (as indicated for at (1969) and Gruber (1978) that Ceratolasma least Hesperonemastoma by the single penis is the closest relative of Ischyropsalis is not muscle and secondarily unsegmented ovi- borne out by this analysis; the characters sug- positor), they belong in the cladogram as gested as supporting the relationship have shown. Martens (1976, 1981) has repeatedly turned out to be plesiomorphies (form ofster- suggested a monotypic family for Crosbycus num and endites, simple penis). but has never formally used the name "Cros- For reasons already discussed in the sec- bycidae." Because of my uneasiness about 10 AMERICAN MUSEUM NOVITATES NO. 2844

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FIGS. 14-17. Scanning electron micrographs. All bars = 100,m. 14-16. Parts of second leg femora. 14. Taracus pallipes (distal to left, x 85). 15. Sabacon occidentalis (distal to lower right, x 120). 16. Ischyropsalis hellwigi (distal below, x 105). 17. Anal operculum of Crosbycus dasycnemus, showing ornamentation of anvil-shaped tubercles on heavily sclerotized parts and trifid scales on membranous parts (x 300). the position ofthe genus, and a prudent wish Ischyropsalidoidea Martens, 1976, p. 64; 1978, p. to base the family name on the oldest generic 186. Gruber, 1978, p. 107. Martens, Hoheisel, name other than Crosbycus, I have decided and G6tze, 1981, p. 37. to call the family Ceratolasmatidae. TYPE GENUS: Ischyropsalis C. L. Koch, 1839. DIAGNOSES OF TAXA DIAGNOSIS: Palpatond opilionids with single muscle in penis; ovipositor secondarily The literature citations given below for de- unsegmented, with common canal for ce- scribed taxa are not meant to be exhaustive, ment glands; metapeltidium with one or more but include the original proposal and sub- sensory cones ofunknown function (these lost sequent substantive discussions. in some genera); males with cheliceral or palpal glands probably producing sex phero- SUPERFAMILY ISCHYROPSALIDOIDEA SIMON mone (this well documented for Ischyrop- Ischyropsalidae Simon, 1879, p. 265. Roewer, salis; see Martens, 1969); palpi usually with 1923, p. 680. Comstock, 1913, p. 80. plumose setae, lacking claw. 1986 SHEAR: ISCHYROPSALIDOIDEA I1I

INCLUDED FAMILIES: Ischyropsalididae Si- bium scarcely sclerotic at all; anterior midline mon; Ceratolasmatidae, new; and Sabacon- of carapace deeply indented to form slot idae Dresco. reaching from margin to eye tubercle; one or DISTRIBUTION: Southwestern, central, and two metapeltidial sensory cones; genital southeastern Europe; central Asia, the Hi- operculum not set off by suture; chelicerae malayas, and Siberia; Japan; western North normal or greatly enlarged, spiny, with or America from southern Alaska to San Fran- without gland in males; palpi with tibia and cisco Bay; in the inland mountains ofcentral tarsus slightly or greatly inflated, very densely Idaho and western Montana, the Sierra Ne- set with plumose setae of uniform length vada, and the Rockies from Alberta to Utah, ("bottle-brush" palpi), tarsus with narrow New Mexico, and Arizona; sporadic in Mis- basal stem, reflexed against asetose area on sissippi River drainage from Minnesota and median surface oftibia. Legs lacking cuticular Michigan to Mississippi; Appalachian sculpture. Mountains from New Hampshire to Georgia. INCLUDED GENERA: Sabacon Simon, Tar- acus Simon. Sabacon (type species S. paradoxum Simon) has been the subject of revi- FAMILY ISCHYROPSALIDIDAE SIMON sions by Suzuki (1974, Japanese species), Ischyropsalidae Simon, 1879, p. 265. Roewer, Shear (1975, American species), and Martens 1923, p. 680 (in part). Martens, 1969, p. 133; (1972, Himalayan species; 1983, European 1978, p. 186; Martens, Hoheisel, and G6tze, species). The best diagnosis is in Martens 1981, p. 37. Shear, 1975, p. 7. Gruber, 1978, p. (1983). Taracus has never been given a com- 107. parative diagnosis in the context of the su- TYPE GENUS: Ischyropsalis C. L. Koch, perfamily, and this is done below. 1839. DISTRIBUTION: For Sabacon only, see the map in Martens (1983, p. 267); Pacific coast DIAGNOSIS: Ischyropsalidoidea with great- of North America from southern Alaska to ly enlarged spiny chelicerae; carapace heavily south of San Francisco Bay; Sierra Nevada sclerotized, with strip ofcuticle extending be- Mountains; Appalachians from New Hamp- tween chelicerae; sclerotization of abdomen shire to North Carolina (isolated records from variable; eye tubercle deeply grooved; genital Ohio, Illinois, Minnesota, Iowa, Wisconsin, operculum separated from second abdominal Michigan, and Arkansas); Pyrenees and sternite by suture; palpi without plumose se- French and Italian Alps; Tun- tae; legs without cuticular sculpture (fig. 16); Himalayas; penis basally broad, muscle in truncus, stylus guska and Altai, Siberia; Korea, Manchuria, usually deflexed; ovipositor with 4 to 12 sem- and Primorje (USSR); Japan. For Taracus, inal receptacles. see below. INCLUDED GENUS: Ischyropsalis C. L. Koch. The genus (type species I. kollari C. L. Koch) Taracus Simon, 1879 is diagnosed, and all known species described, in the revision by Martens (1969). Taracus Simon, 1879, p. 277. Comstock, 1913, p. DISTRIBUTION: Central Europe and the Pyr- 81. Roewer, 1923, p. 693; 1950, p. 52. Shear, enees. 1975, p. 8. TYPE SPECIES: T. packardi Simon, 1879. FAMILY SABACONIDAE DRESCO DIAGNOSIS: Sabaconids with greatly enlarged, spiny chelicerae lacking gland in males; Ischyropsali[di]dae, Roewer, 1923, p. 680 (in part). labrum flat; metapeltidium usually not scler- Comstock, 1913, p. 80 (in part). Shear, 1975, p. otized, metapeltidial sensory cone single, in 7 (in part). Suzuki, 1974, p. 83 (in part). Sabaconidae Dresco, 1970, p. 1210. Martens, 1978, median line; abdomen not sclerotized, or with p. 225; 1983, p. 269. small sclerotized patches, to scutum lami- natum; sternum reduced to two small plates TYPE GENUS: Sabacon Simon, 1879. with single setae; palpi long, slender, with DIAGNOSIS: Ischyropsalidoids with gener- distal teeth on patella in males; legs without ally reduced sclerotization, sternum and la- microtrichia on femora (fig. 14); penis with 12 AMERICAN MUSEUM NOVITATES NO. 2844

*'\- 1 i. I ''/.

* ~K / .

MAP 1. Part ofNorth America, showing distribution ofSabacon species (circles) and Taracus species (squares). Sabacon also occurs in southern Alaska.

MAP 2. Part of North America, showing distribution of Hesperonemastoma species (squares) and Ceratolasma tricantha (circles). 1986 SHEAR: ISCHYROPSALIDOIDEA 13 muscle in truncus; ovipositor slightly longer ber, 1978), Crosbycus Roewer (diagnosis be- than wide. low), Hesperonemastoma Gruber (see DISTRIBUTION: Seven ofthe eight described diagnosis in Gruber, 1970), Acuclavella, new. species (packardi, spinosus, pallipes, silves- DISTRIBUTION: North America, from trii, gertschi, nigripes, and malkini) are lim- southern British Columbia to central Cali- ited to western North America, but their ex- fornia (?); central Idaho; western Montana; act distribution remains uncertain without a Mississippi Valley from southern Illinois to revisionary treatment. The range includes, northern Mississippi; southern Appalachians however, most of montane California, west- from southwestern Virginia to northern ern Oregon and Washington, central Colo- Georgia and Alabama. rado (Utah?), northern New Mexico, and northwestern Arizona. A single species, T. new genus birsteini, was described by Ljovushkin (1971) Acuclavella, from Primorsky Krai, near Vladivostok in TYPE SPECIES: Acuclavella cosmetoides, n. Siberia. sp. ETYMOLOGY: The genus name is a Latin FAMILY CERATOLASMATIDAE, NEW neologism-"acute little spines" -and should be treated as feminine. Ischyropsalidae, Goodnight and Goodnight, 1945, DIAGNOSIS: Closestto Ceratolasma, but dif- p. 5 (in part). Gruber, 1970, p. 130 (in part); fering in lacking lateral anterior extensions of 1978, p. 105 (in part). carapace and in erect, acute (rather than pro- Ischyropsalididae, Martens, 1978, p. 187 (in part). cumbent, blunt) spine ofthe eye tubercle, and Nemastomatidae, Banks, 1894, p. 51; 1901, p. 677 in bearing large, pointed spines on some ab- (in part). Roewer, 1923, p. 650 (in part); 1951, dominal scute areas (Ceratolasma has small p. 95 (in part). bumps on areas 2 and 3 and large blunt spines TYPE GENUS: Ceratolasma Goodnight and on area 4). Goodnight, 1942. DESCRIPTION: Heavily sclerotized, black or DIAGNOSIS: Ischyropsalidoids with very dark brown opilionids of medium size with heavy sclerotization, scutum parvum to mag- sclerotized parts of cuticle ornamented with num; metapeltidium not separate sclerite, distinctive hemispheric warts. Carapace with with two sensory cones or none; carapace and suture immediately behind eye tubercle, sep- scutum heavily ornamented with micro- arating propeltidium from mesopeltidium. sculpture of warts, fine denticles, or anvil- Metapeltidium ("second thoracic tergite") set shaped tubercles; macrosculpture of rows or off by distinct cuticular fold but not by a su- pairs of tubercles on scutum, these some- ture. Metapeltidium with median pair of times very high and acute, or blunt and ap- prominent blunt spines (probably sense or- pressed (Ceratolasma), or absent (Crosby- gans; see Juberthie et al., 1981). Abdomen cus); genital operculum with suture; labium with scutum parvum, first five abdominal and sternum usually well sclerotized; chelic- tergites fused, last three free. Ninth tergite erae with or without (Hesperonemastoma) divided; tenth forms anal operculum. All ab- gland in males, not enlarged; palpi long, with dominal sternites free. Genital operculum (Crosbycus, Hesperonemastoma), or short, sexually dimorphic, clearly delineated by without (Ceratolasma, Acuclavella) plumose transverse furrow that forms membranous setae; legs with microsculpture of apically suture at lateral ends. All coxae with endites. notched, distally raised, scalelike tubercles Sternum broader than long, larger than la- subtending small blunt setae, or acute spikes, bium. Chelicerae and palpi short, chelicerae macrosetae, and recumbent hairs; penis with of males with raised, setose glandular region muscle in truncus; ovipositor about as long on basal article. Vestiture of palpi microtri- as wide, or longer than wide, two or four chia and ordinary setae; claw remnant short, seminal receptacles. very small. Legs moderately long, micro- INCLUDED GENERA: Ceratolasma Good- sculpture offemora, patellae, and tibiae scat- night and Goodnight (see diagnosis in Gru- tered, distally elevated scales infrequently bi- 14 AMERICAN MUSEUM NOVITATES NO. 2844

I / K I

j*s. .2UL7. I

-.---- I / >0 I I .A \ I I

I ,1... ci

. - - r .W. .%, MAP 3. Part of North America, showing distribution of Crosbycus dasycnemus (circles) and species ofAcuclavella (squares).

lobed and subtending short setae. Metatarsi Acuclavella species, on the other hand, have not divided; typical microsculpture limited a robustly convex dorsum with high, sharp at most to basal third, remainder with setae spines on the scute, as well as an erect spine and microtrichia. Tarsi with many segments. on the eye tubercle. There is no impression Penis very simple, distally tapering, glans with of flattening and no suggestion of a camero- sparse microsetae, stylus distinctively twisted stome; the surface of the cuticle is not adapt- in some species. Ovipositor short, with small ed to trap and hold soil particles. There is no apical setae groups, four typical seminal re- stridulatory mechanism. I regard the cutic- ceptacles. ular microsculpture of large, hemispherical DISTRIBUTION: Idaho, Washington. warts as synapomorphic for Acuclavella NOTES: Ceratolasma tricantha (Gruber, species. However, having an eye tubercle 1978) is more specialized than Acuclavella spine at all is a synapomorphy for the two species in a number of obvious ways. It has genera. assumed the troguloid facies (Shear and After due consideration, I have decided to Gruber, 1983) characteristic of soil-dwelling describe four species in this genus. Although opilionids ofmany unrelated genera and fam- each population (except for A. merickeli at ilies, with the profoundly flattened body and Meadow Creek) is represented by a very small typical modifications ofthe carapace and an- sample of one to four animals, the distinc- terior appendages to produce a camerostome. tions in the dorsal armature seem consistent The cuticular sculpture of the body is well within the samples (including Meadow adapted for the trapping of small soil parti- Creek), and there are between-sample differ- cles, a function evidently augmented by an ences in size and proportions. Obviously, fur- adhesive secretion. In addition, there exists ther sampling for this genus in Idaho is re- an unusual chelicera/palpus stridulatory quired to clarify the status of the various mechanism. populations. 1986 SHEAR: ISCHYROPSALIDOIDEA 15 I

FIG.18.Stereo-pairscanning electron moe l e FIG. 18. Stereo-pair scanning electron micrograph ofAcuclavella merickeli female ( x 16).

KEY TO ACUCLAVELLA SPECIES DIAGNOSIS: Easily distinguished from the other species by three pairs of acute spines 1. Dorsum of abdomen with prominent acute on the abdominal scutum. spines ...... 2 DESCRIPTION OF MALE HOLOTYPE: Body (fig. Without such spines, but paired, low, rounded 23) highly arched, convex dorsally, some- tubercles on all five dorsal areas ...... in ...... females of shoshone what rectangular general outline, only 2. A single pair of abdominal spines, on area 2 slightly longer than wide. Nearly all of body ...... merickeli heavily sclerotized, black, with densely scat- More than one pair of abdominal spines .. 3 tered warts; body without pilosity. Total 3. Abdominal spines on area 1 ...... 4 length 4.42 mm; carapace length in midline No spines on area 1, paired spines on areas 2, 1.35 mm, greatest carapace width 2.86 mm; 3, and 4 ...... cosmetoides length of scutum in midline 2.47 mm, great- 4. Four pairs of abdominal spines (on areas 1-4) est width of scutum 3.04 mm...... males of shoshone Eye tubercle (fig. 30) at edge of carapace, Two pairs ofabdominal spines (on areas 1 and 2) ...... females of quattuor oval at base, prolonged into sharp, conical spine standing 1.04 mm above carapace. Eyes new small, dark brown, on rounded basal part of Acuclavella cosmetoides, species tubercle. Carapace surface evenly curved, Figures 19-24, 30, 33, 34, 37 posterior margin arcuate; mesopeltidium (first TYPES: Male holotype and female paratype thoracic tergite) probably fully incorporated, (AMNH) from Pierce, Clearwater Co., Idaho but its side-pieces free, entirely dorsal. Meta- (August 29, 1959; W. J. Gertsch and V. Roth). peltidium (second thoracic tergite) incorpo- ETYMOLOGY: The species name is an ad- rated but clearly delimited by deep fold; sen- jective pointing out the resemblance of this sory cones very prominent, paramedian in species to members of the tropical opilionid position, with shiny brown tips. Ozopores family Cosmetidae. not visible from above, opening in mem- 16 AMERICAN MUSEUM NOVITATES NO. 2844

22 1986 SHEAR: ISCHYROPSALIDOIDEA 17 brane between carapace and first and second TABLE 2 leg coxae. Lengths of Appendage Segments in Holotype Male Scutum ofopisthosoma well-arched, near- Acuclavella cosmetoides ly square in outline when seen dorsally. All (Measurements are in millimeters.) areas with paired tubercles; on area 1 paired Legs small bunches of slightly enlarged cuticular Segment Palpus 1 2 3 4 warts, on areas 2, 3, and 4, tubercles enormously enlarged to form smooth, shining Trochanter 0.52 0.52 0.73 0.52 0.74 spikes with acute tips rising from swollen, Femur 0.78 2.21 3.17 2.00 2.86 nearly circular bases with same warty sculp- Patella 0.65 0.91 1.04 1.04 1.04 1.69 2.24 ture as the scutum surface. Spikes subequal Tibia 0.73 1.66 2.39 Metatarsus - 2.34 3.77 2.42 3.72 in size, but those of area 3 slightly smaller - 4.16 than others. Area 5 with widely separated Tarsus 0.73 2.63 4.94 small tubercles; these barely noticeable on free abdominal tergites. Sclerites of anal region typical, like those uniformly darker than tibia, with setae and of Ceratolasma tricantha. Abdominal ster- dense microtrichia; claw rudiment very small. nites narrow, not modified, with typical Legs with segments long, cylindrical, with sculpture. Spiracles not visible. Genital oper- microsculpture ofevenly spaced, low, flat tu- culum tongue-shaped, separated from second bercles made scalelike by raised distal mar- abdominal sternite by faintly indicated in- gin, widely scattered small setae usually in complete suture. Distal margin rebordered, notch on distal margin of tubercle. All leg glossy. Prosomal sternum 0.09 mm long, 0.13 segments black except metatarsi and tarsi, mm wide, brown, without setae. Labium these bright brown; metatarsi and tarsi lack weakly sclerotized, longer than wide, without tubercles, densely clothed in microsetae. setae. All leg coxae with endites, those oflegs Metatarsi not divided, tarsi divided with III and IV smallest. Endites of leg II about longer basal segment and many short distal twice their size, with single setae. Endites of ones; distitarsus ofthree segments. Claws sin- legs I with free sclerite, oval, brown, setose gle, evenly curved, black, not toothed. (fig. 22). Palpal endites free, with large scler- Penis (figs. 33, 34) 2.5 mm long, widest otized and membranous parts. (0.13 mm) at base of shaft. Shaft evenly ta- Chelicerae (fig. 20) black, surface smooth, pered, broadening slightly at glans. Dorsal shining. Basal article 1.09 mm long, 0.65 mm glans plate with scattered small setae, stylus wide, dorsally with oval, slightly raised glan- spirally twisted, well set offfrom body ofglans. dular area densely set with amber setae. In DESCRIPTION OF FEMALE PARATYPE: Simi- holotype, this area capped with hardened lar in nearly all respects to male, differing in glandular secretion. Palp (fig. 21) with di- dimensions and in secondary sexual charac- mensions given in table 2. Coxae black, ven- ters (fig. 24). Total length 5.2 mm. Midline trally with three seta-bearing tubercles. Tro- lengths: carapace 1.48 mm, metapeltidium chanter very dark brown with ventral row of 0.26 mm, scute 3.30 mm. Widths: carapace four seta-bearing tubercles. Remaining seg- 2.6 mm, scute 3.56 mm (greatest width of ments light yellow-brown except as noted. body). Eye tubercle spike 1. 10 mm above sur- Patella with diffuse darker band on middle face of carapace. third, bearing small setae and few microtri- Sternum 0.13 mm long, 0.31 mm wide. chia distally. Tibia very faintly darkened dis- Chelicerae with basal article lacking glan- tally, with scattered ordinary setae and dense dular area found in male, 1.3 mm long, 0.52 cover of microtrichia. Tarsus slightly and mm wide; distal article 1.3 mm long, 0.44

FIGS. 19-22. Acuclavella cosmetoides. 19. Male, dorsal view. 20. Right male chelicera, lateral view. 21. Right male palpus, mesal view. 22. Anterior ventral surface ofmale, ventral view, genital operculum removed. 18 AMERICAN MUSEUM NOVITATES NO. 2844

TABLE 3 Eye tubercle at edge of carapace, oval at Lengths of Appendage Segments in Paratype base, prolonged into sharp, conical, forward- Female Acuclavella cosmetoides tilted spine standing 0.91 mm above cara- (Measurements are in millimeters.) pace. Eyes small, dark brown, on rounded basal part of tubercle. Features of carapace Legs as in A. cosmetoides. Segment Palpus 1 2 3 4 Scutum of opisthosoma rounded anterior- Trochanter 0.52 0.62 0.65 0.52 0.73 ly, posterior margin nearly square when seen Femur 0.94 2.03 2.83 1.79 2.60 dorsally. All areas with paired tubercles, on Patella 0.65 1.04 1.25 0.96 1.09 areas 1-4, enlarged to form smooth shining Tibia 0.78 1.38 2.08 1.30 1.95 spikes with acute tips, rising from nearly cir- Metatarsus - 1.82 3.17 2.18 3.27 cular, swollen, warty bases; spikes of areas 1 Tarsus 0.68 2.13 4.16 2.60 2.70 and 4 smallest, subequal, those ofareas 2 and 3 nearly twice as large, subequal. Area 5 with paired clusters of four to six slightly larger mm wide. Lengths ofpalpal and leg segments cuticular warts; free tergites without any trace given in table 3. of paired tubercles. Ovipositor (fig. 37) blunt, short, unseg- Anal region, spiracles, and genital oper- mented, symmetrical, with apical tufts of se- culum as in A. cosmetoides. Prosomal ster- tae. num 0.13 mm long, 0.31 mm wide, brown, DISTRIBUTION: Known only from the type without setae. Labium weakly sclerotized, locality. longer than wide, without setae. Endites as in A. cosmetoides. Acuclavella shoshone, new species Chelicerae black, surface smooth, shining. Figures 27, 28, 35 Basal article 1.17 mm long, 0.55 mm wide, TYPES: Male holotype and two female para- dorsally with oval, slightly depressed, rimmed types (AMNH) from Hobo Cedar Grove, glandular area densely set with amber setae. Shoshone Co., Idaho (July 20-29, 1984; F. Gland area not as stongly raised posteriorly W. Merickel). as in A. cosmetoides. Palp with dimensions ETYMOLOGY: The species name is a noun given in table 4. Coxae black, ventrally with in apposition referring to Shoshone County. two seta-bearing tubercles. Trochanter very DIAGNOSIS: Easily distinguished from the dark brown with ventral row of four seta- other species by four pairs of spines on the bearing tubercles. Otherwise palpi as in A. abdominal dorsum in males, and four low cosmetoides. tubercles in the same position in females. Structure and color of legs as in A. cos- DESCRIPTION OF MALE HOLOTYPE: Body (fig. metoides; lengths of segments given in ta- 27) not so highly arched as in type species, ble 4. convex dorsally, more or less oval in general Penis (fig. 35) 3.25 mm long, widest (0.13 outline, significantly longer than wide (length/ mm) at base of shaft. Shaft evenly tapered, width = 1.62). Nearly all of body heavily not broadening at glans. Dorsal glans plate sclerotized, black, with densely scattered small with very few small setae, stylus straight, not warts; body without pilosity. Total length 4.37 perceptibly set off from glans. mm; carapace length in midline 1.30 mm, DESCRIPrION OF FEMALE PARATYPE: Simi- greatest carapace width 2.47 mm; length of lar in nearly all respects to male, differing in scutum in midline 2.50 mm, greatest width dimensions, in secondary sexual characters, of scutum 2.70 mm. and in dorsal ornamentation (fig. 28). Scute

FIGS. 23-30. Structure ofAcuclavella species. 23-29. Lateral view of body. 23. A. cosmetoides male. 24. A. cosmetoides female. 25. A. merickeli male. 26. A. merickeli female. 27. A. shoshone male. 28. A. shoshone female. 29. A. quattuor female. 30. Eye tubercle ofA. cosmetoides male, subdorsal view. Figures 23-29 are not all drawn to the same scale. 1986 SHEAR: ISCHYROPSALIDOIDEA 19

23 24

25 26

27 28

30 29 20 AMERICAN MUSEUM NOVITATES NO. 2844

TABLE 4 TABLE 5 Lengths of Appendage Segments in Holotype Male Lengths of Appendage Segments in Paratype Acuclavella shoshone Female Acuclavella shoshone (Measurements are in millimeters.) (Measurements are in millimeters.) Legs Legs Segment Palpus 1 2 3 4 Segment Palpus 1 2 3 4 Trochanter 0.55 - - - - Trochanter 0.65 0.57 0.73 0.55 0.68 Femur 0.91 2.34 3.17 2.03 2.73 Femur 0.91 1.95 2.86 2.08 2.60 Patella 0.70 0.88 1.17 0.96 1.17 Patella 0.70 0.83 1.35 0.90 1.09 Tibia 0.86 1.69 2.16 1.56 1.95 Tibia 0.78 1.43 2.18 2.08 1.56 Metatarsus - 2.35 3.64 2.60 3.80 Metatarsus - 2.34 3.77 2.47 4.47 Tarsus 0.73 2.50 4.55 2.76 3.64 Tarsus 0.78 2.60 4.29 2.81 3.00

areas lacking characteristic spikes; these re- DIAGNOSIS: Easily distinguished from the placed by low, warty tubercles largest on area other species by having only a single pair of 2, slightly smaller on area 1; those on areas prominent dorsal spikes in both males and 3 and 4 about halfsize oftubercles of area 2. females, and these on area 2. Area 5 with very small bumps at very margin DESCRIPrION OF MALE HOLOTYPE: Body (fig. of scute; first free tergite similar; no bumps 25) not so highly arched as in A. cosmetoides, on second free tergite. convex dorsally, rounded anteriorly, but ap- Total length 5.2 mm. Midline lengths: car- pearing squared off posteriorly; longer than apace 1.69 mm, scute 3.33 mm. Widths: car- wide (length/width = 1.52). Nearly all ofbody apace 2.6 mm, scute 3.38 mm (greatest width heavily sclerotized, black, with densely scat- of body). Eye tubercle spike 0.68 mm above tered warts. Total length 4.16 mm; carapace surface of carapace. length in midline 1.40 mm, greatest carapace Sternum 0.10 mm long, 0.31 mm wide. width 2.60 mm; length of scutum in midline Chelicerae with basal article lacking glan- 2.60 mm, greatest width ofscutum 2.73 mm. dular area found in male, 1.3 mm long, 0.54 Eye tubercle at edge of carapace, oval at mm wide; distal article 1.43 mm long, 0.56 base, prolonged into sharp, conical, forward- mm wide. Lengths ofpalpal and leg segments tilted spine standing 1.06 mm above cara- given in table 5. pace. Eyes small, dark brown, on rounded Ovipositor as in A. cosmetoides. basal part of tubercle. Features of carapace DISTRIBUTION: Known only from the type as in A. cosmetoides. Metapeltidial sensory locality. cones unusually large, much larger than in other species. Acuclavella merickeli, new species Scutum of opisthosoma rounded anterior- Figures 2, 10, 18, 25, 26, 31, 32, 36 ly, posterior margin nearly square when seen TYPES: Male holotype, three male and four dorsally, lateral margins convex. All areas female paratypes (AMNH) from Meadow with paired tubercles, on area 1, as small Creek, 20 miles east-southeast ofLowell, Ida- bunches of slightly enlarged cuticular warts, ho Co., Idaho (August 18, 1984; F. W. Mer- smaller than sensory cones ofmetapeltidium. ickel). Three female paratypes (WAS) from Area 2 with enormously enlarged, shining same locality (May 17, 1983, May 20, 1984; spikes curving slightly posteriad, bases prom- F. W. Merickel), and one additional female inently warty. On areas 3-5, tubercles scarce- paratype (JCC) from same locality (collection ly discernible bunches ofwarts; tubercles not undated; F. W. Merickel). distinct on free tergites. ETYMOLOGY: The species name honors Anal region, spiracles, and genital oper- Frank W. Merickel; with only three excep- culum as in A. cosmetoides. Prosomal ster- tions he has collected all the known speci- num 0.10 mm long, 0.26 mm wide, brown, mens ofAcuclavella species. without setae. Labium well sclerotized, 1986 SHEAR: ISCHYROPSALIDOIDEA 21

TABLE 6 engths of Appendage Segments in Holotype Male Acuclavella merickeli

w,,- (Measurements are in millimeters.)

- o Legs Segment Palpus 1 2 3 4 Trochanter 0.52 0.65 0.73 0.65 0.70 Femur 0.96 2.08 3.43 2.08 2.83 I Patella 0.68 1.04 1.09 0.91 1.09 .11i Tibia 0.83 1.48 2.34 1.51 2.08

-)l Metatarsus - 2.29 3.77 2.55 3.74

I Tarsus 0.73 2.73 4.68 2.80 3.77 .., "z 31' FIG. 31. Scanning electron micrograph ofcheliceral gland mound ofmale Acuclavella merickeli; width of body). Eye tubercle spike 0.94 mm lateral view, x 120. above surface of carapace. Sternum 0.31 mm long, 0.18 mm wide. Chelicerae with basal article lacking glandular area found in male, 1.35 mm long, 0.54 slightly wider than long, without setae. En- mm wide; distal article 1.4 mm long, 0.56 dites as in A. cosmetoides. mm wide. Lengths ofpalpal and leg segments Chelicerae medium brown, surface smooth, given in table 7. shining. Basal article 1.13 mm long, 0.57 mm Ovipositor as in A. cosmetoides. wide, dorsally with oval, slightly depressed, DISTRIBUTION: WASHINGTON: Lewis rimmed glandular area (fig. 31) densely set Co.: 2.3 mi NE of Mossyrock, waterfall on with amber setae. Gland area not more Corn Creek, September 3, 1974 (R. Craw- strongly raised posteriorly than in A. cos- ford), female (BMM). metoides. Palp with dimensions given in table 6. Coxa dark brown, ventrally with two seta-bearing tubercles. Trochanter brown with ventral row of four seta-bearing tubercles. Remaining segments pale yellow, but tarsus shaded darker distally. Structure and color of legs as in A. cosmetoides; lengths of segments given in table 6. Penis (figs. 32, 36) 2.34 mm long, widest (0.09 mm) at base of shaft. Shaft evenly tapered, not broadening at glans. Dorsal glans plate with rows of small setae, stylus spirally curved, well set off from glans, lacking sub- apical tooth. DESCRIPTION OF FEMALE PARATYPE: Simi- lar in nearly all respects to male, differing in dimensions, in secondary sexual characters, and in dorsal ornamentation (figs. 18, 26). Sensory cones smaller, rounder than in male. Spikes of area 2 smaller, less acute than in male; tubercles of areas 3-5 hardly detect- able; free tergites without tubercles. Total length 5.15 mm. Midline lengths: FIG. 32. Scanning electron micrograph ofglans carapace 1.22 mm, scute 3.38 mm. Widths: penis ofAcuclavella merickeli; lateral view, dorsal carapace 2.9 mm, scute 3.51 mm (greatest is to the left (x250). 22 AMERICAN MUSEUM NOVITATES NO. 2844

TABLE 7 DIAGNOSIS: Easily distinguished from the Lengths of Appendage Segments in Paratype other species by having two pairs of spikes Female Acuclavella merickeli on the scutum, on areas 1 and 2. (Measurements are in millimeters.) DESCRIPTION OF FEMALE HOLOTYPE: Body Legs (fig. 29) highly arched, convex dorsally, broader posteriorly, longer than wide (length/ Segment Palpus 1 2 3 4 width = 1.35). Nearly all of body heavily Trochanter 0.57 0.65 0.72 0.65 0.68 sclerotized, dark brown, with densely scat- Femur 0.96 1.90 3.00 1.95 2.73 tered, small, black warts; body without pi- Patella 0.78 1.04 1.04 0.86 1.04 losity. Total length 5.04 mm; carapace length Tibia 0.83 1.38 2.18 1.43 1.95 in midline 1.30 mm, greatest carapace width Metatarsus - 2.16 3.64 2.44 3.67 2.86 mm; length of scutum in midline 3.38 Tarsus 0.73 2.47 4.32 2.66 3.51 mm, greatest width of scutum 3.74 mm. Eye tubercle at edge of carapace, oval at base, prolonged into sharp, conical, forward- tilted spine standing 1.09 mm above cara- NOTES: As with the other species, this one pace. Eyes small, dark brown, on rounded is evidently quite cryptic, with only a few basal part of tubercle. Features of carapace specimens having been taken until the type as in A. cosmetoides. collection was made in fall of 1984. Scutum ofopisthosoma rounded, narrower Regarding this collection, Mr. Merickel anteriorly, posterior margin broadly squared wrote me: "All these were collected next to offwhen seen dorsally. All areas with paired a small creek, on bare, moist ground. Two of tubercles; on area 1, large spikes situated at these, probably a pair, were discovered quite anterior edge of scute on either side of mid- accidentally while I was washing dishes and line. On area 2, spikes about 25 percent larg- I suspect they were disturbed out of hiding. er, about twice as far apart. Areas 3 and 4 Thinking that they might be nocturnal, I re- have small, raised, warty knobs paired near turned to the same place that night and sure the midline, with three noticeably enlarged enough was able to pick up six more walking clusters of warts lateral to them in row on along the [stream] bank. A number of other each side; on area 5 only clusters homologous situations were examined, including moist to spikes visible; free tergites without tuber- logs and mossy areas, but to no avail." The cles, warts irregularly distributed. previous specimens from Meadow Creek Anal region, spiracles, and genital oper- came from Berlese samples oflitter taken near culum as in A. cosmetoides. Prosomal ster- the same stream. num 0.13 mm long, 0.34 mm wide, brown, The specimen from Lewis Co., Washing- without setae. Labium weakly sclerotized, ton, was found under wet rocks near a wa- about as long as wide, without setae. Endites terfall. While very distant from the type lo- as in A. cosmetoides. cality, this specimen is very similar in all Chelicerae light brown, surface smooth, respects to the material from Idaho, and so shining. Basal article 1.19 mm long, 0.57 mm is included here. wide. Palp with dimensions given in table 8. Coxa yellowish, ventrally with two seta-bearing tubercles. Trochanter light brown with Acuclavella quattuor, new species ventral row offour seta-bearing tubercles. Fe- Figure 29 mur white. Patella very light brown, bearing small setae and microtrichia distally. Tibia TYPE: Female holotype (AMNH) from 10 white, with denser setae and microtrichia. miles east of Slate Creek, Idaho Co., Idaho Tarsus darkened distally, claw rudiment ab- (May 10-21, 1983; F. W. Merickel and J. B. sent. Johnson). Structure and color of legs as in A. cos- ETYMOLOGY: The species name is an ad- metoides; lengths of segments given in ta- jective referring to the four prominent spikes ble 8. on the dorsum of the abdomen. Ovipositor as in A. cosmetoides. 1986 SHEAR: ISCHYROPSALIDOIDEA 23

t/33 i'34 3t5 363 37

FIGS. 33-37. Genitalia of Acuclavella species. 33, 34. A. cosmetoides. 33. Penis, lateral view. 34. Glans penis, dorsal view. 35. Glans penis ofA. shoshone, lateral view. 36. Glans penis ofA. merickeli, dorsal view. 37. Ovipositor ofA. cosmetoides, dorsal view.

DISTRIBUTION: Known only from the type Crosbycus goodnighti, from Fountain Cave, locality. Augusta Co., Virginia (no date or collector's NOTES: The holotype specimen was taken name given). Unfortunately, the vial con- from a cantharidin trap. taining the types could not be located at the Senckenberg Museum. As Rambla (1968) Crosbycus Roewer, 1914 pointed out, the illustrations and the descrip- Crosbycus Roewer, 1914, p. 168. See additional TABLE 8 citations below. Lengths of Appendage Segments in Holotype TYPE SPECIES: Nemastoma dasycnemum Female Acuclavella quattuor Crosby (1911). (Measurements are in millimeters.) DIAGNOSIS, ETC.: The only known species Legs is discussed below. NOTES: Giltay (1932), Roewer (195 1), and Segment Palpus 1 2 3 4 Loksa (1962) all described additional Cros- Trochanter 0.57 0.60 0.73 0.58 0.73 bycus species from Europe. Rambla (1968) Femur 0.94 2.18 3.25 2.21 2.73 and Gruber and Martens (1968) demonstrat- Patella 0.78 0.86 1.09 1.04 1.00 ed that all these names were based onjuvenile Tibia 0.89 1.56 2.13 1.43 1.98 nemastomatids. An additional American Metatarsus - 2.34 3.54 2.60 3.80 Tarsus 0.70 2.65 4.16 2.99 3.77 species was also described by Roewer in 1951: 24 AMERICAN MUSEUM NOVITATES NO. 2844 tion leave little doubt that this, too, is a denticles. Metapeltidium fully incorporated juvenile nemastomatid. The only nemasto- into carapace, marked by distinct fold of cu- matid in eastern North America is Nemas- ticle; without sensory cones, with densely toma bimaculatum, a European species that scattered black denticles. Scutum nearly obviously has been imported and has estab- square, anterior and posterior margins con- lished itself. (I found a large number of spec- vex, lateral margins nearly parallel, with five imens in the collection of the AMNH that areas (evidently incorporating five tergites). had been taken in Toronto in 1945 and 1946 First three areas not distinct, marked by by W. Ivie and T. B. Kurata; the species was densely and evenly scattered black denticles; reported from Montreal by LeSage, 1978.) In areas 4 and 5 oval, 4-5 times wide than long, western North America and Mexico are the densely set with black denticles. (In some very distinctive ortholasmatine nema- specimens, area 5 partially separated by white stomatids (Shear and Gruber, 1983). Foun- membrane from remainder of scutum.) Free tain Cave is well known to biospeleologists tergites and anal operculum with same dense (Holsinger, 1964, 1975; personal commun.), scattering of denticles. and no opilionids answering the description Sclerotized parts of ventral surface with of C. goodnighti have been taken there or in scattered denticles and densely set, thin, tri- the vicinity. dentate microscales in irregular rows (fig. 17). The specimens Roewer used must some- Genital operculum very broad, set off by how have gotten mislabeled. Clearly Cros- complete, straight suture, with scattered se- bycus goodnighti, whatever it might be, or tae. Sternum and labium not sclerotized. All from whence it might have come, is not a coxae with large, prominent endites, as in Crosbycus. figure 39. Labrum with long, distinctive median projection. Crosbycus dasycnemus (Crosby) Chelicerae brown, not enlarged. Basal ar- Figures 6, 12, 17, 38-41 ticle without gland. Nemastoma dasycnemum Crosby, 1911, p. 20. Palpus (fig. 40) with dimensions as in table Comstock, 1912, p. 78. 9, extremely long and thin, nearly as long as Crosbycus dasycnemum, Roewer, 1914, p. 168; first and third legs. Trochanter, femur, and 1919, p. 160. patella with few ordinary setae, as in figure Crosbycus dasycnemus, Roewer, 1923, p. 678; 40; tibia and tarsus with plumose setae (fig. 1951, p. 151. Crosby, Wolfe, and Bishop, 1928, 6), becoming more dense towards tip of tar- p. 1074. Comstock, 1940, p. 78. Bishop, 1949, sus, claw rudiment completely absent. p. 172. LeviandLevi, 1952, p. 165. Edgar, 1966, Dimensions of legs as in table 9. Femora, p. 356. Rambla, 1968, p. 70. Gruber, 1970, p. patellae, and tibiae with complex ornamen- 129. Suzuki and Kunita, 1972, p. 89. Suzuki et tation consisting of large, socketed setae, un- al., 1977, p. 126. Martens, 1976, p. 66; 1978, socketed spikes projecting perpendicular to p. 186. segment axis, and unsocketed curly hairs, TYPE: Female holotype (Cornell Colection, procumbent distad, seemingly densely inter- AMNH) from Columbia, Boone Co., Mis- woven (fig. 12). Tarsi consisting of single ba- souri, examined. sitarsal segment, two distitarsal segments on DESCRIPTION: Female (fig. 38) from Dune first legs, three on legs 2-4, claws untoothed. Acres, Porter Co., Indiana. General color dark Ovipositor (fig. 41) very short, two to three brown, with unsclerotized areas white. Legs times longer than wide, apically bifurcate, yellowish. Total length, 0.89 mm. Carapace each furca with two dorsal, two ventral, and 0.17 mm long, 0.41 mm wide; abdominal one apical seta. Seminal receptacles not de- scutum 0.49 mm wide. Carapace arcuate, tected, probably absent. slightly indented in midline, with clusters Males unknown, probably do not exist. of small, acute to anvil-shaped, black den- DISTRIBUTION: NEW HAMPSHIRE: Sul- ticles generally distributed anteriorly in two livan Co.: Meriden, June 28, 1967 (W. B. groups on either side eye tubercle. Eye tu- Muchmore), 3 ff (VM). NEW YORK: Mon- bercle 0.09 mm wide, rounded, at margin of roe Co.: Powder Mills Park, May 2, 1965, carapace, with posteriomedian group ofblack Apr. 16, 1966 (W. B. Muchmore), 2 ff(VM). 1986 SHEAR: ISCHYROPSALIDOIDEA 25

FIGS. 38, 39. Scanning electron micrographs offemale Crosbycus dasycnemus. 38. Body, dorsal view (x 82). 39. Anterior ventral surface, ventral view (x 160).

Steuben Co.: Stoney Brook State Park, Aug. biaville, July 25, 1963 (W. Suter, R. Graves), 1, 1957, 1 f (AMNH). Suffolk Co.: Montauk 2 ff (FMNH). Marquette Co.: 15 mi S Ish- Point, from sphagnum, June 25, 1927 (vial penning (Green Hills Area), Aug. 10, 1956 empty but containing a Crosby determina- (H. Dybas), 2 ff(FMNH). INDIANA: Porter tion label; AMNH). Tompkins Co.: Con- Co.: Dune Acres, Apr. 12, 1949 (H. Dybas), necticut Hill, Sept. 8, 1935 (C. R. Crosby), 1 2 ff(FMNH); 1 mi SW McCool, Oct. 4, 1952 f (AMNH); Freeville, Oct. 12, 1924, 1 f (H. Dybas), 2 ff(FMNH). ILLINOIS: Cham- (AMNH). Wayne Co.: Zurich Bog, Apr. 26, paign Co.: Urbana, Oct. 4, 1925 (V. G. 1970 (W. B. Muchmore), 2 ff (VM). Yates Smith), 1 f(AMNH). WEST VIRGINIA: Po- Co.: Keuka Lake, July 3, 1926, 1 f; Sept. 24, cahontas Co.: Hills Creek Falls Scenic Area, 1920, 1 f (AMNH). PENNSYLVANIA: Ly- coming Co.: near Williamsport, Sept. 18, 1969 TABLE 9 (W. B. Muchmore), 1 f (VM). WISCONSIN: Lengths of Appendage Segments in Female Cros- Vernon Co.: 5 mi N Viola, mixed hardwood bycus dasycnemus from Dune Acres, Porter Co., litter, Sept. 2, 1954 (H. W. Levi), 1 f(MCZ). Indiana MINNESOTA: Hennepin Co.: Lake Min- (Measurements are in millimeters.) netonka, Aug. 28, 1924 (C. Fletcher) (two Legs empty vials with Crosby determination labels, AMNH). MICHIGAN: Alger Co.: Shel- Segment Palpus 1 2 3 4 ter Bay, 6 mi W Au Train, July 16, 1953 (H. Trochanter 0.26 - - - - Dybas), f(FMNH). Alpena Co.: T30N, R7E, Femur 0.50 0.52 0.68 0.47 0.64 Sec. 5, beech-maple forest, July 16-20, 1969 Patella 0.41 0.13 0.14 0.13 0.14 (G. V. Manley), 2 ff (MSU). Leelanau Co.: Tibia 0.43 0.39 0.71 0.43 0.64 Metatarsus - 0.37 0.59 0.39 0.52 near Empire, June 7, 1946 (C. Goodnight), 1 f (AMNH). LaPeer Co.: Bog 4 mi E Colum- Tarsus 0.27 0.38 0.63 0.43 0.50 26 AMERICAN MUSEUM NOVITATES NO. 2844

40 41 FIGS. 40, 41. Crosbycus dasycnemus female. 40. Left palpus, mesal view. 41. Ovipositor, ventral view. Note lack of seminal receptacles. Drawing courtesy of Jurgen Gruber. above Middle Falls, June 19, 1971 (W. Shear), house Mt., 4 mi NW Walhalla, Apr. 4, 1929 2 ff (WAS). VIRGINIA: Shenandoah Na- (C. R. Crosby), 1 f(AMNH). GEORGIA: Ra- tional Park: Big Meadows (3500 ft), July 2, bun Co.: Chatooga River bank, 1.5 mi NE 1963 (W. B. Muchmore), 2 ff(VM). Giles Co.: jct GA 28 and Jeep Road, June 1, 1975 (L. Mountain Lake, July 2 and 12, 1970 (W. B. Driskill), 2 ff (WAS). Muchmore), 5 ff (VM). NORTH CAROLI- NOTES: As Bishop (1949) pointed out, less NA: Buncombe Co.: Craggie Pinnacle on Blue well-sclerotized specimens may appear to Ridge Parkway (5640 ft), July 14, 1977 (J. have tergites 4 and 5 not incorporated in the D. Pittillo), 2 ff(WAS). Macon Co.: 5 mi NE abdominal scutum. From an examination of Highlands, ericaceous litter, Oct. 25, 1969 a series ofspecimens it seems likely that with (W. Shear), 2 ff(WAS); Wayah Bald Summit age (within the adult instar) additional scler- (5300 ft) June 21, 1963 (W. B. Muchmore), otization or additional deposition ofpigment 1 f (VM). Watauga Co.: Grandfather Moun- leads to the typical scutum parvum. In all the tain, Fork Creek (4080 ft), Rhododendron lit- specimens examined for this study, as well ter, May 16, 1977 (J. D. Pittillo) 1 f(WAS). as those in the Vienna Museum studied by SOUTH CAROLINA: Oconee Co.: Stump- Gruber, no males were found. Using light 1986 SHEAR: ISCHYROPSALIDOIDEA 27 microscopy at high magnifications, no sem- 1974). For a further discussion of the bio- inal receptacles could be detected in the ovi- geographical situation, see Suzuki et al. positors. This strongly suggests that the (1977), and Shear and Gruber (1983). species is parthenogenetic, but relatively few specimens are available, and males might be LITERATURE CITED highly seasonal. Likewise, there are no juvenile specimens in collections, suggesting Banks, Nathan 1894. The Nemastomatidae and Trogulidae that a different habitat might be occupied ear- ofthe United States. Psyche, vol. 7, pp. ly in life. Prey probably consists ofeven tinier 11-12. mites, collembolans, and myriapods. 1901. Synopses of North American inverte- In North America, C. dasycnemus is found brates. XVI. The Phalangida. Amer. in boreal regions, high mountains, or in cool Nat., vol. 35, pp. 669-679. microhabitats. In the midwestern states, col- Bishop, Sherman C. lection labels suggest an association with 1949. The Phalangida (Opiliones) of New mosses, particularly sphagnum. From West York, with special reference to the Ed- Virginia south, the species occurs in the mund Niles Huyck Preserve, Rensse- mountains, at moderate to higher elevations, laerville, N.Y. Proc. Rochester Acad. Sci., vol. 9, pp. 159-235, figs. 1-110. but evidently not in coniferous forests (col- Cokendolpher, James C. lection labels mention beech and maple), and 1984. A new species of Sabacon Simon from often near streams. Usually taken in Berlese Oregon (Arachnida: Opiliones: Saba- samples, this animal would be very difficult conidae). Can. J. Zool., vol. 62, pp. 989- to spot during hand sorting, especially if it 991, figs. 1-7. were slow-moving or tonically immobile. In Comstock, John Henry no case is there a record of how much litter 1913 (1912). The spider book. Garden City, was sorted, but only two collections con- N.Y., 721 pp., 770 figs. tained more than one or two individuals. 1940. The spider book, revised and edited by Specimens may be overlooked in the field or Willis J. Gertsch. Ithaca, N.Y., 730 pp., 771 figs. misfiled in museums because oftheir resem- Crosby, Cyrus R. blance to mites, just as Opilioacarus (an opil- 1911. A new species of phalangida from Mis- ionid-like mite) can often be found in mu- souri. Can. Ent., vol. 43, pp. 20-22, fig. seums' opilionid collections. 1. The wide distribution of this species in- Crosby, Cyrus R., A. Wolf, and Sherman C. Bish- cludes Japan (Suzuki et al., 1977), as well as op the North American localities listed above. 1928. Orders Araneae and Opiliones. In M. The Japanese specimens were compared by D. Leonard (ed.), A list ofthe insects of Suzuki with North American ones and are New York. Ithaca, N.Y., 1076 pp. virtually identical; likewise there is no de- Dresco, E. 1970. Recherches sur la variabilite et la phy- tectable variation over the range ofthe species logenie chez les opilions du genre Is- in North America. Other opilionid species chyropsalis C. L. Koch (Fam. Ischyrop- and genera have a similar discontinuous dis- salidae), avec creation de la famille tribution: Caddo agilis, C. pepperella, and nouvelle des Sabaconidae. Bull. Mus. Acropsopilio boopis (Caddidae), Dendrolas- Hist. Nat. Paris, ser. 2, vol.41, pp. 1200- ma (Nemastomatidae), and Paranonychus 1213, figs. 1-8. brunneus (= Mustsunonychusfuscus Suzuki; Edgar, Arlan E. Paranonychidae). The connection seems to 1966. Phalangida of the Great Lakes region. be between eastern North America and Amer. Midl. Nat., vol. 75, pp. 347-366, southern and western North America figs. 1-17. Japan, Giltay, Louis and northern Japan. For three ofthe disjunct 1932. Arachnides recueillis par M. D'Orchy- species, males are not known (C. pepperella, mont au cours de ses voyages aux Bal- A. boopis, C. dasycnemus), and for a fourth kans et en Asie Mineure. Bull. Mus. Roy. (C. agilis), single males have been found just Hist. Nat. Belg., vol. 8, pp. 1-40, figs. once in Japan (Suzuki and Tsurusaki, 1983) 1-22. and just once in North America (Gruber, Goodnight, Clarence J., and Marie L. Goodnight 28 AMERICAN MUSEUM NOVITATES NO. 2844

1942. New Amerian Phalangida. Amer. Mus. Acad. Sci. Arts. Lett., vol. 41, pp. 163- Novitates, no. 1164, pp. 1-4, figs. 1-6. 167, figs. 1-9. 1945. A representative of the Ischropsalidae Ljovushkin, S. I. [sic] Simon (Phalangida) from Mexico. 1971. Biospeleologica Sovietica XLIII. Re- Trans. Conn. Acad. Arts Sci., vol. 36, port ofthe first finding ofthe American pp. 249-251, figs. 1, 2. genus Taracus Simon, 1879 (Opiliones, Gruber, Jiirgen Ischyropsalididae) in Asia with a de- 1969. Uber Stridulationsorgane bei einem Is- scription of T. birsteini sp. n., from a chyropsaliden: Ceratolasma tricantha cave of the southern part of Primorsky Goodnight und Goodnight (Opiliones, Krai. Byul. Mosk. Obshch. Ispyt. Pr. ir., Arachnida). Anz. Akad. Wiss. Wien, Biol., vol. 76, pp. 127-133, figs. 1-5, Math.-Naturwiss. Ki., 1968, pp. 249- map 1. 255, figs. 1-4. Loksa, I. 1970. Die "Nemastoma-"Arten Nordameri- 1962. Beitrage zur kenntnis der Weber- kas (Ischyropsalididae, Opiliones, knechte. Fauna Ungarns I. Zool. Anz., Arachnida). Ann. Naturhistor. Mus. vol. 68, pp. 265-269. Wien, vol. 74, pp. 129-144, figs. 1-24. Martens, Jochen 1974. Bemerkungen zur Morphologie und sys- 1969. Die Abgrenzung von Biospezies auf tematischen Stellung von Caddo, Ac- biologisch-ethologischer und morpho- ropsopilio und verwandten Formen logischer Grundlage am Beispiel de (Opiliones, Arachnida). Ibid., vol. 78, Gattung Ischyropsalis C. L. Koch 1839 pp. 237-259, figs. 1-7. (Opiliones, Ischyropsalididae). Zool. 1978. Redescription ofCeratolasma tricantha Jahrb. Syst., vol. 96, pp. 133-264, figs. Goodnight and Goodnight, with notes 1-68. on the Family Ischyropsalidae (Opil- 1972. Opiliones aus dem Nepal-Himalaya. I. iones, Palpatores). Jour. Arachnol., vol. Das Genus Sabacon Simon (Arachnida: 6, pp. 105-124, figs. 1-29, map 1. Ischyropsalididae). Senckenbergiana Gruber, Jiirgen, and Jochen Martens Biol., vol. 53, pp. 307-323. 1968. Morphologie, Systematik und Okologie 1976. Genitalmorphologie, System und Phy- der Gattung Nemastoma C. L. Koch (s. logenie der Weberknechte (Arachnida: str.) (Opiliones, Nemastomatidae). Opiliones). Ent. Germanica, vol. 3, pp. Senckenbergiana Biol., vol. 49, pp. 137- 51-68, figs. 1-46. 172, figs. 1-74. 1978. Weberknechte, Opiliones. In Die Tier- Hansen, Henrik J., and William S0rensen welt Deutschlands, vol. 64, Jena, Gus- 1904. On two orders ofArachnida. Cambridge tav Fischer Verlag, 464 pp. Univ. Press, 178 pp. 1981. Versuch eines phylogenetischen Sys- Holsinger, John R. tems der Opiliones. Proc. 8th Internat. 1964. The biology ofVirginia caves. In H. H. Congr. Arachnol., pp. 355-360. Douglas, Caves of Virginia. Falls 1983. Europaische Arten der Gattung Saba- Church, Va., 761 pp. con Simon 1879 (Arachnida: Opiliones: 1975. Descriptions ofVirginia caves. Virginia Sabaconidae). Senckenbergiana Biol., Div. Min. Res. Bull., vol. 85, pp. 1-450. vol. 63, pp. 265-296, figs. 1-56, maps Juberthie, Christian, Andre Lopez, and Lysiane 1,2. Juberthie-Jupeau Martens, Jochen, Ulrich Hoheisel, and Marianne 1981. Etude ultrastructurale des sensilles tho- Gotze raciques dorsales et paramedianes chez 1981. Vergleichende Anatomie der Legeroh- Sabacon paradoxum Simon (Palpa- ren der Opiliones als Beitrage zur Phy- tores, Sabaconidae). Atti Soc. Tosc. Sci. logenie der Ordnung (Arachnida). Zool. Nat., Mem., ser. B, vol. 88 (suppl.), pp. Jahrb. Anat., vol. 105, pp. 13-76, figs. 27-33, fig. 1. 1-44. LeSage, Laurent Martens, Jochen, and Wolfgang Schawaller 1978. Premi'ere mention de Nemastoma bi- 1977. Die Cheliceren-Driisen der Weber- maculatum (Fab.) pour l'Amerique du knechte nach rastoptischen und lichtop- Nord (Opiliones: Nemastomatidae). tischen Befunden (Arachnida: Opil- Nat. Can., vol. 104, p. 485. iones). Zoomorphol., vol. 86, pp. 223- Levi, Lorna R., and Herbert W. Levi 270, figs. 1-11. 1952. Preliminary list of harvestmen of Wis- Martens, Jochen, and Seisho Suzuki consin with a key to genera. Wisconsin 1966. Zur systematischen Stellung ostasia- 1986 SHEAR: ISCHYROPSALIDOIDEA 29

tischer Ischyropsalis-arten (Arachnida, Tomicomerus in America (Opiliones, Opiliones, Ischyropsalididae). Annot. Troguloidea, Ischyropsalidae). J. Ar- Zool. Japonenses, vol. 39, pp. 215-221, achnol., vol.3, pp. 5-29, figs. 1-37, maps figs. 1-7. 1, 2. Platnick, Norman I. Shear, William A., and Jiirgen Gruber 1977. Monotypy and the origin ofhigher taxa: 1983. The opilionid subfamily Ortholasma- a reply to E. 0. Wiley. Syst. Zool., vol. tinae (Opiliones, Troguloidea, Nema- 26, pp. 355-357. stomatidae). Amer. Mus. Novitates, no. Rambla, Maria 2757, pp. 1-65, figs. 1-210, maps 1-3. 1968. Sobre el genero Crosbycus Roewer 1914 Simon, Eugene (Opiliones, fam. Nemastomatidae). Pub. 1879. Les Arachnies de France, vol.7, pp.266- Inst. Biol. Apl., vol. 44, pp. 65-79, figs. 332, Paris. 1-8. Suzuki, Seisho Roewer, Carl-Friedrich 1974. The Japanese species of the genus Sa- 1914. Die Familien der Ischyropsalidae und bacon (Arachnida, Opiliones, Ischyrop- Nemastomatidae der Opiliones Palpa- salididae). J. Sci. Hiroshima Univ., ser. tores. Arch. Naturg., vol. 80A, pp. 99- B, div. 1 (Zool.), vol. 25, pp. 83-108, 169. figs. 1-13. 1919. Uber Nemastomatiden und ihre Ver- Suzuki, Seisho, and Toshiaki Kunita breitung. Ibid., vol. 83A, pp. 140-160, 1972. Opiliones ofEhime Pref., Shikoku. Ehi- figs. 1-3. me-no-Seibutsu, vol. 8, pp. 89-94, figs. 1923. Die Weberknechte der Erde. Jena, 1116 1-6 [in Japanese]. I?.P- Suzuki, Seisho, Keiji Tomijima, Shizuko Yano, 1950. Uber Ischyropsalididae und Troguli- and Tenji Tsurusaki dae. Weitere Weberknechte XV. Sen- 1977. Discontinuous distributions in relict ckenbergiana, vol. 31, pp. 11-56, plates harvestmen (Opiliones, Arachnida). 1-10. Acta Arachnol., vol. 27, pp. 121-138, 1951. Uber Nemastomatiden. Weitere We- figs. 1-5 [in Japanese]. berknechte XVI. Ibid., vol. 32, pp. 95- Suzuki, Seisho, and Nobuo Tsurusaki 153, plates 1-9. 1983. Opilionid fauna ofHokkaido and its ad- Shear, William A. jacent areas. J. Fac. Sci. Hokkaido Univ., 1975. The opilionid genera Sabacon and ser. 4, Zool., vol. 23, pp. 195-243.

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