1999. The Journal of Arachnology 27:44±52

ON THE PHYLOGENETIC RELATIONSHIPS OF SISICOTTUS HIBERNUS (ARANEAE, , ERIGONINAE)

Jeremy Zujko-Miller: Department of Biological Sciences, The George Washington University, Washington, D.C. 20052, and Department of Entomology, National Museum of Natural History, NHB-105, Smithsonian Institution, Washington, D.C. 20560

ABSTRACT. Carorita hiberna NEW COMBINATION, a species with many putative autapomorphies known from one sex and few specimens, is transferred from Sisicottus. This transfer is based on a modi®ed version of a cladistic analysis of erigonine relationships by G. Hormiga which incorporated 43 taxa scored for 73 characters. The modi®ed analysis features 46 taxa scored for 74 characters. The resulting cladogram placed C. hiberna sister to C. limnaea, the type species of Carorita. It is concluded that C. hiberna is better placed in Carorita than in either a new monotypic or in Sisicottus. Carorita hiberna is redescribed and the monophyly of Carorita as currently circumscribed is discussed.

Carorita hiberna (Barrows 1945) NEW tion using a temporary slide mount (Codding- COMBINATION (Linyphiidae, Erigoninae) ton 1983). I made sketches using a camera was inexplicably described as a member of the lucida ®tted to a Leica DMRM compound mi- genus Sisicottus Bishop & Crosby 1938. Ca- croscope at 400ϫ. Further observations were rorita hiberna is a very unusual erigonine spe- made using a Leica MZ APO dissecting mi- cies that shares none of the synapomorphies croscope. Museum acronyms for specimen de- that unite Sisicottus (Zujko-Miller 1999). Ca- positories appear in the acknowledgments. rorita hiberna is known only from three male specimens from the Great Smoky Mountains CLADISTIC ANALYSIS National Park, North Carolina, USA. After re- The cladistic analysis by Hormiga (in press) vising Sisicottus (Zujko-Miller 1999), I was is the most rigorous hypothesis of erigonine left with three alternatives as to the fate of C. hiberna: I could keep it in Sisicottus, which relationships to date and is the logical starting would leave Sisicottus polyphyletic; I could point for questions of relationships within the erect a new monotypic genus for it; or I could Erigoninae. The original analysis incorporates transfer it to the genus which contains its clos- 43 terminal taxa, including 31 erigonine gen- est relatives. Carorita hiberna features many era, scored for 73 characters. My modi®ed apparently apomorphic character states in the version of Hormiga's analysis incorporates form of the male palpus, and it was not ob- three additional taxa, one new character, and vious to me what genus contained its closest one recoded character. relatives. I chose to seek the closest relatives Carorita hiberna, C. limnaea, and Sisicot- of C. hiberna using phylogenetic methods. By tus montanus (Emerton 1882) were added to placing C. hiberna in a phylogenetic context, Hormiga's (in press) matrix. Carorita limnaea I was able to formulate a testable phylogenetic was included because it appears to share some hypothesis. My cladistic analysis identi®ed potentially synapomorphic character states Carorita limnaea (Crosby & Bishop 1927), with C. hiberna including a looped sperm duct the type species of Carorita Duffey & Merrett in the tegulum, a tuberculate radical tailpiece, 1963, as the sister taxon of C. hiberna. With and a suprategulum separated from the tegul- the transfer of C. hibernus, Carorita currently um by a membranous region so that it appears contains three species. to form a distinct sclerite. Sisicottus was in- METHODS cluded to test the implicit phylogenetic hy- I cleared specimens in methyl salicylate pothesis of Barrows (1945) that C. hiberna (Holm 1979) and positioned them for illustra- plus Sisicottus form a monophyletic group.

44 ZUJKO-MILLERÐSISICOTTUS HIBERNUS 45

Several character states remain unknown for this character. The junction between the for C. hiberna because females are unknown tegulum and the suprategulum (character 12) and males are rare and not available for irre- was recoded. Character 12 documents the versible methods of examination. The male membranous hinge between the tegulum and cephalothorax was not examined using scan- the suprategulum in Stemonyphantes Menge ning electron microscopy to search for cutic- 1886 (van Helsdingen 1968; Hormiga 1994). ular pores in the clypeal region (character 50) In the context of Hormiga's analysis, this or to examine details of the stridulatory striae character state is autapomorphic and character on the chelicerae (character 56). No abdomens 12 is not phylogenetically informative. I have were digested for examination of the tracheal added a third state to character 12 and it is system (characters 51, 52). Carorita hiberna now coded as follows: Suprategulum: 0 ϭ was coded as follows: 0001310110 continuous with tegulum; 1 ϭ articulated; 2 1210110101 1201000101 3????????? ϭ separate from tegulum (Figs. 7, 9). In Ca- 000000000? ??001?0??1 00011?0??1 1??1 rorita limnaea, C. hiberna, Asthenargus pa- Carorita limnaea was coded as follows: ganus (Simon 1884), Gongyliellum vivum (O. 0001310110 1210110101 1501000101 Pickard-Cambridge 1875) and Erigone psy- 300±000100 0000000000 0±00120111 chrophila Thorell 1871, there is a membra- 0011100101 1??1. Carorita limnaea was giv- nous division between the sclerotized parts of en a unique character state for the shape of the tegulum and the suprategulum so that the the radical tail piece (character 22). In C. lim- suprategulum appears to be a distinct sclerite naea, the tailpiece extends both dorsally and rather than a more heavily sclerotized distal ventrally from its origin distal to the origin of portion of the tegulum. This is the new char- the embolus. Coding of C. limnaea was based acter state coded as 2. In most other liny- on examination of the following specimens: phiids, the tegulum and the suprategulum are UNITED STATES: Maine: Piscataquis joined by a region of continuous sclerotization County, 2.3 km ESE of Soubunge Mtn., dense and only part of the junction between the te- spruce-®r forest, Line I, Stn, 1, T4 R11, gulum and the suprategulum is membranous. WELS, 1 June 1978, pitfall collection, 1(, In Stemonyphantes, the junction between the (D.T. Jennings, M.W. Houseweart, USNM); tegulum and the suprategulum is a wide ¯ex- New York: McLean, mud pond, 42Њ32ЈN, ible hinge (Hormiga 1994, ®g. 2c). Also, the 76Њ18ЈW, 30 May 1921, 8(17&, (C.R. Cros- tegular-suprategular junction is unusual in Ste- by, AMNH). monyphantes because it is on the ventral face Sisicottus montanus was coded as in Zujko- rather than the mesal face of the palpal bulb. Miller (1999). Character 74 was scored with Analysis.ÐI used PAUP version 3.1 a zero. Coding of S. montanus was based on (Swofford 1993), Hennig86 version 1.5 (Far- examination of the following specimens: ris 1988) and NONA version 1.6 (Goloboff UNITED STATES: Massachusetts: Berk- 1993) to search the data (46 taxa, 74 charac- shire County, Mt. Greylock, 3400 feet, decid- ters) for the most parsimonious topology. In uous litter, 15 October 1990, 2(1&, (R.L. Ed- PAUP, I ran a heuristic search with 100 rep- wards, USNM). licates of random taxon addition subjected to Hormiga's (in press) analysis was modi®ed tree bisection-reconnection branch swapping. by the addition of one new character and the In Hennig86, I used the ``mh*,bb*'' search recoding of an existing character. Both char- strategy. In NONA, I ran a search under the acters pertain to structures that are synapo- ``ambϭ'' setting (modi®ed rule 3; see Cod- morphies of the Linyphiidae (the suprategul- dington & Scharff 1994; Zujko-Miller 1999) um and the linyphiid radix) so character states with the ``mult*'' random taxon addition al- for linyphiid taxa other than C. limnaea, C. gorithm for 100 replicates followed by the hiberna, and S. montanus were determined us- ``max*'' branch-swapping algorithm. I used ing Hormiga (1994, in press). Character 74 is MacClade (Maddison & Maddison 1992) to the texture of the radical tailpiece which is analyze character optimization. tuberculate in C. limnaea and C. hiberna.In Successive character weighting (Farris all other taxa with a radical tailpiece, this 1969; Carpenter 1988) by the maximum value sclerite is more or less smooth. Taxa without of the rescaled consistency index was per- a radical tailpiece were coded as inapplicable formed in PAUP with the base weight set to 46 THE JOURNAL OF ARACHNOLOGY

Figures 1±6.ÐPalpus of male Carorita hiberna from Thomas Ridge, North Carolina. 1. Embolic di- vision, dorsal view; 2. Ventral view with embolic division removed; 3. Ventral view; 4. Mesal view; 5. Ectal view; 6. Palpal tibia, dorsal view. Abbreviations: DRP, dorsal radical process; DSA, distal suprate- gular apophysis; E, embolus; EM, embolic membrane; ETP, ectal tibial process; F, fundus; P, paracymbium; PT, protegulum; PTA, palpal tibial apophysis; R, radix; SD, sperm duct; SPT, suprategulum; ST, subtegu- lum; T, tegulum; TP, radical tail piece.

1000. Trees found by Hennig86 and NONA unique trees were then ®ltered to exclude po- were imported into PAUP. NONA trees were lytomous trees when more highly resolved saved using the ``ksv*'' command. PAUP will compatible trees were found (Coddington & arbitrarily resolve polytomies in trees saved Scharff 1996). This set of trees was re-weight- using the ``sv'' command in NONA. The so- ed and the data re-analyzed in PAUP. lution set from all three programs (PAUP, Results.ÐPAUP, Hennig86, and NONA Hennig86, and NONA) was combined. Dupli- and all found multiple trees of 236 steps (cal- cate trees were eliminated. The remaining culated after excluding uninformative charac- ZUJKO-MILLERÐSISICOTTUS HIBERNUS 47

Figures 7±9.ÐGenitalia of Carorita species. 7. Male palpus of C. hiberna from Thomas Ridge, North Carolina, mesoventral view with embolic division removed; 8, 9. Carorita limnaea from McLean New York; 8. Cleared epigynum, ventral view; 9. Male palpus, ventromesal view. Abbreviations: CD, copu- latory duct; CDC, copulatory duct capsule; DP, dorsal plate of epigynum; DSA, distal suprategular apoph- ysis; E, embolus; EM, embolic membrane; FD, fertilization duct; P, paracymbium; PT, protegulum; R, radix; S, spermatheca; SPT, suprategulum; ST, subtegulum; T, tegulum; TP, radical tail piece; VP, ventral plate of epigynum. ters) with a consistency index (CI) of 0.377 agreement among the six trees is found in two and a retention index (RI) of 0.673. PAUP places: the relationships among the linyphi- found 63 trees, Hennig86 found 33 trees, and ines, micronetines and all other linyphiids and NONA found 45 trees. A total of 78 unique the relationships among Drepanotylus Holm most parsimonious trees were found. Of these, 1945, Sciastes Bishop & Crosby 1938 and the 24 trees were more highly resolved but oth- ``distal erigonines'' clade. The ``distal erigon- erwise compatible with other most parsimo- ines'' clade is fully resolved and identical in nious trees. All trees place Carorita limnaea all six trees. The ``distal erigoinines'' clade sister to Carorita hiberna. A strict consensus features a monophyletic Carorita clade sister of all most parsimonious trees has a trichot- to Gongylidiellum vivum. Sisicottus remains in omy composed of the Carorita clade, Asthen- the position reported by Zujko-Miller (1999), argus paganus and Gongylidiellum vivum. sister to Oedothorax gibosus (Blackwall This clade is part of an 11-tomy composed of 1841). various erigonine terminals and clades. Suc- Implications.ÐThe two species which for- cessive character weighting stabilizes on six merly composed Carorita have traditionally trees. All six trees are 236 steps long under been united based on their chaetotaxy (espe- equal weights and were among the original set cially the presence of a prolateral macroseta of 78 trees. Except for the additional taxa, this on tibia I), the large paracymbium, the form set of six trees is identical to the result found of the suprategular apophysis, overall palpal in Hormiga's (in press) original analysis. Dis- conformation, the general form of the palpal 48 THE JOURNAL OF ARACHNOLOGY tibia, the relatively long tarsi, and proportional through the tegulum. However, when I at- characteristics of the eyes (Duffey 1971; Mil- tempted to code this as a cladistic character, I lidge 1977). Millidge (1977) considered C. found a spectrum of conditions rather than a limnaea and C. paludosa Duffey 1971 to be small number of discrete character states. members of a monophyletic group despite Nevertheless, the presence of a kink in the conspicuous differences in the form of the em- sperm duct is a useful part of the diagnosis of bolic division and cited it as evidence of how Carorita. greatly the embolic division can vary within Missing data.ÐAttributes of the tracheal an otherwise ``good'' genus. Unfortunately, I system have long been relied upon as char- have been unable to test this claim since I acters in linyphiid classi®cation (Blest 1976; have not been able to examine specimens of Millidge 1984, 1986). Hormiga (in press) C. paludosa and existing descriptions and il- demonstrated that these characters have high lustrations are not adequate for scoring many consistency. This analysis optimizes C. hiber- of the characters in the matrix. na as having the haplotracheate condition, al- Optimization.ÐThis analysis indicates though no observation of the tracheal system that the genus Carorita can be de®ned phy- of C. hiberna has been made. If C. hiberna is logenetically on the basis of two unambiguous speculatively coded as having the desmitra- synapomorphies: the presence of a radical tail- cheate condition (1: character 51) with no piece (character 21) and the loss of a lamella taenidia (0: character 52), 328 unique most characteristica (character 27). Although C. parsimonious trees of 237 steps result (192 paludosa has a large mesal sclerite of the em- trees in Hennig86, 246 trees each in NONA bolic division (Millidge 1977, ®g. 159), I have and PAUP; CI ϭ 0.376, RI ϭ 0.670). been unable to determine from published de- A total of 102 of these trees are more re- scriptions whether it is a radical tailpiece or a solved than otherwise compatible trees. The lamella characteristica. Five characters can be strict consensus of these 102 trees has little optimized to provide additional support for phylogenetic structure. It features a polytomy the monophyly of Carorita. However, these of 21 clades and individual taxa at the node ®ve characters could also be placed on alter- which represents the Erigoninae. Carorita native tree nodes without affecting tree length. limnaea and C. hiberna are not consistently These are the presence of papillae on the pro- monophyletic and form two of the branches tegulum (character 9; unknown in C. paludo- in this 21-tomy. Successive character weight- sa), a long embolus (character 17; short in C. ing of these 102 trees stabilizes on 30 trees paludosa), the shape of the radical tailpiece (CI ϭ 0.372, RI ϭ 0.666). These trees are two (character 22; unknown in C. paludosa), en- steps longer under equal weights than the capsulation of the epigynum (character 38; most parsimonious trees. The strict consensus unknown in C. hiberna and C. paludosa), and of these 30 trees forms a topology unlike that a tuberculate radical tailpiece (character 74; found by Hormiga (1999), especially in the unknown in C. paludosa). Although C. lim- deep nodes. Carorita limnaea and C. hiberna naea and C. hiberna are the only taxa in the form a paraphyletic assemblage rather than a analysis with a tuberculate radical tailpiece, monophyletic group. Under the assumption their two closest relatives, Asthenargus pa- that C. hiberna is haplotracheate rather than ganus, and Gongylidiellum vivum, both lack a desmitracheate without taenidia, the data are radical tailpiece and were therefore coded as found to be more internally consistent and the inapplicable for this character. A generic re- resulting phylogenetic hypothesis that is con- vision of Carorita with a strong phylogenetic sistent with the previous hypothesis (Hormiga component should be undertaken in the near in press), one which suffered from few miss- future to address the placement of C. palu- ing observations. dosa. All three Carorita species feature a strong DISCUSSION kink or loop in the path of the sperm duct in Phylogenetic hypotheses can be communi- the tegulum near the junction between the te- cated either implicitly as higher taxonomic gulum and the suprategulum (Figs. 7, 9; Mil- names or explicitly in the form of a clado- lidge 1977, ®g. 159). Most other erigonines gram. If is to re¯ect phylogenetic have a sperm duct that follows a smooth curve history, genera should serve as implicit hy- ZUJKO-MILLERÐSISICOTTUS HIBERNUS 49 pothesis that a particular group of species TAXONOMY share a unique common ancestor. However, monotypic genera do not provide this group- Carorita Duffey & Merrett 1963 ing information and thus cannot be considered Carorita Duffey & Merrett 1963:573±576, ®gs. 1± phylogenetic hypotheses. On the other hand, 8((,&). Duffey 1971: 14±15, ®gs. 1±8 ((,&). C cladistic analyses by their nature explicitly Locket, Millidge & Merrett 1974: 92±94, ®gs. convey detailed phylogenetic hypotheses. The 56a-g ((,&). Blest 1976: 188. Millidge 1977: 40, ranks and labels assigned to nodes in a clad- ®gs. 158, 159 ((); 1984: 245±246. Brignoli ogram are less critical since the hypothesis of 1983: 328. Roberts 1987: 108, ®gs. 53d, 53e relationships is fully expressed in the tree. As ((,&). Platnick 1989: 223; 1993: 252; 1997: 328. part of an explicit phylogenetic hypothesis, Heimer & Nentwig 1991: 124, ®gs. 352.1±353.4 detailed grouping information is available and ((,&). Type species by original designation and monotypic genera are less problematic. by monotypy, Oedothorax limnaeus Crosby & Bishop 1927: 149±150, ®gs. 11±14. The placement of Carorita hiberna was dif- ®cult because of the many apparently auta- Diagnosis.ÐMales of Carorita can be dis- pomorphic character states exhibited by this tinguished from most other erigonines by the species. However, no matter how many auta- form of the suprategulum which has a distinct pomorphic character states are found in a par- boundary at its origin and by the kinked or ticular species, it must still have a sister taxon looped path of the sperm duct through the te- (Platnick 1976). Placing Carorita hiberna in gulum (Figs. 7, 9). They are distinguished a new monotypic genus without identifying its from males of other erigonines with these putative sister taxon would have been an ab- character states by the absence of a lamella dication; a lack of a grouping hypothesis. As characteristica and the presence of a radical it stands, my circumscription of Carorita is tail piece (uncertain in C. paludosa). Females based on explicit evidence and is subject to can be distinguished from other erigonines by falsi®cation given suf®cient new evidence. the complex, anteriorly-projecting looped path Although these conclusions re¯ect the best of the copulatory ducts (Fig. 8). available evidence, missing data has led to the Description.ÐTibial macrosetae 2-2-1-1 optimization of some character states based on (except in C. hiberna: 2-2-2-1); Tm IV absent. inference (Platnick et al. 1991). When addi- Tibia I with one distal prolateral macroseta tional specimens are discovered, the predic- (absent in C. hiberna). At least in C. limnaea, tions of this analysis should be tested. chelicerae with imbricated stridulatory ®les Between 1981 and 1995, over 70% of new and median tracheal trunks unbranched, short- linyphiid genera have been monotypic (Plat- er than laterals, con®ned to abdomen (Blest nick 1989, 1993, 1997). The volume of new 1976; haplotracheate sensu Millidge 1984). monotypic genera generated within the Liny- Males: Palpal tibia with one prolateral and phiidae indicates a critical lack of phyloge- one retrolateral trichobothrium. Carorita lim- netic consideration. A phylogenetic approach naea and C. hiberna (but not C. paludosa) based on morphological characters demands with long embolus in frontal plane, radix with careful examination of comparative anatomy tuberculate tailpiece and no anterior radical but the result is a rich hypothesis not only of process (Figs. 3, 9). Distal part of cymbium taxonomic relationships but of character evo- and ectal side of palpal tibia with clusters of lution. A broad phylogenetic context will be macrosetae. Females: Females of C. paludosa very helpful for associating monotypic genera have not been examined and details of female with their relatives. This will no doubt lead to genitalia cannot be unambiguously interpreted the synonymization of many existing genera. using published descriptions and illustrations; Ultimately, the utility of the linyphiid taxo- females of C. hiberna unknown; females of nomic system will be greatly improved. How- C. limnaea with posteriorly oriented ventral ever, I have attempted to demonstrate that we plate invagination leading to copulatory open- do not need to wait for some grand phyloge- ings; copulatory ducts encapsulated with loop netic hypothesis of the Linyphiidae. The basic at anterior maximum and again posterior to context for applying phylogenetic criteria to spermathecae near junction. Fertilization ducts new work in linyphiid systematics is already project posteriomesally from spermathecae available. (Fig. 8). 50 THE JOURNAL OF ARACHNOLOGY

Composition.ÐThree species: Carorita North Carolina: Swain County, Great Smoky limnaea (Crosby & Bishop 1927), C. palu- Mountains National Park, Mingus Creek, 1 Febru- dosa Duffey 1971 and C. hiberna (Barrows ary 1943, 1(, (holotype, Barrows, OSU); Swain 1945). County, Great Smoky Mountains National Park, Distribution.ÐNorth America (C. lim- Thomas Ridge, ca. 200 m from trailhead at route 441, west-facing slope below trail, 4540 feet, old naea, C. hiberna) and Europe (C. limnaea, C. growth mixed hardwood, UTM: E3107, N39436, paludosa). 22 September 1994, 1 m2 litter sample, 1(, (Cribbs team, GSMNP); Haywood County, Great Smoky Carorita hiberna (Barrows 1945) Mountains National Park, Cataloochee, 150 m NEW COMBINATION south of mouth of Palmer Branch at Caldwell Fork, Figs. 1±7 2800±3000 feet, old growth hemlock, UTM: Sisicottus hibernus Barrows 1945: 74, ®gs. 1, 2 ((). E3107, N39436, 23±31 March 1997, pitfall trap, Brignoli 1983: 356. 1(, (Coyle, Edwards & Wright, GSMNP) North Carolina, Great Smoky Mountains National Park. Type.ÐMale holotype from United States: North Carolina, Great Smoky Mountains Na- Natural history.ÐCarorita hiberna is tional Park, Mingus Creek, 1 February 1943, known only from three male specimens col- in OSU, examined. lected in the Great Smoky Mountains National Diagnosis.ÐMales of C. hiberna are read- Park. The rarity of this species in the face of ily distinguished from other Carorita species intensive collecting efforts by Dr. Frederick by the unusual, complex shape of the embolus Coyle (Western Carolina University) and his which runs in a more or less frontal plane collaborators within the park raise questions (Fig. 3E), by their long, ectally projecting, tu- about the long term prospects for the contin- berculate, radical tailpiece (Fig. 3TP), by the ued survival of this species. The rare diplurid horn-like process on the ectal side of the pal- spider Microhexura montivaga Bishop & pal tibia (Fig. 6ETP), by the presence of a Crosby 1925 is known only from spruce and distal macroseta on tibia III, by the absence Fraser ®r forests in the southern Appalachians of a prolateral macroseta on tibia I, and by the and is currently listed as a federally protected presence of a dorsal radical process (Fig. 1, endangered species (Coyle 1981; Fridell DRP). 1995). The staphylinid beetle Dasycerus bi- Description.ÐMale: (from Thomas Ridge, color Wheeler & McHugh 1994 and the lin- North Carolina). Carapace length ϭ 0.6 mm. yphiid spider Sisicottus montigenus Bishop & Tibial macrosetae weak; 2-2-2-1; Tm I ϭ Crosby 1938 are both endemic to this same 0.34. Chelicerae with 5±6 promarginal teeth; region and habitat type, have experienced re- 5 retromarginal teeth with proximal 2 larger cent and dramatic declines in their popula- than distal 3. Embolus describing a semicir- tions, and may be worthy of similar protected cular arc in more or less frontal plane with status (Wheeler & McHugh 1994; Zujko-Mill- outside of curve ectal; tip of embolus arced in er 1999). However, C. hiberna has been found nearly transverse plane with outside of curve in both mixed hardwood and hemlock forests. dorsal (Figs. 1, 3E). Embolic membrane long, It does not appear to be associated with the narrow, curved with outside ectal (Fig. 2EM). declining spruce-®r habitat and though rare, Radix with anteriomesally projecting tuber- there is no evidence that its population has culate tailpiece (Fig. 3TP) and ectodorsally actually declined since its discovery. projecting dorsal radical process (Fig. 1DRP). ACKNOWLEDGMENTS Protegulum on ectal side of bulb (Fig. 2PT). The following persons and institutions Palpal tibia with long, straight apophysis with kindly loaned specimens for this study: R. ¯at distal margin; horn-like process on ectal Bradley, Ohio State University Museum of side of apophysis; ectal and mesal sides of Biodiversity (OSU); J.A. Coddington, Nation- palpal tibia both with regions of semitrans- al Museum of Natural History, Smithsonian parent chitin (Figs. 4-6). Female: Unknown. Institution (USNM); F.A. Coyle, Western Car- Distribution.ÐKnown only from the Great olina University -Great Smoky Mountains Na- Smoky Mountains National Park, North Car- tional Park Collection (GSMNP); N.I. Plat- olina. nick, American Museum of Natural History Material examined.ÐUNITED STATES: (AMNH). ZUJKO-MILLERÐSISICOTTUS HIBERNUS 51

G. Hormiga, J. Coddington, N. Scharff, N. Farris, J.S. 1969. A successive approximations ap- Platnick, B. Opell, and J. Berry made helpful proach to character weighting. Syst. Zool., 18: comments on an earlier draft of this manu- 374±385. script. I would like to thank G. Hormiga, J. Farris, J.S. 1988. Hennig86, version 1.5. Program Coddington, F. Coyle, N. Platnick, N. Scharff, and documentation. Computer program distrib- uted by D. Lipscomb, Dept. of Biol. Sci., The C. Griswold, D. Lipscomb, M. Allard, and J. George Washington Univ., Washington, D.C. Clark for stimulating discussion and advice. Fridell, J. 1995. Endangered and threatened wild- G. Hormiga generously allowed me to use life and plants; spruce-®r moss spider determined drawings and data from a manuscript in press to be endangered. Federal Register, 60:6968± in addition to unpublished data. F. Coyle kept 6974. an eye out for new specimens of Carorita hi- Goloboff, P.A. 1993. NONA. Noname (a bastard berna. The George Washington University son of Pee-Wee), version 1.6 (32 bit version). and National Museum of Natural History Program and documentation. Computer program (Smithsonian Institution) provided valuable distributed by J.M. Carpenter, Dept. of Entomol- lab space, materials, and equipment. Equip- ogy, American Museum of Natural History, New York. ment and ®nancial support was provided by Heimer, S. & W. Nentwig. 1991. Spinnen Mitte- an NSF-PEET grant to G. Hormiga and J. leuropas. Ein Bestimmungsbuch. Verlag Paul Coddington (DEB-9712353). Additional ®- Parey, Berlin. 542 pp. nancial support was provided by a Weintraub Holm, AÏ . 1979. A taxonomic study of European fellowship from The George Washington Uni- and east African species of the genera Pelecopsis versity. and Trichopterna (Araneae, Linyphiidae), with descriptions of a new genus and two new species LITERATURE CITED of Pelecopsis from Kenya. Zool. Scripta, 8:255± Barrows, W.M. 1945. New from the Great 278. Smokey [sic] Mountains National Park. Ann. En- Hormiga, G. 1994. Cladistics and the comparative tomol. Soc. America, 38:70±76. morphology of linyphiid spiders and their rela- Blest, A.D. 1976. The tracheal arrangement and tives (Araneae, Araneoidea, Linyphiidae). Zool. the classi®cation of linyphiid spiders. J. Zool., J. Linnean Soc., 111:1±71. London, 180:185±194. Hormiga, G. In press. Higher level phylogenetics Brignoli, P.M. 1983. A Catalog of the Araneae De- of erigonine spiders (Araneae, Linyphiidae, Eri- scribed Between 1940 and 1981. Manchester goninae). Smithsonian Contrib. Zool. Univ. Press, Manchester. 755 pp. Locket, G.H., A.F. Millidge & P. Merrett. 1974. Carpenter, J.M. 1988. Choosing among multiple British Spiders. Volume III. The Ray Society, equally parsimonious cladograms. Cladistics, 4: London. 314 pp. 291±296. Maddison, W.P. & D.R. Maddison. 1992. Mac- Coddington, J.A. 1983. A temporary slide mount Clade: Analysis of phylogeny and character evo- allowing precise manipulation of small struc- lution. Version 3.0. Sinauer Assoc., Sunderland, tures. Verh. Naturwiss. Ver. Hamburg, 26:291± Massachusetts. 292. Millidge, A.F. 1977. The conformation of the male Coddington, J.A. & N.I. Scharff. 1994. Problems palpal organs of linyphiid spiders, and its appli- with zero-length branches. Cladistics, 10:415± cation to the taxonomic and phylogenetic anal- 423. ysis of the family (Araneae, Linyphiidae). Bull. Coddington, J.A. & N.I. Scharff. 1996. Problems British Arachnol. Soc., 4:1±60. with ``soft'' polytomies. Cladistics, 12:139±146. Millidge, A.F. 1984. The taxonomy of the Liny- Coyle, F.A. 1981. The mygalomorph spider genus phiidae, based chie¯y on the epigynal and tra- Microhexura (Araneae, Dipluridae). Bull. Amer- cheal characters (Araneae: Linyphiidae). Bull. ican Mus. Nat. Hist., 170:64±75. British Arachnol. Soc., 6:229±267. Crosby, C.R. & S.C. Bishop. 1927. New species Millidge, A.F. 1986. A revision of the tracheal of Erigoneae and Theridiidae. J. New York En- structures of the Linyphiidae (Araneae). Bull. tomol., Soc., 35:147±157. British Arachnol. Soc., 7:57±61. Duffey, E. 1971. Carorita paludosa n. sp., a new Platnick, N.I. 1976. Are monotypic genera possi- linyphiid spider from Ireland and eastern Eng- ble? Syst. Zool., 25:198±199. land. Bull. British Arachnol. Soc., 2:14±15. Platnick, N.I. 1989. Advances in Spider Taxonomy Duffey, E. & P. Merrett. 1963. Carorita limnaea 1981±1987. Manchester Univ. Press, Manchester. (Crosby & Bishop), a linyphiid spider new to 637 pp. Britain, from Wybunbury Moss, Cheshire. Ann. Platnick, N.I. 1993. Advances in Spider Taxonomy Mag. Nat. Hist., 6:573±576. 1988±1991. New York Entomol. Soc. 846 pp. 52 THE JOURNAL OF ARACHNOLOGY

Platnick, N.I. 1997. Advances in Spider Taxonomy tes Menge (Araneida, Linyphiidae). ZooÈl. Med- 1992±1995. New York Entomol. Soc. 976 pp. ed. Leiden, 43:117±139. Platnick, N.I., C.E. Griswold & J.A. Coddington. Wheeler, Q.D. & J.V. McHugh. 1994. A new 1991. On missing entries in cladistic analysis. southern Appalachian species, Dasycerus bicolor Cladistics, 7:337±343. (Coleoptera: Staphylinidae: Dasycerinae), from Roberts, M.J. 1987. The Spiders of Great Britain declining endemic ®r forests. The Coleopts. and Ireland. Vol. 2. Linyphiidae and Check List. Bull., 48:265±271. Harley Books, Martins. 204 pp. Zujko-Miller, J. 1999. Revision and cladistic anal- Swofford, D.L. 1993. Phylogenetic analysis using ysis of the erigonine spider genus Sisicottus (Ar- aneae, Linyphiidae, Erigoninae). J. Arachnol., parsimony, version 3.1. Illinois Nat. Hist. Sur- 27(2):000±000. vey, Champaign, Illinois. van Helsdingen, P.J. 1968. Comparative notes on Manuscript received 1 May 1998, revised 7 Decem- the species of the Holarctic genus Stemonyphan- ber 1998.