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FIRST FOSSIL FILISTATIDAE: A NEW SPECIES OF MISIONELLA IN MIOCENE AMBER FROM THE DOMINICAN REPUBLIC PDF

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2005. The Journal of Arachnology 33:93-100 FIRST FOSSIL FILISTATIDAEi A NEW SPECIES OF MISIONELLA IN MIOCENE AMBER FROM THE DOMINICAN REPUBLIC Da¥M Penney: Earth Sciences, The University of Manchester, Manchester, Ml3 9PL, United Kingdom ABSTRACT. Misionella didicostae new species is described from 15-20 Ma Miocene amber from the Dominican Republic as the first fossil record of the family Filistatidae. The biogeography of the extant (Brazil and Argentina) and the new fossil species supports the hypothesis that the developing northern Greater Antilles and northwestern South America were briefly (33-35 Ma) connected by a landspan centered on the ero.ergeiit Aves Ridge. Undiscovered extant species ofMisionella may existon Hispaniola. The autospasized firstpair oflegs suggestthatthe spiderwas engulfed in a flowingresin seep ofrelatively low viscosity, rather than having wandered onto a sticky exudate, becoming stuck and then covered by a subsequent resin flow. Keywords: Hispaniola, Araneae, spider, biogeography, autospasy The spider family Filistatidae has an almost amber as a syninclusion. The holotype and worldwide distribution in tropical and warm only known specimen is held in the collection temperate regions, and consists of 107 species of the Museo del Ambar Dominicano, Puerto and one subspecies in 16 genera (Platnick Plata, Dominican Republic. 2003). These small to medium-sized, cribeh Measurements and drawings,—All mea- late spiders represent one of the most basal surements were made using an ocular grati- branches of the Haplogynae (Platnick et al. cule and are in mm. Drawings were done un- 1991). Filistatidae have not previously been der incident light with a camera lucida described in the fossil record, although Eskov attached to an Olympus SZH stereomicro- (1990) mentioned a fossil specimen from the scope and photographs were taken with a Ni- Upper Jurassic of Kazakhstan. The current kon DIX digital camera attached to a Wild evolutionary tree (including the new fossil de- M8 stereomicroscope then manipulated in scribed in this paper) of the Haplogynae (Fig. Adobe Photoshop. 1) predicts the presence of Filistatidae in the Abbreviations used in the text and fig- — fossil record back to the Upper Cretaceous. ures.- In the leg formula (e.g. 1423), the legs However, this is a youngest age prediction are ranked in order of length (longest first). based on the presence of Ooeopidae and Se- Abbreviations are as follows: cx = coxa; cy gestriidae in amber from New Jersey (Penney = cymbium; e = embolus; fe = femur; h = 2002a, 2004). Here, the first fossil Filistatidae haem,olymph; la = labium; mt == metatarsus; is described in the genus Misionella, from op = opisthosoma; pa = patella; pi = paraem- Miocene (15-20 Ma; e.g, Iturralde-Vineet & bolic lamina; pom = leg segment present but MacPhee 1996) Dominican Republic amber. not measurable; pp = pedipalp; rex = retro- lateral excavation; s = long setae; sp = spine; METHODS spr = spinneret region; st = sternum; ta — Preservation,—The spider is preserved tarsus; ti = tibia; 1-4 = walking legs 1-4. close to the surface of a clear yellow, tear- SYSTEMATIC PALEONTOLOGY shaped piece of Dominican Republic amber 4 cm long X 1.7 cm wide; for details oflocality Family Filistatidae Ausserer 1867 and stratigraphy see Iturraide-Vineet & Subfamily Prithinae Gray 1995 MacPhee (1996). The spider is best observed Misionella Ra—mirez & Grismado 1997 in ventral view. There are two partial insect Type species, Filistata mendensis Mello- legs and the associated thoracic stereite in the Leitao 1920 by original designation. 93 94 THE JOURNAL OF ARACHNOLOGY L<U U<J Q UJ i5u<j % ^Q < L<JJ ^ < Sym Q5O “<gaUJ H= y^<QLy3LU UoOmJ ^^“^9 L<Q~U L<Q“U ^QQ1^ ^QC<O UQwJUCJe_OOJn9jhQ- D>Xf- oh-^UJE<hP-iP O _CD X ^0 UJ o ^ o a Age(Ma) u. O O .Q CO a. Q CL O U_JJ UhJ- CO CO Q DRA/*SSC MA B/BA AeP NJA 135 o oN 0rco) O >Oo.) Q. X<S 180 KEY 205 rangeextension based on phylogeneticrelationships range based on published record ofdescribed specimen(s) describedfossilspiders hypothesized relationships ghostlineage based on phylogenetic relationships 250 — — PENNEY—FOSSIL FILISTATIDAE IN DOMINICAN AMBER 95 — Distribution. Recent species in Brazil wide, fine detail of anal tubercle, spinnerets and Argentina, fossil species in Dominican and cribellum not clear, situated ventrally, ad- Republic amb—er. vanced from the posterior margin (Figs. 2, 3). Remarks. Misionella was erected as a Leg formula 1423; leg 1 cx 0.33, fe 1.86, monotypic genus by Ramirez & Grismado ti 2.13, mt 1,64, ta 1.00; leg 2 cx 0.30, fe 1.34, (1997) for Fiiistata mendensis from Brazil ti 1.47, mt 0.70, ta 0.60; leg 3 cx 0.30, fe 0.93, and Argentina. Grismado & Ramirez (2000) ti 0.93, mt0.83, ta 0.43; leg 4 cx 0.33, fe 1.36, described a second species from Brazil. ti pnm, mt LOO, ta not preserved; mt 2 with a retrolateral excavation in the distal halfbear- Misionella didicostae new species ing a single prominent retrolateral spine (Figs. Figs. 2-5 2-4); mt 1 with a single long, ventral terminal Material examined. ^Holotype, adult spine; remaining leg segments without spines. male in Dominican Republic amber, held in All leg segments, opisthosoma and sternum the Museo del Ambar Dominicano, Puerto with clearly visible, long setae. Pedipalp fe Plata, Dominican Republic. long, ti globose without projections, cymbium Diagnosis.' Misionella didicostae can be vestigial and partially fused to tegulum, em- distinguished from the two known extant spe^ bolus spirally twisted distally which extends cies by having a spiralled embolus distally ex- well beyond the paraembolic lamina (Fig. 5). tending well beyond the paraembolic lamina. Female.—Unknown. In addition, it lacks small spinules in the re- Distribution and age.—Dominican Repub- trolateral excav—ation of metatarsus 2. lic amber; Miocene (see Iturralde-Vinent & Etymology.' ^The specific epithet is a ma- MacPhee [1996]). tronym in honor of Mrs Ada (Didi) Benelli Remarks.—This specimen conforms with Costa who co-foeeded the Museo del Ambar the diagnostic characters of the genus given Dominicano, Puerto Plata, Dominican Repub- by Ramirez & Grismado (1997). Ramirez & lic in 1982, based on collections amassed over Grismado (1997) performed a cladistic anal- 33 years. ysis of the filistatid genera, which are placed Description of holotype.^Body length in two subfamilies, Filistatinae and Prithinae. 2.21; carapace 0.84 long, width not measur- Only males of the genera Misionella Ramirez & able; ocular region slightly raised, with eight Grismado 1997 and Pikelinia Mello-Leitio closely grouped eyes. Clear views of the eye 1946 have the metatarsus of the second leg arrangement and carapace shape are not af- modified, presumably to perform a clasping forded by the specimen, but there are no ob- function during copulation. Extant species of vious differences between the details visible both these genera are known only from South and the figure ofthe type species provided by America and the Galapagos Islands (Grismado Ramirez & Grismado (1997: fig. 96). Only & Ramirez 2000; Ramirez & Grismado 1997; distal tips of chelicerae visible: weak, unmod- Mtiller 1987). These genera are distinguished ified with a small fang. Labium longer than from one another by the structure of the male broad, with curved sides converging to form palpal tibia. Pikelinia has a prominent dorsal a pointed tip; fused to sternum. Maxillae lon- retrolateral projection, whereas Misionella ger than wide, convergent. Sternum subcir- does not (Ramirez & Grismado 1997). The cular, width 0.40, length offused sternum and new fossil species differs from the two known labium 0.77. Opisthosoma 1.21 long, 0.54 extant species ofMisionella in the structure of Figure L—Evolutionary tree oftheHaplogynae. DRA = DominicanRepublic amber(Wunderlich 1988; Penney 2001, this paper); SSC = Shanwang, Shandong, China (Dysderidae only; Zhang et al. 1994); MA = Mexican amber(Petrunkevitch 1971); B/BA = Baltic andBitterfeldambers (Menge 1854;Petrankevitch 1950; Wunderlich 1981, 1991, 1993); AeP = Aix en Provence (Gourret 1888); NJA = New Jersey amber (Penney 2002a, 2004). Cladogram follows Coddington & Levi (1991); for details of evolutionary tree construction and terminology see Smith (1994). Not illustrated: Segestriidae describedby Hickman (1957) and Oonopidae from Japan and Kenya (Nishikawa 1974 and Wunderlich 1981 respectively) may beextant species in Recent copal (e.g, Poinar 1992). 96 THE JOURNAL OF ARACHNOLOGY PENNEY^FOSSIL FILISTATIDAE IN DOMINICAN AMBER 97 the pedipalp= In M. mendensis (Mello=Leitao MacPhee (1999) proposed that during the Eo- 1920) the embolus is not spiralled but has cene-Oligoceee transition, the developing only a slight kink and it does not extend be^ northern Greater Antilles and northwestern yoed the paraembolic lamina; M. jaminawa South America were briefly (33-35 Ma) con- Grismado & Ramirez 2000 has a hook-shaped nected by a landspae (a subaerial connection embolus and lacks a paraembolic lamina. In between a continent and one or more off-shelf addition, both extant species have small spi- islands) centered on the emergent Aves Ridge. Eules in the retrolateral excavation of mt2; This landspan consisted of a series of large, these are absent in the new fossil species. closely spaced islands or possibly a continu- Based on the pedipalp morphology the new ous peninsula stretching from northern South species is more closely related to M. menden- America to the Puerto RicoA/irgin Islands sis. Block (Iturralde-Vinent & MacPhee 1999). DISCUSSION The massive uplift that apparently permitted these connections was finished by 32 Ma The fossil and extant Hispaeiola&n spider (Iturralde-Vinent & MacPhee 1999). The faunas were reviewed by Penney Perez- Greater Antilles in their current guise are rel- Gelabert (2002). The only known filistatid atively young geographical features, probably from the island was the extant Kukulcania no older than the middle Miocene. Therefore, Mbernalis (Hentz 1842), which is most cer- all oe-islaed lineages forming the Recent fau- tainly an introduced species (Penney 1999). na must be younger than Middle Eocene (Itur- Based on this observation, Penney (1999) hy- ralde-Vinent & MacPhee 1999). The known pothesized that Filistatidae may not have been distribution of Recent and fossil species of present on Hispaniola during the Miocene Misionella supports the Greater Antillean- when the amberwas formed, and thatthe fam- iTlhyehdaidsccoovleornyizoefdthteheneiswlafnodssimlorfaelsirfeiceesnttlhyi.s rSaoludteh-VAimneenrtic&a MlaancdPshpeaen h(y1p99o9t)h,esriasthoefr Itthuarn- the Greater Antillean-Cefitiai .kmericae land hypothesis and undiscovered extant species of connection hypothesis of Petrunkevitch Misionella may also exist on the island today. This new family record for the fossil fauna (1928). & brings the number of families recorded from Roth Roth (1984) defined autospasy as named species in Dominican Republic amber the separation of appendages, segments or to 36 and the total number offamilies record- parts thereof at a predetermined locus of ed to 45 (see Penney & Perez-Gelabert 2002). weakness when the appendages or segments Misionella was formerly only recorded are restrained by any external (not self-in- from extant species in Brazil and Argentina duced) source. Entrapment in amber would (Ramirez & Grismado 1997) although it prob- constitute such a source. Autospasy occurs ably has a wider distribution in South Amer- rarely in haplogynes but is known to occur in ica. It is not included in species lists for Pan- Filistatidae at the patellar-tibia joint. In the ama (Nentwig 1993), Costa Rica (Vega 1980), fossil, right legs 1 and 4 and left legs 1 and 3 Cuba (Alaybn-Garcia 2000) and is notrecord- have autospasized at this point and there are ed from Noith America (Ffiamick 2003). Pe- what I interpret as haemolymph exudations trunkevitch (1928) considered the GreaterAn- from the patella of right leg 1 and possibly tillean spider fauna to represent an eastern also left leg 1 preserved in the amber (Figs. outgrowth of the Central American fauna by 2, 3), which indicates it had happened shortly way ofapresumed earlierlandconnection and after becoming entrapped in the resin. Indeed, subsequent continent-island vicariaece. How- the first pair oflegs are preserved in the same ever, such a land connection appears never to piece of amber close to the spider, however & have existed (Ross Scotese 1988; Iturralde- they have rotated around approximately 180 Vieent & MacPhee 1999). Iturralde-Vinent & degrees (Figs. 2, 3), This observation suggests — Figure 2. Misionella didicostae new species. Holotype male, DominicanRepublic amber. Ventralview of whole specimen. Scale line 1.0 mm. 98 THE JOURNAL OF ARACHNOLOGY — Figure 3. Misionella didicostae new species. Holotype male, Dominican Republic amber. Camera lucida drawing of whole specimen. Scale line 1.0 mm. Refer to text for abbreviations. PENNEY—FOSSIL FILISTATIDAE IN DOMINICAN AMBER 99 — Figures 4—5, Misionella didicostae new species. Holotype male, Dominican Republic amber. 4, meta- tarsus 2. 5, pedipalp. Scale line 0.5 mm. Refer to text for abbreviations. that the spider was engulfed in a flowing resin Gourret, M.P. 1888. Recherches sur les arachnides seep of relatively low viscosity, rather than Tertiaires d’Aix en Provence, Recueil Zoolo- having wandered onto a sticky exudate, be- gique Suisse 1888:431-496. coming stuck and then covered by a subse- Gray, M.R. 1995. Morphology and relationships quent resin flow, as is known to have occured within the spider family Filistatidae (Araneae: Araneomorphae). Records of the Western Aus- in many cases of invertebrate amber preser- tralian Museum, Supplement 52:79-89. vation (Penney 2002b). Although Recent spe- Grismado, C.J. & M.J. Ramirez. 2000. Notes onthe cies ofMisionella are primarily synanthropic, spider genus Misionella with a description of a some have been collected from tree trunks new species from Brazil (Araneae: Filistatidae). (Ramirez & Grismado 1997). The discovery Studies on Neotropical Fauna and Environment of this genus in amber is consistent with eco- 35:161-163. logical observations of its Recent relatives. Hentz, N.M. 1842. Descriptions and figures of the araneides ofthe United States. Boston Journal of ACKNOWLEDGMENTS Natural History 4:54-57,223-231. I thank Michele Costa, Ada (Didi) Benelli Hickman, V.V. 1957. A fossil spider from Tertiary Costa and Annie Del Rosario for their hospi- retinite at Allendale, Victoria. Proceedings ofthe Royal Society of Victoria 69:25-27. tality, including provision ofresearch facilities Iturralde-Vinent, M.A. & R.D.E. MacPhee. 1996. and access to the collections held in the Mu- Age and paleogeogeographical origin ofDomin- seo del Ambar Dominicano, Puerto Plata, Do- ican amber. Science 273:1850-1852. minican Republic. I thank M. Ramirez, Uni- Iturralde-Vinent, M.A. & R.D.E. Macphee. 1999. versity of Buenos Aires for reprints, R A. Paleogeography of the Caribbean region: impli- Selden, University ofManchester for his com- cations for Cenozoic biogeography. Bulletin of ments on the manuscript and I acknowledge a the American Museum of Natural History 238: Leverhulme Trust grant. 1-95. Mello-Leitao, C.F. 1920. Algumas aranhas novas. LITERATURE CITED Revista Sociedade Brasileira de Ciencia 3:169- Alayon-Garcia, G. 2000. Las aranas endemicas de 176. 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Advances in the taxonomy ofspi- tectonic model ofthe GulfofMexico and Carib- ders in Miocene amber from the Dominican Re- bean region. Tectonophysics 155:139-168. public (Arthropoda, Araneae). Palaeontology 44: Roth, V.D. & Roth, B.M. 1984. A review ofappen- 987-1009. dotomy in spiders and other arachnids. Bulletin Penney, D. 2002a. Spiders in UpperCretaceousam- ofthe British Arachnological Society 6:137-146. ber from New Jersey (Arthropoda:Araneae). Pa- Smith, A.B. 1994. Systematics and the Fossil Re- laeontology 45:709-724. cord: documenting evolutionary patterns. Black- Penney, D. 2002b. P—aleoecology ofDominican am- well Science, Oxford. ber preservation spider (Araneae) inclusions Vega, C.M.Z. 1980. Lista anotada de especies de demonstrate abias foractive, trunk-dwellingfau- arahas de Costa Rica. Brenesia 18:301-352. nas. Paleobiology 28:389-398. Wunderlich, J. 1981. Fossile Zwerg-Sechsaugen- Penney, D. 2004. 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Entomologische Museum of Comparative Zoology. Bulletin of Zeitschrift 101:21-25. the Museum of Comparative Zoology 103:259- Wunderlich J. 1993. Dieersten fossilen Speispinnen 337. (Fam. Scytodidae) im Baltischen Bernstein Petrunkevitch, A. 1971. Chiapas amber spiders 2. (Arachnida, Araneae). Mitteilungen aus dem University of California Publications in Ento- Geologisch-Palaontologischen Institut der Univ- mology 63:1-44. ersitat Hamburg 75:243-247. Platnick, N.I. 2003. The world spider catalog, ver- Zhang J., B. Sun & X. Zhang. 1994. Miocene in- sion 3.0. American Museum of Natural History, sects and spiders from Shanwang, Shandong. online at http://research.amnh.org/entomology/ Science Press, Beijing. spiders/catalogS1-87/index.html Platnick, N.I., J.A. Coddington, R.R. Forster& C.E. Manuscript received 19 June 2003, revised 17No- Griswold. 1991. Spinneret morphology and the vember 2003.

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