new synonymies and combinations for orb-weaving

Zootaxa 2073: 22–30 (2009)www.mapress.com/zootaxa/Copyright © 2009 · Magnolia PressArticleISSN 1175-5326 (print edition)ZOOTAXAISSN 1175-5334 (online edition)Not from “Down Under”: new synonymies and combinations for orb-weavingspiders (Araneae: Araneidae) erroneously reported from AustraliaVOLKER W. FRAMENAU 1,2,5 , NIKOLAJ SCHARFF 3 & HERBERT W. LEVI 41Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia, 6986, Australia.E-mail: volker.framenau@museum.wa.gov.au2School of Animal Biology, University of Western Australia, Crawley, Western Australia 6009, Australia3Natural History Museum of Denmark, Zoological Museum, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen,Denmark: E-mail: nscharff@snm.ku.dk4Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA. E-mail: levi@fas.harvard.edu5Corresponding authorAbstractThe examination of type material of presumed Australian orb-weaving spiders as part of a revision of the Araneidae ofthis country revealed that a number of species are not from Australia. The Natural History Museum, Vienna (Austria)holds the type material of three species of orb-weaving spiders that were originally described from Australia, however allof the species are undoubtedly of American, most likely southern Brazilian, origin and it is unlikely that they werecollected in Australia. We propose the following synonymies and generic transfers: Acacesia tenella (L. Koch, 1871)comb. nov. (= Acacesia cornigera Petrunkevitch, 1925 new synonymy); Alpaida navicula (L. Koch, 1871) comb. nov.(= Alpaida roemeri (Strand, 1908) new synonymy); and Eustala mucronatella (Roewer, 1942) comb. nov. In addition,Novearanea queribunda (Keyserling, 1887) comb. nov. (= Araneus quaesitus (Keyserling, 1887) new synonymy; =Novaranea laevigata (Urquhart, 1891) new synonymy) is a New Zealand orb-weaving spider based on the labels thatwere found with the type specimens housed at the Natural History Museum, London (England). In the originaldescription no locality data was given for N. queribunda and “Australien” was erroneously listed for A. quaesitus.Key words: New Holland, Neuholland, Brazil, Nearctic, New Zealand, new combinationIntroductionColonialism by the Netherlands from the 17 th into the 20 th century resulted in many localities world-widebearing names of Dutch origin. New Holland (initially “Nova Hollandia”) is a historic name for Australiagiven by the seafarer Abel Tasman who named it after the Dutch province of Holland (Sharp 1968). Butbetween 1624 and 1664 parts of northeastern Brazil (Rio Grande do Norte to Bahia states) were also referredto as “Nova Holanda” and localities in Acre and Rondônia and many hotels and lodgings in northeasternBrazil are still named Nova Holanda (with various spellings). Today there is also a municipality, VilaHolanda, in São Paulo and a favela, Nova Holanda, in Rio de Janeiro (E. Buckup personal communication).There is also a village in Santa Catarina, southern Brazil, with the name Nova Hollandia (A.A. Lise personalcommunication) and there are at present six New Holland post offices in the United States.Consequently, historic specimens simply labeled “New Holland” (“Neuholland” in German) are difficultto correctly associate to a locality without geographic reference. Two of three species of orb-weaving spiders,of which the type material is lodged at the Natural History Museum in Vienna, were included in historicaccounts of the Australian fauna based on their collection labels reading “Neuholland”, Araneus tenellus (L.Koch, 1871) and Araneus mucronatellus (Roewer, 1942) (originally described as Epeira mucronata L. Koch,1871) (L. Koch 1871). A third species, Araneus navicula (L. Koch, 1871), was also thought to be from22 Accepted by G. Hormiga: 7 Mar. 2009; published: 16 Apr. 2009

Australia based on the original locality label “Sidney”, mistaken for the Australian settlement Sydney. Recentexamination of the type material of these species revealed that they are not of Australian origin, butundoubtedly belong to the American fauna.Keyserling (1887) described two species of Araneidae, Araneus queribundus (Keyserling, 1887) andAraneus quaesitus (Keyserling) which in past and current catalogs (Roewer 1942; Bonnet 1955; Platnick2009) are listed as Australian. The type material of both species is lodged at the Natural History Museum,London, and is clearly labeled “New Zealand”. We were able to identify both species as belonging to the NewZealand genus Novaranea Court & Forster, 1988.MethodsSpiders were examined in ca. 70% ethanol under different Leica stereomicroscopes. Images of MNHW typeswere taken in different focal planes with a Nikon D300 digital SLR camera attached via a C-mount adapterfrom LM-Scope (http://www.lmscope.com/index_e.html) to a Leica M16A stereomicroscope and combinedwith Auto Montage (vers. 5.02) software from Syncroscopy to increase depth of field. We used 2 Nikon R1C1wireless speedlights instead of fiber optics to illuminate the exposures. The latter were used as guide-light forfocusing. The photograph of the male syntype of Epeira queribunda was taken with a Canon G6 digitalcamera through the ocular tube of the microscope provided by the BMNH.AbbreviationsAMNH American Museum of Natural History, New York, USABMNH The Natural History Museum, London, EnglandMNHN Muséum national d’Histoire naturelle, Paris, FranceNHMW Naturhistorisches Museum, Vienna, AustriaPMY Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USASMF Naturmuseum und Forschungsinstitut Senckenberg, Frankfurt/Main, GermanyTaxonomyAcacesia tenella (L. Koch, 1871) comb. nov.Figs 1A–D, 2A–EEpeira tenella L. Koch, 1871: 76–78, pl. 6, figs 5, 5a, 6, 6a; Butler 1876: 352.Aranea tenella (L. Koch). Roewer 1942: 834.Wixia tenella (L. Koch). Archer, 1951: 15.Araneus tenellus (L. Koch). Rainbow 1911: 194; Bonnet 1955: 609.Acacesia cornigera Petrunkevitch, 1925: 117, figs 32–41; Glueck, 1994: 73–75, figs 2–3, 9–16. New synonymy.Types. Syntypes of Epeira tenella L. Koch, 1871: 1 male, 5 females, 1 penultimate female, 1 juvenile, listedin L. Koch (1871) from “Neuholland” (NHMW 1826.II.11) (examined).Syntypes of Acacesia cornigera Petrunkevitch, 1925: male, female, juvenile, San Lorenzo, Panama (PMY)(examined).Remarks. Ludwig Koch (1871) listed 1 male and 9 females as syntypes in his original description ofEpeira tenella (p. 78, “Neun entwickelte Weibchen und ein Männchen von Neuholland ohne nähere Angabeim k. k. Museum zu Wien.”), but the vial only contained the 8 specimens listed above. This also coincideswith a copy of the NHMW register which lists 8 specimens. It appears that L. Koch (1871) erroneously listed9 females and also misinterpreted the developmental stage of some of the spiders.NEW SYNONYMIES FOR ORB-WEAVING SPIDERSZootaxa 2073 © 2009 Magnolia Press · 23

FIGURE 1. Syntype male and female of Acacesia tenella (L. Koch, 1871) comb. nov. A, male, dorsal view; B, maleright pedipalp, mesal view; C, female, dorsal view; D, epigyne, ventral view. Scale bar: A, C, 1 mm; B, D, 0.1 mm.Acacesia cornigera was reported as widespread from central Mexico to southern Brazil and is found onboth sides of the Andes (Glueck 1994). The species has quite variable male pedipalps and the posterior viewof the epigynum is generally heavily sclerotised and sometimes covered with debris and therefore difficult toexamine; the female illustrated here has apparently recently moulted with less sclerotised epigyne (Figs 1D,2B). All males from Mexico to Brazil have similar cone-shaped emboli, although it may be hidden in part bythe conductor. The paracymbium is variable, Central American specimens have two distal prongs, theproximal short, the South American generally have only one (Fig. 2D). However, there is little doubt that thesyntypes of Epeira tenella and those of Acacesia cornigera represent the same species and both aresynonymised here as Acacesia tenella (L. Koch, 1871) comb. nov.24 · Zootaxa 2073 © 2009 Magnolia PressFRAMENAU ET AL.

Based on the locality label “Neu-Holland” L. Koch (1871) erroneously assumed that the spiders werecollected in Australia. It is possible that the spiders were collected in the Dutch colony of the same name innortheastern Brazil, although this colony had long seized to exist when the specimens were given to theMNHW in 1826.FIGURE 2. Syntype female and male of Acacesia tenella (L. Koch, 1871) comb. nov. A, female, dorsal view; B,epigyne, ventral view; C, epigyne, posterior view; D, paracymbium of left male pedipalp, dorsolateral view, E, male leftpedipalp, mesal view. Scale bar: A, 1 mm; B–E, 0.1 mm.Alpaida navicula (L. Koch, 1871) comb. nov.Figs 3A–BEpeira navicula L. Koch, 1871: 102–103, pl. 8, figs 4, 4a, 4b.Aranea roemeri Strand, 1908: 3; Strand 1916: 109. New synonymy.Araneus naviculus (L. Koch). Rainbow 1911: 189; Roewer 1942: 830; Bonnet 1955: 548.Araneus roemeri Strand. Petrunkevitch 1911: 312; Mello-Leitão 1927: 400; Roewer 1942: 851; Bonnet 1955: 586.Alpaida roemeri (Strand). Levi, 1988: 469–470, figs 551–554.Types. Holotype of Epeira navicula L. Koch, 1871: female, labeled “Sidney” and “collected by Novara”(NHMW 1866.I.10) (listed in L. Koch (1871) from Sydney, 33º53’S, 151º13’E, New South Wales, Australia)(examined).Holotype Aranea roemeri Strand, 1908: female, Joinville [Joinvile] [26º19’S, 48º52’W, Santa Catarina,Brazil], W. Ehrhardt, 1906 (SMF 3380) (examined).Remarks. A comparison of the holotype of Epeira navicula with the holotype of Alpaida roemeriundoubtedly supports a synonymy of both species; we can not see any differences in somatic and genitalicNEW SYNONYMIES FOR ORB-WEAVING SPIDERSZootaxa 2073 © 2009 Magnolia Press · 25

morphology between the two specimens that warrant a recognition as separate species. We therefore transferAraneus naviculus to Alpaida, A. navicula (L. Koch, 1871) comb. nov. and consider it a senior synonym of A.roemeri. The species is currently only known from the holotype collected in southern Brazil (Levi 1988), andthe type locality “Sydney” published by L. Koch (1871) must be considered erroneous. The register of theMNHW lists “Sidney” as locality and “collected by Novara”, presumably referring to the Novara-Expedition(1857–1859), a circumnavigation of the globe by the Austrian Imperial war navy that visited both SouthAmerica and Australia (Treffer 1973).FIGURE 3; Holotype female of Alpaida navicula (L. Koch, 1871) comb. nov. A, dorsal view; B, epigyne, ventral view.Scale bar: A, 2 mm; B, 0.1 mm.Eustala mucronatella (Roewer, 1942) comb. nov.Figs 4A–HEpeira mucronata L. Koch, 1871: 74–76, pl. 6, figs 2, 3, 3a; Butler 1876: 352; Karsch 1878: 789; (preoccupied by Epeiramucronata Blackwall, 1862).Aranea mucronatella Roewer, 1942: 830 (replacement name).Araneus mucronatus (L. Koch). Bonnet 1955: 547.Types. Syntypes of Epeira mucronata L. Koch, 1871: 2 penultimate males, 4 females, listed in L. Koch(1871) from “Neuholland” (NHMV 1826.II.3) (examined).Description. Female bleached, only bare indication of pattern on abdomen (Figs 4A, B). Abdomenpointed posteriorly with two posterior humps (Fig. 4E). Eyes subequal in size. Anteriors 1.2 their diameterapart, 2.2 from laterals. Posterior median eyes their diameter apart, 2.0 from laterals. Total length 4.6 mm.Carapace 1.9 mm long, 1.7 mm wide. First Femur 2.2 mm; third metatarsus 1.4 mm (most leg articles lost).Remarks. Somatic and genital morphology (Figs 4A-H), in particular the shape of the epigynal scapewhich projects anteriad (Figs 4F–H) leave no doubt that Araneus mucronatellus is a member of the Americangenus Eustala to which we transfer it here: Eustala mucronatella comb. nov. The species is similar to Eustalataquara (Keyserling, 1892) described from Rio Grande do Sul, Brazil (see Levi 2007), but E. mucronatelladiffers by an additional abdominal hump between the posterior hump and the spinnerets, by a pointedepigynum scape (that of the type of E. taquara is blunt) and the shorter distance between the posterioropenings of epigyne (Figs 4F–H). Pending a revision of the genus Eustala and the examination of additionalspecimens, there is a possibility that E. mucronatella is a variable specimen of E. taquara.26 · Zootaxa 2073 © 2009 Magnolia PressFRAMENAU ET AL.

FIGURE 4. Syntype females of Eustala mucronatella (Roewer, 1942) comb. nov. A, dorsal view; B, another female,abdomen, dorsal view; C, abdomen, ventral view; D, epigyne, ventral view; E, abdomen, lateral view; F, epigyne, ventralview; G, epigyne, lateral view; H, another epigyne, ventral view. Scale bar: A–C, E, 1 mm; D, F–H, 0.1 mm.NEW SYNONYMIES FOR ORB-WEAVING SPIDERSZootaxa 2073 © 2009 Magnolia Press · 27

Novaranea queribunda (Keyserling, 1887) comb. nov.Fig. 5Epeira queribunda Keyserling, 1887: 195–197, pl. 17, figs 4, 4a, b, 5. 5a, b.Epeira quaesita Keyserling, 1887: 197–198, pl. 17, figs 6, 6a. New synonymy.Epeira laevigata Urquhart, 1891: 171–172, pl. 21, fig. 6; Urquhart 1892: 227. New synonymy.Araneus quaesitus (Keyserling). Rainbow 1911: 191; Bonnet 1955: 580.Araneus queribundus (Keyserling). Rainbow 1911: 191; Bonnet 1955: 580.Aranea laevigata (Urquhart). Roewer 1942: 829.Aranea queribunda (Keyserling). Roewer 1942: 832.Aranea quaesita (Keyserling). Roewer 1942: 832.Araneus laevigatus (Urquhart). Bonnet 1955: 526.Novaranea laevigata (Urquhart). Court & Forster 1988: 114–115, figs 361, 539–552.Types. Lectotype (here designated) of Epeira queribunda Keyserling, 1887: male, New Zealand [no exactlocality] (BMNH 1890.7.1.4134) (examined).Paralectotype of Epeira queribunda Keyserling, 1887: female, same data as lectotype (BMNH1890.7.1.4135) (misidentification; this specimen is Colaranea verutum (Urquhart, 1887) (examined).Holotype of Epeira quaesita Keyserling, 1887: female, New Zealand [no exact locality], BradleyCollection (BMNH 1890.7.1.4163) (examined).Holotype of Epeira laevigata Urquhart, 1891: female, forest near Stratford [39º20’S, 174º17’E, NewZealand], A.T. Urquhart (lost; Court & Forster 1988) (not examined).Remarks. In his original description, Keyserling (1887) did not mention a type locality for Araneusqueribundus but Rainbow (1911, p. 191) listed “Hab.–Australia” for this species. This was later perpetuatedas distribution by Roewer (1942) and Bonnet (1955). Our examination of the syntype male and female in theBMNH revealed that the specimens are clearly labeled “New Zealand” which was omitted in Keyserling’s(1887) description. Comparison with New Zealand Araneidae illustrated by Court and Forster (1988) showedthat both male and female syntype of Epeira queribunda represent two different species. To providenomenclatural stability for the species concept of E. queribunda, the male is here designated as lectotype. Themale pedipalp morphology, in particular the shape of the terminal apophysis of this species, is unique withinthe New Zealand Araneidae and clearly the same as that of Novaranea laevigata (Urquhart, 1891), the typespecies of the monotypic genus Novaranea Court & Forster, 1988 (Fig. 5; see also Court & Forster 1988, figs546–547). Therefore Araneus queribundus is here transferred to Novaranea and considered a senior synonymof Novaranea laevigata: Novaranea queribunda (Keyserling, 1887) comb. nov. The female syntype(paralectotype) of Epeira queribunda is Colaranea verutum (Urquhart, 1887).Examination of the holotype of Epeira quaesita revealed through the original label, that this specimen wasalso collected in New Zealand, and not in Australia as originally stated by Keyserling (1887, p. 198:“Australien, Sammlung Bradley”). The morphology of the epigyne clearly conforms to that of Novaraneaqueribunda comb. nov. as illustrated in Court & Forster (1988, sub N. laevigata; see above) who also state(pg. 115): “The loss of the type [of Epeira laevigata] is fortunately of little importance because Urquhartillustrated the epigynum and this cannot be confused with any other New Zealand species.” Consequently,Araneus quaesitus is here considered a junior synonym of N. queribunda.Our exhaustive examination of Australian Araneidae did not reveal any specimens of N. queribundacollected in Australia and we do not consider this genus and species part of the Australian fauna.28 · Zootaxa 2073 © 2009 Magnolia PressFRAMENAU ET AL.

FIGURE 5. Lectotype male of Novaranea queribunda (Keyserling, 1887) comb. nov., male pedipalp, retrolateral view.Scale bar: 0.2 mm.AcknowledgementsWe are particularly indebted to Janet Beccaloni (BMNH) and Christoph Hörweg (NHMW) who facilitatedthis study either by assisting when visiting their institution or by the loan of material. David Court (Singapore)confirmed the synonymy of A. queribundus and A. quaesitus with N. laevigata based on the photographs weprovided to him. We thank Jagoba Malumbres-Olarte and Cor Vink (both Lincoln, New Zealand) forproviding photographs of an expanded N. queribunda pedipalp to clarify the terminology of its sclerites. Wethank Gustavo Hormiga and an anonymous reviewer for their constructive suggestions that improved thismanuscript considerably. This study was financially supported by the Australian Biological Resources Study(ABRS) (grant no. 205–24) to VWF and NS and by the Danish Agency for Science, Technology andInnovation to NS (grant no. 272-05-0431).ReferencesArcher, A.F. (1951) Studies in the orbweaving spiders (Argiopidae). 1. American Museum Novitates, 1487, 1–52.Bonnet, P. (1955) Bibliographia Araneorum. Tome II (part 1: A–B). Douladoure, Toulouse, pp. 1–918.Butler, A.G. (1876) On a small collection of Arachnida from Queensland, with descriptions of three apparently newspecies. Cistula Entomologica, 1, 349–354.Court, D.J. & Forster, R.R. (1988) Family Araneidae. Subfamily Araneidae [sic]. Otago Museum Bulletin, 6, 70–124.Glueck, S. (1994) A taxonomic revision of the orb weaver genus Acacesia (Araneae: Araneidae). Psyche, 101, 59–84.Karsch, F. (1878) Exotisch-araneologisches. Zeitschrift für die gesamten Naturwissenschaften, 51, 322–333, 782–826.Keyserling, E. (1987) Die Arachniden Australiens, nach der Natur beschrieben und abgebildet. (Theil 2, 3. Lieferung).Bauer & Raspe, Nürnberg, pp. 153–202.Koch, L. (1971) Die Arachniden Australiens, nach der Natur beschrieben und abgebildet. (Theil 1, 1. Lieferung). Bauer& Raspe, Nürnberg, pp. 1–104.Levi, H.W. (1988) The Neotropical orb-weaving spiders of the genus Alpaida (Araneae: Araneidae). Bulletin of theMuseum of Comparative Zoology, 151, 365–487.Levi, H.W. (2002) Keys to the genera of araneid orbweavers (Araneae, Araneidae) of the Americas. Journal ofNEW SYNONYMIES FOR ORB-WEAVING SPIDERSZootaxa 2073 © 2009 Magnolia Press · 29

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