NEW SPIDERS IN UPPER CRETACEOUS AMBER FROM NEW JERSEY IN THE AMERICAN MUSEUM OF NATURAL HISTORY (ARTHROPODA: ARANEAE)

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1 NEW SPIDERS IN UPPER CRETACEOUS AMBER FROM NEW JERSEY IN THE AMERICAN MUSEUM OF NATURAL HISTORY (ARTHROPODA: ARANEAE) by DAVID PENNEY ABSTRACT. The oldest described fossils of the extant spider family Araneidae (Araneinae; gen. et sp. indet.), the extant genus Orchestina (Oonopidae; O. sp. indet.) and the new fossil genus Palaeosegestria (Segestriidae; P. lutzzii gen. et sp. nov.) are presented from Upper Cretaceous amber of New Jersey. The known fossil range of the extant family Araneidae is extended approximately 50 myr from the previously oldest described araneid from the Middle Eocene oil shales of the Messel pit in Hesse, Germany. The fossil range of the extant genus Orchestina is also extended 50 myr from the previously oldest described specimen in Eocene Baltic amber. KEY WORDS: fossil, spider, Oonopidae, Segestriidae, Araneidae, Orchestina. F OSSILIFEROUS resins have been known from the Cretaceous of New Jersey for over a century and a half, but have had only brief mentions in the literature until recently (Grimaldi et al. 1989). Numerous authors have described a diverse fauna from these deposits (see Grimaldi 2000), but the only description of spiders from this deposit is that of Penney (2002a) who described the oldest known spider fossils, at that time, of the families Oecobiidae, Lagonomegopidae, Dictynidae, and Linyphiidae, and presented the oldest described Segestriidae and Oonopidae, from the same source. Grimaldi et al. (2002) reported a specimen of the extant jumping spider family Salticidae (previously unknown from the Mesozoic but the most diverse spider family today) in the New Jersey amber collections of the American Museum of Natural History; however this was a misidentification (pers. obs.). Penney (2002a) listed the spiders described from Cretaceous sources; since then a female Linyphiidae: Linyphiinae has been described from myr old (late Neocomian earliest Aptian) Cretaceous amber from the Kdeirji/Hammana outcrop, Lebanon (Penney and Selden 2002), which represents the oldest record of the family and the oldest described amber spider. Selden (2002) described the first British Mesozoic spider, identified as Nemesiidae, from amber of the Isle of Wight, and Selden and Penney (2003) described the first spider specimens from the Lower Cretaceous (Barremian) of Las Hoyas, Spain, which they placed in the Tetragnathidae. In a reassessment of the spiders previously described from the Lower Cretaceous of Montsech, Spain, Selden and Penney (2003) placed Palaeouloborus lacasae Selden, 1990 and Cretaraneus vitaltae Selden, 1990 in the families Uloboridae and Tetragnathidae: Nephilinae respectively. For a general review of spiders preserved in amber see Penney (2002b). Recently, new spiders have been discovered in the New Jersey amber collections of the American Museum of Natural History (AMNH), representing the families Oonopidae, Segestriidae and Araneidae, which are described below. LOCALITY AND STRATIGRAPHY The amber originates from the Raritan Formation, a locally restricted deposit from the Turonian (90 94 Ma) of central New Jersey, USA. The locality corresponds closest to site no. 5 in Grimaldi et al. (1989). Botanical inclusions indicate that the biological source of this amber was a pine (Pinaceae) or a member of the Taxodiaceae (Grimaldi et al. 2000), and not of araucarian origin as previously proposed (Grimaldi et al. 1989). The amber occurs in veins of compacted, lignitic peat, within deltaic deposits formed in coastal swamps (Grimaldi et al. 1997). These lie just above the South Amboy Fire Clay, which [Palaeontology, Vol. 47, Part 2, 2004, pp , 1 pl.] q The Palaeontological Association

2 368 PALAEONTOLOGY, VOLUME 47 marks the Raritan Formation. For a detailed account of the stratigraphy, taphonomy and palaeoecology of this site, see Grimaldi et al. (2000). All the specimens described in this paper were collected from White Oaks Pit, Sayreville, Middlesex County, New Jersey. MATERIAL AND METHODS The specimens upon which this paper is based are deposited in the American Museum of Natural History (AMNH), New York. Prior to receiving the specimens, they were embedded in clear synthetic resin for protection and to facilitate handling, then ground and polished by AMNH staff to reveal the inclusions. Much of the amber is opaque and has flaws running through it, which often obscure features of the inclusions, some of which are coated, in part, with a white substance of indeterminate origin. All measurements were made using an ocular graticule. Drawing was done under incident light with a camera lucida attached to an Olympus SZH stereomicroscope, and photographs were taken with a Nikon D1X digital camera attached to a Leica DMLM or Leica Wild M8 stereomicroscope. Drawings and photographs were manipulated in Adobe Photoshop and Adobe Illustrator. In the leg formula (e.g. 4123), the legs are ranked in order of length (longest first). Tti is the ratio of the distance that a trichobothrium is located from the base of the tibia, e.g. Tti1 ¼ 0 4 indicates that the trichobothrium is located four-tenths of the way along tibia 1, from the proximal end of the segment. Explanations of the morphological abbreviations used in the text, text-figures and plates are given in the appendix. SYSTEMATIC PALAEONTOLOGY Order ARANEAE Clerk, 1757 Suborder OPISTHOTHELAE Pocock, 1892 Infraorder ARANEOMORPHAE Smith, 1902 Family OONOPIDAE Simon, 1890 Genus ORCHESTINA Simon, 1882 Type species. Schoenobates pavesii Simon, 1873, by monotypy, Recent, from Spain, southern France, Corsica, Algeria. Orchestina sp. indet. Plate 1, figures 1 2; Text-figure b Oonopidae gen. et sp. indet.; Penney, p. 4, fig. 3. Material. Male, AMNH no. NJ 1012, in Cretaceous amber from New Jersey. The only known specimen. Coll. Ed Otte, Description. Body length 1 14 mm; carapace 0 57 mm long, 0 40 mm wide, raised but height not measurable, sides rounded, narrower in the ocular region, fovea absent; erect setae mid-dorsally and in the cephalic region, some of these are missing but their past presence is evident from the setal insertion sockets visible in the cuticle; clypeus approximately twice diameter PME with two pairs of long bristles curving inwards, converging at their tips (Pl. 1, fig. 1; Text-fig. 1). Six eyes, closely grouped in two rows but not contiguous, AME missing, PME slightly larger than ALE and PLE which are subequal; PER recurved and slightly wider than AER (Text-fig. 1). Chelicerae projecting downwards, long and thin, depressed on the disto-medial margin, and with scattered long setae projecting outwards, fangs short and unmodified, mouth parts not visible. Sternum shield-shaped, with scattered long setae and not projecting between the coxae. Opisthosoma 0 61 mm long, 0 57 mm wide, globular, with scattered long setae and lacking scuta; spinneret region ground off during preparation of the specimen prior to examination. Leg formula 1/423; leg 1 fe 0 43 mm, pa 0 14 mm, ti 0 33 mm, mt 0 31 mm, ta 0 23 mm, total 1 44 mm; leg 2 fe 0 40 mm, pa 0 14 mm, ti 0 31 mm, mt 0 31 mm, ta 0 21 mm, total 1 37 mm; leg 3 fe 0 29 mm, pa pnm, ti pnm, mt 0 30 mm, ta 0 20 mm; leg 4 fe 0 43 mm, pa 0 14 mm, ti 0 30 mm, mt 0 33 mm, ta 0 24 mm, total 1 44 mm; covered with long setae, leg spines absent. Femur 4 considerably thicker than fe 1 3 (Pl. 1, fig. 1; Text-fig. 1), Tti 1 4 ¼ 0 4, at least

3 PENNEY: CRETACEOUS SPIDERS 369 TEXT-FIG. 1. Camera lucida drawing of Orchestina sp. indet.; male (AMNH no. NJ 1012) in Upper Cretaceous amber from New Jersey. mt 1 þ 2 with one long and one short T distally, all leg tarsi with two claws on an onychium, each claw with approximately six teeth which become shorter and thicker distally. Pedipalp haplogyne and typical for the genus, consisting of a simple, swollen subspherical bulb 0 14 mm in diameter, which tapers off distally to form a long thin embolus. Both pedipalps are visible when the specimen is viewed ventrally; however the tip of neither embolus is visible (Pl. 1, fig. 2). Remarks. The spider is exquisitely preserved in a small mm piece of clear yellow amber. There are numerous small air bubbles but these do not obscure the detail of the specimen; there are no syninclusions. Wunderlich (1981) provided a diagnosis for Orchestina, which contains 36 extant species, six of which occur in the USA (Platnick 2002). In addition, numerous fossil species are known from various Tertiary ambers and semi-fossilized copals (Wunderlich 1981; Penney 2000), including a pair preserved during copulation (Wunderlich 1981). The high abundance of oonopids in amber is not surprising. They are small hunting spiders that actively pursue their prey, a behavioural trait which increases their likelihood of becoming trapped in resin (Penney 2002c). Unfortunately, the tip of the embolus of the palp is not visible due to the position of preservation, so it is not possible to differentiate this specimen from other known species. This specimen differs from that described by Penney (2002a) as gen. et sp. indet. by the position of the eyes and the leg spination. This is the oldest record of the genus Orchestina, extending the known range of this extant genus by a further 50 myr. The oldest fossil Orchestina prior to the discovery of this specimen was described from Baltic amber (e.g. Wunderlich 1981). Family SEGESTRIIDAE Simon, 1893 Genus PALAEOSEGESTRIA gen. nov. Derivation of name. Greek, palaios, old, and the extant genus, Segestria, which the fossil genus resembles. Type and only known species. Palaeosegestria lutzzii sp. nov.

4 370 PALAEONTOLOGY, VOLUME 47 Diagnosis. Male palp without accessory processes and with large subspherical bulb which tapers to form an elongated spiniform embolus. Each chelicera antero-distally with a distinct truncated conical cuticular outgrowth, upon which are situated at least four closely-grouped long, strong setae. The new genus is easily distinguishable by the structure of the chelicerae, which are unmodified in other segestriid genera. Female unknown. Description. See description of the type species below. Palaeosegestria lutzzii sp. nov. Plate 1, figures 3 5; Text-figure 2 Derivation of name. Named after K., J. and R. Luzzi, the collectors of the specimen, at the request of the AMNH. Type specimen. Male, AMNH no. NJ 1087, in Cretaceous amber from New Jersey. The only known specimen. Coll. K., J. and R. Luzzi. Diagnosis. As for the genus. Description. Body length 3 75 mm; carapace 1 75 mm long, 1 43 mm wide, covered with a fine pubescence of short setae, narrowed slightly but distinctly in the ocular region where there are a few longer setae; fovea absent. Eyes typical of the extant genus Segestria: six in three diads, occupying approximately two-thirds the carapace width, PME largest and contiguous, PLE slightly larger than the ALE which they almost touch, PER recurved and slightly wider than AER; clypeus short, 0 5 diameter PME (Pl. 1, figs 3 4; Text-fig. 2A). Chelicerae 0 64 mm long, 0 36 mm wide, projecting forwards slightly and without lamella, dentition if present not visible. Each chelicera with a brush of long, downward-projecting setae proximally and antero-distally with a distinct truncated, conical cuticular outgrowth, upon which are at least four closely-grouped long, strong setae (Pl. 1, fig. 5; Text-fig. 2A). Fangs, mouthparts and sternum not visible. Opisthosoma only visible dorsally, unmodified, 2 00 mm long, 1 25 mm wide, without scuta and uniformly covered with medium length setae; spinnerets not visible. Leg formula 1432 or 4132, left legs 2 and 4, and right leg 3 missing, left leg 1 detatched from the specimen but present in the amber; leg 1 fe 2 15 mm, pa 0 63 mm, ti 2 38 mm, mt 1 85 mm, ta 0 80 mm, total 7 81 mm; leg 2, fe 2 00 mm, pa 0 63 mm, ti 2 38 mm, mt 1 65 mm, ta 0 63 mm, total 7 29 mm; leg 3 fe pnm, pa 0 50 mm, ti 1 75 mm, mt 1 25 mm, ta 0 58 mm; leg 4 fe pnm, pa 0 63, ti pnm, mt pnm, ta pnm. Only the spination of leg 1 is particularly clear and is as follows: fe 2d-1p-0v-2r, pa 0d-0p-0v-0r, ti 0d-3p-3pairsv-3r, mt 0d-0p-1pairv-0r, ta 0d-0p-0v-0r; numerous erect hairs on ti, mt and ta, which are presumably trichobothria; mt and ta scopulate. Tarsi with three claws: inferior claw small, unmodified; superior claws each with a single row of teeth. Claw tufts absent; this is clearly visible in left leg 1 which has become separated from the spider but is preserved in the same piece of amber. Pedipalp fe 0 64 mm long with at least two slender dorsal spines; bulb large, subspherical 0 47 mm diameter, tapering to form an elongate spiniform embolus, accessory processes absent (Pl. 1, figs 3, 5; Text-fig. 2B). Remarks. The spider is preserved close to the surface of a relatively clear block (approximately mm) of yellowish amber. There are numerous air bubbles of various sizes, a reasonable amount of inorganic debris and numerous flaws in the piece, which obscure some features of the specimen, particularly when viewed ventrally. Extant species of Segestria are a common component of the tree-trunk EXPLANATION OF PLATE 1 Figs 1 2. Orchestina sp. indet.; male, AMNH no. NJ 1012 in Cretaceous amber from New Jersey. 1, dorsal view; 35. 2, anterior region, ventral view; 85. Figs 3 5. Palaeosegestria lutzzii gen. et sp. nov.; male, AMNH no. NJ 1087 in Cretaceous amber from New Jersey. 3, dorsal view; 9. 4, anterior view showing eyes; 17. 5, anterior view showing chelicerae and pedipalp; 17. Fig. 6. Araneidae: Araneinae, gen. et sp. indet.; juvenile?, AMNH no. NJ 364 (PN 157) in Cretaceous amber from New Jersey; dorsal view; 20.

5 PENNEY, Cretaceous spiders PLATE 1

6 372 PALAEONTOLOGY, VOLUME 47 TEXT-FIG. 2. Camera lucida drawing of Palaeosegestria lutzzii gen. et sp. nov. (AMNH no. NJ 1087) in Upper Cretaceous amber from New Jersey. A, anterior region of carapace, pedipalps and chelicerae which show the diagnostic features of the genus. B, pedipalp bulb and embolus. fauna. In the appropriate habitat a single tree can contain hundreds, if not thousands, of specimens (pers. obs.), identifiable solely by the presence of their highly distinctive web, which consists of numerous straight trip lines radiating outwards from a central tubular retreat. Furthermore, these webs are frequently found on damaged parts of the tree, which are more prone to secrete resin, for example, where branches have broken off. Penney (2002c) quantitatively identified a bias for amber preservation towards treedwelling faunas and it is not surprising, therefore, to find fossils of this family frequently preserved as amber inclusions (see Penney 2002a). Family ARANEIDAE Simon, 1895 Subfamily ARANEINAE Simon, 1895 Gen. et sp. indet. Plate 1, figure 6; Text-figure 3 Material. Juvenile?, AMNH no. NJ 364 (PN 157), in Cretaceous amber from New Jersey. The only known specimen. Coll. Paul Nascimbene, Description. Body length 4 13 mm; carapace length 1 50 mm, width 1 50 mm, flat, narrowed anteriorly. Eight subequal eyes in two rows, laterals appear to be touching, the typical araneid arrangement, although the right PME appears to be set back slightly; clypeus low. Opisthosoma 2 63 mm long, 2 50 mm wide, globose; a hole has accidentally been

7 TEXT-FIG. 3. Camera lucida drawing of Araneidae: Araneinae, gen. et sp. indet. [AMNH no. NJ 364 (PN 157)] in Upper Cretaceous amber from New Jersey. PENNEY: CRETACEOUS SPIDERS 373 ground into the dorsal surface during preparation of the specimen; the same is true for the patella and distal aspect of the femur of left leg 1. The specimen is preserved such that it is not possible to make leg measurements, but by eye, the leg formula appears to be either 1432 or The specimen is interesting in that distinct leg annulations have been preserved (Pl. 1, fig. 6; Text-fig. 3). These and the numerous stong leg macrosetae present on many of the segments, in addition to the general habitus of the specimen, are typical of many extant Araneidae: Araneinae. Remarks. The spider is preserved close to the surface and the end of a yellow-orange block of amber (approximately mm). The end with the spider inclusion is relatively clear although there are numerous flaws, densely packed tiny air bubbles and many tiny pieces of organic and inorganic matter, which hinder a clear view of the specimen. In addition the spider is coated ventrally with a white film typical of many specimens preserved in amber. Although it is not possible to see any autapomorphic characters that would unequivocally place this specimen in the extant family Araneidae, I think it is correctly placed in this family. The above combination of characters visible in the fossil and its general habitus are practically identical to those of many extant araneids and I am unaware of another family that shares these attributes to the same degree. It is now evident that many extant spider families have a long geological history (e.g. Selden and Penney 2001) and there is no reason to believe the Araneidae are any different. Indeed, it is considered the most basal family of the aerial web-weaving superfamily Araneoidea (i.e. it is sister to the remaining araneoids) and the presence of fossil Tetragnathidae in the earliest Cretaceous (Selden and Penney 2003) predicts the presence of the Araneidae then too (Selden and Penney 2001). In addition, Wunderlich (1986a) suggested that the fossil described by Eskov (1984) as Juraraneus in the extinct family Juraraneidae from Transbaikalia, Siberia may actually belong in the Araneidae, which would extend the known range of the family back to the Jurassic. However, this synonymy has not yet been formally established and thus the fossil described here represents the oldest known member of the family, extending the known geological range by approximately 50 myr from the previously oldest described araneids in Baltic amber (e.g. Petrunkevitch 1942) and the Messel oil shales, Germany (Wunderlich 1986b). The family has been reported, but not described, from Cretaceous (Campanian) Canadian amber (McAlpine and Martin 1969).

8 374 PALAEONTOLOGY, VOLUME 47 DISCUSSION The New Jersey amber spider fauna is now the best recorded source of Mesozoic fossil Araneae in terms of described specimens (Penney 2002a and herein). There are ten specimens described and figured from six extant and one extinct family. Three genera are extant (Oecobius, Orchestina and Segestria) and two are currently only known from fossils (Lagonomegops and Palaeosegestria). I have examined approximately 30 spiders in the New Jersey amber collections of the AMNH but unfortunately many of the specimens are juvenile or poorly preserved. Both prohibit the identification of specimens certainly to genus and often even to family level. It is unlikely that this source will yield many more fossil spiders as the excavation of the New Jersey amber site by the AMNH staff is now finished. There are a few New Jersey amber collections in private hands, but the AMNH received the best specimens from this locality. The fossil spiders from this source are important because they represent the oldest examples of a number of extant spider taxa and can also predict the presence of other extant spider taxa at the same point in geological time (e.g Penney 2002a). However, the assemblage is not particularly useful as it stands for addressing palaeoecological problems, because none of the spider taxa identified to date from this fossil resin can be considered indicators for specific climate or habitat types. The genera Oecobius and Orchestina are considerably more common in the tropics and subtropics, but they are also found in temperate regions. However, the assemblage may prove useful for addressing these problems when considered in conjunction with other taxa from the same source or when compared with spiders from other fossil deposits. A number of other Mesozoic deposits such as Burmese and Lebanese ambers appear very promising from a palaeoarachnological perspective and this material is currently under study. Acknowledgements. I thank David Grimaldi (AMNH) for the preparation and loan of these specimens and Paul Selden (University of Manchester) for providing comparative Recent material. This paper was completed under Leverhulme Trust grant F/00 120I, Mesozoic arachnids. REFERENCES CLERK, C Aranei suecici, descriptionibus et figuris oeneis illustrati, ad genera subalterna redacti speciebus ultra 60 determinati. Svenska Spindlar, uti sina hufvud-slagter indelte samt. L. Salvii, Stockholm, 154 pp., 6 pls. ESKOV, K. Y A new fossil spider family from the Jurassic of Transbaikalia (Araneae, Chelicerata). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 1984, GRIMALDI, D. (ed.) Studies in fossils in amber, with particular reference to the Cretaceous of New Jersey. Backhuys, Leiden, 498 pp. AGOSTI, D. and CARPENTER, J. M New and rediscovered primitive ants (Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic relationships. American Museum Novitates, 3208, BECK, C. W. and BOON, J. J Occurrence, chemical characteristics and palaeontology of the fossil resins from New Jersey. American Museum Novitates, 2948, ENGEL, M. S. and NASCIMBENE, P. C Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance. American Museum Novitates, 3361, SHEDRINSKY, A. and WAMPLER, T A remarkable deposit of fossiliferous amber from the Upper Cretaceous (Turonian) of New Jersey In GRIMALDI, D. (ed.). Studies on fossils in amber, with particular reference to the Cretaceous of New Jersey. Backhuys, Leiden, 498 pp. MCALPINE, J. F. and MARTIN, J. E. H Canadian amber a paleontological treasure chest. Canadian Entomologist, 101, PENNEY, D Miocene spiders in Dominican amber (Oonopidae, Mysmenidae). Palaeontology, 43, a. Spiders in Upper Cretaceous amber from New Jersey (Arthropoda: Araneae). Palaeontology, 45, b. Arachnological gems: spiders in amber. Newsletter of the British Arachnological Society, 94, c. Paleoecology of Dominican amber preservation: spider (Araneae) inclusions demonstrate a bias for active, trunk-dwelling faunas. Paleobiology, 28, and SELDEN, P. A The oldest linyphiid spider, in Lower Cretaceous Lebanese amber (Araneae, Linyphiidae, Linyphiinae). Journal of Arachnology, 30, PETRUNKEVITCH, A A study of amber spiders. Transactions of the Connecticut Academy of Arts and Sciences, 34, , 69 pls.

9 PENNEY: CRETACEOUS SPIDERS 375 PLATNICK, N. I The world spider catalogue version catalog81-87/ POCOCK, R. I Liphistius and its bearing upon the classification of spiders. Annals and Magazine of Natural History, Series 6, 10, SELDEN, P. A Lower Cretaceous spiders from the Sierra de Montsech, north-east Spain. Palaeontology, 33, First British Mesozoic spider, from Cretaceous amber of the Isle of Wight. Palaeontology, 45, and PENNEY, D Phylogeny of Araneae: the fossil evidence and its interpretation. p In: Abstracts of the 15th International Congress of Arachnology, Badplaas, South Africa, March Lower Cretaceous spiders (Arthropoda: Arachnida: Araneae) from Spain. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 2003, SIMON, E Aranéides nouveaux ou peu connus du didi de l Europe (2 ème Mémoire). Mémoires de la Société Royale des Sciences de Liège, 5, 1 174, 3 pls Etudes arachnologiques. 13 e Mémoire. 20. Descriptions d espèces et de genres nouveaux de la famille des Dysderidae. Annales de la Société Entomologique de France, 2, Etudes arachnologiques. 22 e Mémoire. 34. Etude sur les Arachnides de l Yemen. Annales de la Société Entomologique de France, 10, Histoire naturelle des Araignées. Volume 1, part 2. Roret, Paris, pp , figs Histoire naturelle des Araignées. Volume 1, part 4. Roret, Paris, pp , figs SMITH, F. P The spiders of Epping Forest. The Essex Naturalist, 12, WUNDERLICH, J Fossile Zwerg-Sechsaugenspinnen (Oonopidae) der Gattung Orchestina Simon, 1882 im Bernstein, mit anmerkungen zur Sexual-Biologie (Arachnida, Araneae). Mitteilungen aus dem Geologisch- Paläontologischen Institut der Universität Hamburg, 51, a. Spinnenfauna gestern und heute. Fossile Spinnen in Bernstein und ihre heute lebenden Verwandten. Erich Bauer Verlag bei Quelle und Meyer, Wiesbaden, 283 pp. 1986b. Die ersten Spinnen aus dem Mittel-Eozän der Grube Messel. Senckenbergiana Lethaea, 67, Typescript received 29 July 2002 Revised typescript received 6 January 2003 DAVID PENNEY Department of Earth Sciences University of Manchester Manchester M13 9PL, UK David.Penney@man.ac.uk APPENDIX Abbreviations used in text, text-figures and plates AER, anterior eye row; ALE, anterior lateral eye(s); AME, anterior median eye(s); b, bulb; c, clypeus; car, carapace; cb, cheliceral brush; ch, chelicera(e); co, cheliceral cuticular outgrowth; cs, closely-grouped long, strong cheliceral setae; cx, coxa; d, dorsal; e, embolus; f, flaw in amber; fe, femur; h, hole at the surface of the resin where the inclusion has been accidentally ground away during preparation prior to receipt by the author; la, leg annulations; m, macroseta; mt, metatarsus; op, opisthosoma; p, prolateral; Pp, pedipalp; pa, patella; PER, posterior eye row; PLE, posterior lateral eye(s); PME, posterior median eye(s); pnm, leg segment present but not measurable; r, retrolateral; T, trichobothrium; ta, tarsus; tc, tarsal claws; ti, tibia; tr, trochanter; v, ventral; 1 4, legs 1 4.