Mesothele spiders in the Museum für Naturkunde Berlin

An annotated catalogue of the rare mesothele spiders (Araneae: Mesothelae: Liphistiidae) held in the Museum für Naturkunde Berlin is presented. The museum hosts non-type specimens of nine species representing all three currently recognised genera, namely: Liphistius desultor Schiödte, 1849, L. malayanus cameroni Haupt, 1983, L. cf. thaleban Schwendinger, 1990, Heptathela kikuyai Ono, 1998, H. kimurai (Kishida, 1920), H. yanbaruensis Haupt, 1983, Ryuthela ishigakiensis Haupt, 1983, R. nishihirai (Haupt, 1979) and R. tanikawai Ono, 1997 (spec. reval.). The geographical focus of this collection is Malaysia and Japan, and most of the material was collected by the Berlinbased zoologist Joachim Haupt.

Mesothelae is a fairly small suborder of spiders (currently 87 species in three genera, Platnick 2014) which are nevertheless of considerable phylogenetic interest.On first appearance they resemble mygalomorph spiders ("tarantulas", etc.), but in fact they are widely accepted as the most basal spider lineage retaining plesiomorphic characters such as a segmented opisthosoma bearing spinnerets near the middle of the underside.The latter character is the source of the name 'meso' -'thelae'.All other spiders have their spinnerets located at or close to the rear of the opisthosoma and are grouped in the suborder Opisthothelae.Fossil data indicate that mesotheles -or at least similar-looking spiders with a segmented opisthosoma and similar carapace and eye morphology -were found across Euramerica during the late Carboniferous.For a recent account of new fossils, which also drew on the material documented here for comparative purposes, see Selden et al. (in press).Today the group is restricted to eastern Asia (see below).
Living mesotheles are medium to large-sized spiders which construct a burrow covered by one or two trap-doors.Up to ten silken 'trip-wires' radiate from the burrow entrance.The spider lurks inside the burrow and is alerted to prey touching the silk threads which effectively act as a sort of 'proto-web'.A detailed account of mesothele anatomy and biology can be found in Haupt (2003).
The Museum für Naturkunde in Berlin (MfN) hosts a small, but significant collection of these quite rarely collected spiders.As the first of a planned series of papers documenting the spider collections of this museum -particularly groups not covered previously by the Berlin type catalogues by Manfred Moritz and Sophie-Charlotte Fischer (e.g.Moritz & Fischer 1990, Moritz 1992) -we present an annotated catalogue of the Mesothelae holdings.
Much of the mesothele material of the MfN was assembled by the Berlin-based zoologist Joachim Haupt (Fig. 1), formerly of the Free-University Berlin and later of the city's Technical University, who died in April 2013.As well as studying arthropod groups such as myriapods, mites, hexathelid spiders and whip scorpions -with a particular focus on micro-morphology and ultrastructure -Joachim Haupt worked extensively on the biology and systematics of mesothele spiders (Haupt 1977, 1979, 1981, 1982, 1983, 1984, 1986, 1990, 1993, 2002, 2003, Haupt & Traue 1986, Haupt & Kovoor 1993).He also had strong links to Japan, where he often collected as can be seen from the specimen labels and associated dates.We dedicate this present work to his memory.It should be noted that -contrary to published data -the types of his new species were not deposited in the MfN (formerly in East Berlin), having been described at the time when Berlin was still a divided city.Other spider specimens collected by Joachim Haupt can be found in the Zoological Museum of the University of Hamburg and in the zoological collections of the University of Rostock (both Germany), but the precise wherabouts of some type material remains equivocal.See below for notes on individual species.

Material and methods
All specimens listed here are stored in the wet collections (in 70 % alcohol) and have all been given ZMB (for Zoologisches Museum Berlin) repository numbers, which is the traditional acronym for the MfN.The data will be added to the database Systax.Some changes in nomenclature proved necessary to reflect the recognition of certain subspecies as distinct species in more recent publications.Individual case studies are discussed below and the specimen labels are amended accordingly.During the course of this work we also realised that a number of locality names were either incorrect or had at least been wrongly transcribed from the original labels.These have all been corrected here.

Results
Order Araneae Clerck, 1757 Suborder Mesothelae Pocock, 1892 Family Liphistiidae Thorell, 1869 Subfamily Liphistiinae Thorell, 1869 Remarks: According to authors such as Schwendinger & Ono (2011, and references therein), living mesotheles can be divided into two subfamilies: the South-East Asian Liphistiinae and the East Asian Heptathelinae.The latter subfamily was considered in some schemes -particularly Haupt (1983) -to be a distinct family, Heptathelidae, but see Raven (1985) for counterarguments.

Liphistius Schiödte, 1849
Liphistius desultor Schiödte, 1849 (Fig. 2a) Material: ZMB 10074, 1(; "Pulu Pinang" [Penang Island], Peninsular MALAySIA; date uncertain, "Kauf Roesen 27.3.47"[purchased from Roesen].ZMB 48315, "Falltür" [trapdoor]; Teluk Bahang, Penang Island, Peninsular MALAySIA, leg.J. Haupt, 2.III.1981.Remarks: This species -the first mesothele to be described (Schiödte 1849) -is restricted to Peninsular Malaysia.There is no further data about whom ZMB 10074 was purchased from.The locality is, in both cases, most likely Penang Island since the literal translation of the locality of the label is "areca palm island" (Schwendinger pers. comm.).This species was also reported from the nearby mainland (Platnick et al. 1997).Haupt, 1983 Material: ZMB 48532, 2 juv.; "Berinchan" [Brinchang or Berincang, Peninsular MALAySIA]; leg.Haupt, 16.IV.1984.Remarks: Also endemic to Peninsular Malaysia; as its name implies, this subspecies occurs in the Cameron Highlands.The holo-and paratypes of this species were explicitly noted as having been deposited in the Zoologisches Museum Hamburg (see Haupt 1983: 282), and as having been collected in 1981.Both specimens in Berlin are also from the type locality, but both are juveniles and since they postdate the original description, they cannot be part of the type series.
Genus Heptathela Kishida, 1923 Remarks: In his 1983 paper, Joachim Haupt regarded all Heptathela from Kyushu to Okinawa as part of a single species -a concept reflected in the original MfN labels -albeit recognising several subspecies: H. kimurai kimurai (Kishida, 1920) (type locality: Shiroyama, Kagoshima-shi), H. kimurai higoensis Haupt, 1983 (Kumamoto, North Kyushu), H. kimurai amamiensis Haupt, 1983 (Amami-oshima Island) and H. kimurai yanbaruensis Haupt, 1983 (Okinawa Island).Subsequently, Ono (2009) regarded all of these (plus some new taxa) as distinct species and this view has been adopted in the World Spider Catalog of Platnick (2014).Applying this current species concept to the Berlin material we now have voucher material from three Heptathela species.Note that Tanikawa's attempts to explain the species diversity of Japanese Heptathela (see Tanikawa et al. 2006; pers.comm. of A. Tanikawa to H. Ono) have not found favour;  (Kishida, 1920).c.Ryuthela nishihirai (Haupt, 1979).The latter two originated from the Joachim Haupt collection.
partly because of the large number of heterogeneous populations with a scattered distribution, but still often adjacent to one another.Ono, 1998 Material: ZMB 48317, 1 exuvia; Oita, Kyushu, JA-PAN; [leg.J. Haupt], VIII.2004. ZMB 48318, 48342, 48345-47, 5 specimens; Oita, Kyushu, JAPAN; leg.J. Haupt, 25.III.2004.Remarks: Originally labelled as H. kimurai, the geographical origin of these specimens -from Oita in Kyushu, the most southerly of Japan's four main Islands -implies that they should probably be H. kikuyai (sensu Ono 1998) which is common there; although we concede that several Heptathela species are known from this island (P.Schwendinger pers.comm.).The locality details are nonetheless a little vague as there is both an Oita Prefecture and a more specific locality of Oita-shi (= Oita City) in Kyushu.We assume the specimens come from somewhere in the wider Oita area.(Kishida, 1920) (Fig. 2b) Material: ZMB 48319, 1 specimen; Shiroyama, Kagoshima, Kyushu, JAPAN; leg.J. Haupt, date uncertain.ZMB 48341, 1 specimen; Kirishima, Kyushu, JAPAN; leg.J. Haupt, 23.III.2004. Remarks: Schwendinger & Ono (2011) noted that this species is known from several localities in southern Kyushu where the species is endemic.ZMB 48319 is topotypic; the type specimen also originated from Shiroyama.ZMB 48341 could be from the Kirishima Shrine at Kirishima-shi, from Kirishimashi (= Kirishima City) itself or from Mt. Kirishima situated on the border of Kagoshima and Miyazaki Prefectures.Haupt, 1983 Material: ZMB 48316, 1); JAPAN, locality uncertain; leg.J. Haupt, I.2006.ZMB 48320, 1 specimen; Iii (as "Jii"), Kunigami-son, Okinawa Island, JAPAN; leg.J. Haupt, 15.IV.1979.Remarks: These specimens were originally labelled Heptathela kimurai yanbaruensis, as per Haupt's (1983) original description.However, as noted above, Ono (2009) recognised H. yanbaruensis as a distinct species; see also Schwendinger & Ono (2011).The species is currently recorded as endemic to Okinawa in the Ryukyu Islands (cf.Platnick 2014) which form an island chain from Kyushu in the north towards Taiwan in the south.On the basis of the current data the distributional range of H. yanbaruensis can be restricted to the northern part of Okinawa Island (the yanbaru area), thus ZMB 48316 probably comes from this part of Okinawa too.The holo-and paratypes of this species were explicitly noted as having been deposited in the Zoologisches Museum Hamburg (cf. Haupt 1983: 284).The present material, collected in 2006, cannot thus be part of the type series.

Fig. 1 :
Fig. 1: Joachim Haupt (died 2013), photographed in 2008.He collected much of the mesothele material held in the Museum für Naturkunde in Berlin (MfN) and worked extensively on these spiders.Image courtesy of the European Society of Arachnology (http://www.european-arachnology.org/).