PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3462, 7 pp., 8 ®gures December 30, 2004

On a Third Group of Flattened Ground from Australia (Araneae, Lamponidae)

NORMAN I. PLATNICK1

ABSTRACT A new genus and species, Platylampona mazeppa, are described for an unusual, newly discovered ground from Queensland and northern New South Wales. Although the body of these is greatly ¯attened, as is characteristic of both trochanteriids and hemicloeine gnaphosids, the new genus belongs to neither of those groups but rather to the subfamily Lamponinae, and thus represents a third lineage of Australian gnaphosoids that has acquired a greatly ¯attened habitus.

INTRODUCTION dorsoventral height of the appendages). Out- side of Australia, such taxa are relatively Among the numerous Australian spiders belonging to the superfamily Gnaphosoidea rare, and have thus provided inducement for are several genera notable for having greatly the establishment of such classical family- ¯attened bodiesÐso much so that many of group names as the Trochanteriidae Karsch these taxa look like animals that have been (1879) and Platoridae Simon (1890). In Aus- smashed under an unabridged dictionary. In- tralia, however, they form a signi®cant pro- deed, their bodies are often so ¯attened that portion of the ground spider fauna, and are the legs are laterigrade (i.e., resemble those frequently found under the peeling bark of of crab spiders in being twisted, with the eucalypts. morphologically anterior surfaces directed Some authors (e.g., Wunderlich, 1987) dorsally, thereby minimizing the effective have considered all these extremely ¯attened

1 Peter J. Solomon Family Curator, Division of Invertebrate Zoology, American Museum of Natural History; Ad- junct Professor, Department of Biology, City College, City University of New York; Adjunct Professor, Department of Entomology, Cornell University; Adjunct Senior Research Scientist, Center for Environmental Research and Con- servation, Columbia University ([email protected]).

Copyright ᭧ American Museum of Natural History 2004 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3462 taxa to be members of a single monophyletic the area of the major ampullate gland spig- group, but spinneret morphology indicates ots. As a result, the area surrounding the pir- that they actually belong to two separate iform gland spigots consists of a mound of families (Platnick, 1990, 2002). Most of the soft, expandable cuticle. These features in- ¯attened genera are ``lower gnaphosoids'', dicate that Platylampona is a ``higher'' rather belonging to the Trochanteriidae and retain- than ``lower'' gnaphosoid (i.e., a member of ing a sclerotized, complete distal ring on the the Lamponidae, Prodidomidae, or Gnaphos- anterior lateral spinnerets, but Hemicloea idae, rather than the Ammoxenidae, Cithae- Thorell (1870) is a ``higher gnaphosoid'', ronidae, , or Trochanteriidae; having lost that distal ring. By losing the dis- Platnick, 1990, 2002). tal ring, the ``higher gnaphosoids'' (i.e., Although insuf®cient specimens have been members of the families Lamponidae, Prod- available to examine the spinnerets of Pla- idomidae, and Gnaphosidae) have gained the tylampona with scanning electron microsco- ability to expand and contract the soft cuticle py, it seems clear from light microscopy that surrounding the piriform gland spigots on the piriform gland spigots of these spiders their anterior lateral spinnerets. Hemicloea are both few in number (perhaps only one or has thus generally (and correctly) been two) and small in size, relative to the major placed in the family Gnaphosidae, as it also ampullate gland spigots. These features in- has the greatly enlarged and widened piri- dicate that the species is a lamponid, rather form gland spigots, situated on an expand- than a prodidomid or gnaphosid (members of able mound, that are putatively synapo- those families have enlarged piriform gland morphic for that family (Platnick, 1990). spigots that are either greatly elongated or It was surprising, however, to discover an- widened; see Platnick, 1990). other Australian ground spider with a greatly The placement of Platylampona in the ¯attened habitus that seems to be neither a Lamponidae is corroborated by the presence trochanteriid nor a hemicloeine gnaphosid. of the pair of invaginated abdominal scler- The present paper is devoted to a description ites, situated just behind the booklung open- of this unusual species, and an exploration of ings, that appear to be synapomorphic for its relationships. The format of the descrip- that family (Platnick, 2000). These sclerites tion follows that of Platnick (2000). Material are small and inconspicuous in females of has been examined from the collections of Platylampona, but in males they are enor- the Australian Museum, Sydney (AMS), the mously expanded, and cover about one-®fth Australian National Insect Collection, Can- of the abdominal venter (®g. 2). Such ex- berra (ANIC), and the Queensland Museum, panded sclerites have been found also in Brisbane (QMB). some males of Lamponina elongata Platnick (2000: 129). Both sexes of Platylampona RELATIONSHIPS also have a greatly enlarged and bent seta on the cheliceral promargin, adjacent to the base The intriguing species described below as of the fang. This character optimized as a Platylampona mazeppa (®gs. 1±3) seems lamponid synapomorphy in the analysis of clearly to be a member of the subfamily Platnick (2000) but is also widespread out- Lamponinae. Its placement as a gnaphosoid side the group, and its distribution needs to is supported by the irregularly rectangular be documented in detail across a wide spec- and completely ¯attened posterior median trum of dionychans before its phylogenetic eyes, which have their tapeta obliquely signi®cance can be assessed. aligned and converging posteriorly (implying Specimens of Platylampona obviously that these eyes are used not to focus an image lack the elevated pars cephalica typical of the but rather to determine the direction of po- lamponid subfamily Pseudolamponinae, and larized light; Dacke et al., 1999). have the distinctive longitudinal groove near The anterior lateral spinnerets of this spe- the median edge of the endites that is char- cies are composed of a single segment; the acteristic of the subfamilies Centrothelinae sclerotized distal ring representing a second and Lamponinae instead. Females lack the segment is reduced to a crescent surrounding row of three greatly enlarged cylindrical 2004 PLATNICK: AUSTRALIAN FLATTENED GROUND SPIDERS 3

Figs. 1±3. Platylampona mazeppa, new species, male. 1. Dorsal view. 2. Ventral view. 3. Lateral view of cephalothorax and abdomen. gland spigots on the posterior median spin- nig86, version 1.5 (Farris, 1988), the h* and nerets that is characteristic of centrothelines. bb* tree-generating options produced 2070 Their placement in the subfamily Lamponi- cladograms of 52 steps; three rounds of suc- nae is supported by the typically lamponine cessive approximations weighting stabilized pedicel sclerites, which form a tube. The en- on 48 cladograms with a length of 276. Even dites are also evenly rectangular, as in other the strict consensus of those 48 cladograms lamponines, although (as one would expect resolves all but one of the previously hy- in such a ¯attened ), the labium and pothesized nodes (the genera Pseudolam- endites are longer than in other lamponines pona Platnick and Paralampona Platnick do (®g. 2). not always cluster together), but in all cases To test this placement, the cladistic anal- Platylampona forms a trichotomy with the yses of Platnick (2000, ®gs. 1, 2) were rerun, lamponine genera Lamponega Platnick and with Platylampona added to the matrix; the Lamponicta Platnick. character coding for this extra row is 01101 In Pee-Wee, version 2.6 (Goloboff, 1997) 01011 11111 10000 01000 00001. In Hen- implied weighting of the expanded matrix 4 AMERICAN MUSEUM NOVITATES NO. 3462 produced just three cladograms at all six set- rather than enlarged, piriform gland spigots tings of the concavity function. Here again, on the anterior lateral spinnerets). Platylampona always clustered with Lam- DESCRIPTION: Small spiders, total length ponega and Lamponicta, sharing with those 2.9±3.8. Carapace dark red, greatly ¯attened, two genera the fusion of the chilum with the tuberculate, coated with long setae originat- anterior margin of the carapace. ing from bases of tubercles, tubercles pro- It is common, among gnaphosoids with truding from sinuous lateral margin (®g. 1); such a ¯attened body, for the fourth pair of thoracic groove short, almost obsolete. Eight trochanters to be elongated. This character is eyes in two rows, eyes large, subequal in conspicuous in some trochanteriids (hence size; anterior medians circular, dark, other the family name) and also in some hemiclo- eyes light, laterals oval, posterior medians ir- eine gnaphosids. In Platylampona, however, regularly rectangular, ¯attened; from above, the laterigrade leg position is accommodated both eye rows slightly procurved, from front, by elongation of the fourth pair of coxae, anterior row slightly procurved, posterior rather than trochanters (®gs. 1, 2). row strongly procurved; anterior medians separated by almost their diameter, slightly FAMILY LAMPONIDAE closer to anterior laterals; posterior medians separated by almost their diameter, separated SUBFAMILY LAMPONINAE by more than their diameter from posterior Platylampona, new genus laterals; anterior and posterior laterals sepa- rated by more than their diameter; median TYPE SPECIES: Platylampona mazeppa, ocular quadrangle about as wide in front as new species. in back, slightly longer than wide. Chelicer- ETYMOLOGY: The generic name refers to ae, sternum, and mouthparts dark red; chilum the general appearance of these ¯attened apparently fused to carapace, which is pro- lamponids, and is feminine in gender. longed at midline; second, short, posterior DIAGNOSIS: In the existing key to lampon- chilum (extremely narrow sclerite separating ine genera (Platnick, 2000: 14), specimens of bases of chelicerae posteriorly) present; che- Platylampona will key out to Lamponova licerae with deeply excavated ridge opposite Platnick because the ventral surface of meta- carapace margin, followed distally by dis- tarsi and tarsi I and II does have a divided tinct lateral boss and large median boss oc- (if weak) scopula, the sternal surface is cupying most of paturon width, promargin smooth with circular punctations rather than with series of short setae originating in line rugose, and the female epigynum lacks an along base of fang, with long, thick, modi®ed anterior hood (males of Lamponova remain seta originating closest to fang; promargin unknown). The greatly ¯attened habitus with teeth obsolete, represented by fused (®gs. 1±3) and sinuous lateral margins of the ridge, retromargin without teeth; cheliceral carapace (®g. 1) immediately separate the ge- gland openings not scanned. Labium elon- nus from Lamponova and all other lampon- gated, truncate anteriorly and posteriorly, ids. In having the chilum fused to the anterior distinctly depressed medially; anterior sur- margin of the carapace, producing a medial face not scanned. Endites obliquely de- prolongation of that margin, Platylampona pressed, with sharply demarcated, deep specimens resemble those of Lamponega and groove along margin near labium; serrula Lamponicta; they can be distinguished from long, with single row of teeth; anterior sur- the former by the ¯attened (rather than great- face not scanned. Sternum ¯attened, with in- ly elevated) sternum and from the latter by clined sides, fused to epimeric sclerites so the presence of a retrolateral tibial apophysis that coxae originate from foramina complete- on the male palp and the absence of a pos- ly enclosed by sclerotized cuticle (except be- terior spermathecal bulb in females. As in- tween legs III and IV of females, where the dicated above, specimens might be confused fusion is not entire, with tiny strip of unscler- with either trochanteriids (but lack a distal otized cuticle remaining between sternum ring on the anterior lateral spinnerets) or and epimeric sclerite); surface smooth, with hemicloeine gnaphosids (but have small, circular punctations. Epimeric sclerites not 2004 PLATNICK: AUSTRALIAN FLATTENED GROUND SPIDERS 5 fused with carapace. Pedicel elongated, com- retrolateral tibial apophysis; tegulum expand- posed of two small, ¯at dorsal sclerites and ed, extending beyond posterior rim of tarsus; rounded sclerite covering venter and sides, embolus laterally situated, inset into mem- rounded sclerite with slight median longitu- branous conductor; median apophysis absent. dinal keel, without anterior protrusion ex- Epigynum with v-shaped median septum; tending toward sternum. spermathecae on short stalks. Anterior edge of abdomen in males with rectangular dorsal scutum extending to al- Platylampona mazeppa, new species most half of abdominal length; dorsal scutum Figures 1±8 anteriorly separated from epigastric scutum only by narrow strip of unsclerotized cuticle; TYPE: Male holotype taken by pyrethrum females without dorsal scutum; cuticle with sampling of gidgee (an indigenous group of long dark setae; epigastric scutum accompa- Acacia species) trunks at an elevation of 240 nied posterolaterally by pair of oval, deeply m at the north end of Mazeppa National invaginated sclerites bearing clearly elevated Park, 22Њ14ЈS, 147Њ15ЈE, Queensland (Mar. anterior rim; anterior edge of oval sclerites 27, 2001; G. Monteith), deposited in QMB ®tting under epigastric scutum; sclerites of (S55138). male expanded into rectangular plates occu- ETYMOLOGY: The speci®c name is a noun pying about one-third of postepigastric por- in apposition taken from the type locality. tion of abdominal venter; colulus represented DIAGNOSIS: With the characters of the ge- by transverse row of setae posterior of tiny nus and genitalia as in ®gures 4±8. transverse sclerite presumably marking po- MALE (holotype): Total length 2.9. Dorsal sition of small posterior spiracle. Anterior and postepigastric abdominal scuta light or- lateral spinnerets tubular, separated by rough- ange, unsclerotized portion of abdominal ly their diameter, cuticle representing distal, dorsum mottled gray, venter unmarked; legs second spinneret segment restricted to semi- light orange, unmarked, anteriors darkest. circle surrounding major ampullate gland Palpal femur with distinct ventral enlarge- spigots (piriform gland spigots surrounded ment at about one-third its length; patella only by soft cuticle, apparently only one or dorsoventrally ¯attened, laterally widened; two present); posterior median spinnerets tibia short, retrolateral portion about as long small, tubular, without anteriorly expanded as retrolateral tibial apophysis, dorsally and tips; posterior lateral spinnerets two-seg- prolaterally prolonged, with one or two thick mented, spigots unscanned. macrosetae on prolongation; palpal tegulum Legs spineless, laterigrade; most surfaces enlarged, protruding ventrally beyond base with long, dark setae; anterior coxae with of cymbium, embolus situated prolaterally, protuberant posterolateral corners; coxae tu- apically extended into ®ne point (®gs. 4±6). berculate, coxae IV much longer than others; FEMALE (from type locality): Total length trochanters unnotched; anterior metatarsi and 3.8. Coloration as in male (but without ab- tarsi with weak but distinct, divided scopulae dominal scuta). Epigynum with anterior mar- composed of short, thickened, bent setae; gins and median septum forming m-shaped posterior metatarsi with thick, dark, distal ridge, openings apparently at anterolateral preening brushes (stronger on leg III than corners of median septum; spermathecae IV); posterior tarsi with weak, entire scopu- oval, on short stalks (®gs. 7, 8). lae; tarsi with two ®nely dentate claws, claw VARIATION: The male palpal tibia can have tufts composed of lateral pads of closely ap- either one strong dorsal macroseta (as shown pressed setae; trichobothria present on tibiae, in ®gs. 4, 6) or two, but the distinction does metatarsi, and tarsi, in 2±3 irregular rows, not seem to be taxonomically signi®cant, as bases unscanned; tarsal organ unscanned. Fe- the number can differ on the left and right male palpal tarsus without spines, with tiny palps of a single specimen. claw. OTHER MATERIAL EXAMINED: AUSTRA- Male palp with retrolateral tibial apophysis LIA: New South Wales: Atholwood Loop accompanied by strong macroseta, cymbial Road, 500 m N of road at point opposite Ret- surface with retrobasal excavation opposite ford Springs, Severn State Forest, 6 AMERICAN MUSEUM NOVITATES NO. 3462

Figs. 4±8. Platylampona mazeppa, new species. 4. Left male palp, prolateral view. 5. Same, ventral view. 6. Same, retrolateral view. 7. Epigynum, ventral view. 8. Same, dorsal view.

29Њ04Ј28ЉS, 151Њ00Ј53ЉE, Nov. 22±Dec. 13, elev. 160 m, pyrethrum, brigalow (G. Mon- 2001, pitfall (L. Wilkie, H. Smith, AMS teith, QMB S52782), 1(; 5 km SW Mount KS81891), 1(; 16 km S Texas (Queensland), Robert, 21Њ24ЈS, 148Њ27ЈE, Mar. 26, 2001, 29Њ00ЈS, 151Њ09ЈE, Nov. 24, 1983 (D. Rentz, elev. 300 m, pyrethrum, tree trunks and logs, M. Harvey, ANIC), 2&. Queensland: 3km brigalow (G. Monteith, QMB S55136), 1(. NW Langlo Crossing, 26Њ07ЈS, 145Њ39ЈE, DISTRIBUTION: Known only from central May 4, 2001, pyrethrum, gidgee (G. Mon- and southern Queensland and northeastern teith, QMB), 1&; S end, Mazeppa National New South Wales. Park, 22Њ16ЈS, 147Њ16ЈE, Dec. 18, 2000±Mar. 26, 2001, elev. 240 m, ¯ight intercept trap, ACKNOWLEDGMENTS brigalow (D. Cook, G. Monteith, QMB), 1&; site 3, Mazeppa National Park, 22Њ16ЈS, This work was supported by a National 147Њ17ЈE, Mar. 27, 2001, elev. 240 m, py- Science Foundation PEET (Partnerships for rethrum, gidgee trunks (G. Monteith, QMB Enhancing Expertise in ) award S55141), 1(; on road 7 km NNE Mount (DEB-9521631). I'm indebted to Barbara Bluffkin, 22Њ36ЈS, 149Њ14ЈE, June 2, 2002, Baehr, Val Davies, and Robert Raven 2004 PLATNICK: AUSTRALIAN FLATTENED GROUND SPIDERS 7

(QMB), Mike Gray and Graham Milledge Platnick, N.I. 2000. A relimitation and revision of (AMS), and Bruce Halliday (ANIC) for ac- the Australasian ground spider family Lampon- cess to specimens and help during museum idae (Araneae: Gnaphosoidea). Bulletin of the visits and ®eldwork, and to Mohammad American Museum of Natural History 245: 1± Shadab (AMNH) for his work on the illus- 330. Platnick, N.I. 2002. A revision of the Australasian trations. ground spiders of the families Ammoxenidae, , Gallieniellidae, and Trochanter- REFERENCES iidae (Araneae: Gnaphosoidea). Bulletin of the Dacke, M., et al. 1999. Built-in polarizers form American Museum of Natural History 271: 1± part of a compass organ in spiders. Nature 401: 243. 470±473. Simon, E. 1890. E tudes arachnologiques. 22e MeÂ- Farris, J.S. 1988. Hennig86, version 1.5. Com- moire. XXXIV. E tude sur les arachnides de puter program distributed by its author. l'Yemen. Annales de la SocieÂte Entomologique Goloboff, P. 1997. Pee-Wee, version 2.6. Com- de France (6) 10: 77±124. puter program distributed by its author. Thorell, T. 1870. Araneae nonnullae Novai Hol- Karsch, F. 1879. Arachnologische BeitraÈge. Zeit- landiae, descripte. OÈ fversigt af Konglige Veten- schrift fuÈr die Gesammten Naturwissenschaften skaps-Akademiens FoÈrhandlingar 27: 367±389. 52: 534±562. Wunderlich, J. 1987. Die Spinnen der Kanarisch- Platnick, N.I. 1990. Spinneret morphology and the en Inseln und Madeiras: Adaptive Radiation, phylogeny of ground spiders (Araneae, Gna- Biogeographie, Revisionen und Neubeschrei- phosoidea). American Museum Novitates 2978: bungen. Langen, West Germany: Triops Verlag, 1±42. 435 pp. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last ®ve years are available at World Wide Web site http://library.amnh.org. Or address mail orders to: American Museum of Natural History Library, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-5545. FAX: (212) 769- 5009. E-MAIL: [email protected]

a This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).