A catalogue of the scutigeromorph centipedes in the Museum für Naturkunde, Berlin

An annotated catalogue of the type and non-type scutigeromorph centipedes (Chilopoda: Scutigeromorpha) held in the Museum für Naturkunde Berlin is presented. The collection includes material from throughout the world, with a focus on former German colonies. Notes on collectors and localities are provided where appropriate. Type material for 42 Recent species or subspecies is present; all described in the early 20th century by Karl Verhoeff. However, only seven of these names remain valid with the other thirty-five currently regarded as junior synonyms. The collection in its entirety includes material from nineteen currently valid species, making up about 20% of the known world fauna. The type of a fossil species in Eocene Baltic amber is also listed for completeness.


Introduction
Scutigeromorphs (Chilopoda: Scutigeromorpha) -colloquially known as house centipedes -are distinctive, long-legged myriapods widely perceived as the earliest branching clade within the Chilopoda. In fact they are one of the most ancient terrestrial arthropod groups, with fossils known from the late Silurian (Shear et al. 1998). Today they are mostly found throughout warmer regions of the world, although Scutigera coleoptrata Linnaeus, 1758 can show a synanthropic distribution into colder regions. Approximately 95 valid species are currently recognized (Giribet and Edgecombe 2013), conventionally divided into three familes: the Neotropical-Afrotropical Pselliodidae, the Afro-Malagasy Scutigerinidae and the largely cosmopolitan Scutigeridae; the latter containing the bulk of the species. Phylogenetic relationships have been investigated by Edgecombe andGiribet (2006, 2009) and Giribet and Edgecombe (2013). Scutigeromorph centipedes tend to be fast-moving animals, their long legs strongly adapted for running after prey. General notes on their biology can be found in reviews such as Acosta (2003) and Edgecombe (2011), and references therein.
Scutigeromorph types in the Museum für Naturkunde, Berlin (MfN) were initially listed by Moritz and Fischer (1979), as part of their wider catalogue of centipede type material which recognized 319 types and 709 species across Chilopoda as a whole. The present work draws extensively on their efforts. Today, we have additional online resources -in particular the excellent ChiloBase 2.0 (Bonato et al. 2016) -which enable us to check the taxonomic status of both our type and non-type material. In what we hope will be a series of papers on the major centipede clades, this permits us to offer a full catalogue of the entire scutigeromorph collection with species placed according to the latest taxonomic opinions. Historical resources and online encyclopedias also mean we are now in a position to provide additional data and clarifications about type localities and historical notes on individual collectors which were not available to Moritz and Fischer. The Berlin collection is unusual in that all 42 Recent scutigeromorph type series stem from a single worker, Karl [Carl] Verhoeff, who described them in seven publications between 1904 and 1939. Among myriapodologists it is widely recognized that Verhoeff was a 'splitter', emphasizing minor differences between specimens and using these to define a substantial number of species and subspecies. As a consequence, an unusually large percentage (83%: 35 out of 42) of our scutigeromorph types have since been reinterpreted as junior synonyms and only seven names currently remain valid: Thereuopoda chinensis Verhoeff, 1905, Parascutigera dahli Verhoeff, 1904, Ballonema gracilipes Verhoeff, 1904, Podothereua insularum Verhoeff, 1905; Parascutigera noduligera pahangiensis Verhoeff, 1937, Thereuopodina tenuicornis Verhoeff, 1905 and Thereuonema turkestana Verhoeff, 1905. In fairness, Verhoeff was not alone in creating synonyms for scutigeromorphs. A cursory glance at the synonymy lists for the more common and widespread species (Bonato et al. 2016) often reveals more than a dozen names proposed (unnecessarily) for these distinctive animals.
One could argue that this high level of synonymy weakens the overall significance of the Berlin collection, which at the present time (Table 1) consists of nineteen valid species (seven as types); or about 20% of the world fauna. We would argue that in fact this is an example of a thoroughly revised collection. Excessive numbers of unrecognized synonyms give a misleading picture of biodiversity, both at a local and at a global level. Thanks to the efforts of Muralewitsch (1910), Würmli (1975aWürmli ( , b, 1977Würmli ( , 1978Würmli ( , 1979Würmli ( , 2005, Stoev and Geoffroy (2004) and Giribet and Edgecombe (2013) most of these problems have hopefully already been addressed for the scutigeromorphs in Berlin, leaving us with a small but well-resolved collection which we hope will be valuable for studies of systematics and/ or biogeography. In this context, we offer the following data as a resource to future specialists. A record of a fossil scutigeromorph type in the palaeontological collections of the Berlin Museum is also listed for completeness.
Remarks. "Tsingtau" was a German colonial possession from 1898-1914. The collector may have been a naval officer, Dr H. Glaue, who was a student of the zoologist H. Korschelt from Kiel and who subsequently tried to establish a biological station in Tsingtau in 1912.
annuligera Verhoeff, 1904 Pselliophora annuligera Verhoeff, 1904 Type material. Syntypes, 2♂, 1♀, ZMB 3837 / 3837a-r (Verhoeff slide nrs 2568-2583); "Tanagebiet, D. O. Afrika" [Tanzania or Kenya?]; leg. Denhardt, 23.VIII.1895. Present name. Junior synonym of Sphendononema rugosa (Newport, 1844); synonymized by Würmli (2005). ; one of a pair of brothers who undertook several expeditions to east Africa and were active in promoting the exploration and use of the Tana river. Gustav was still in the area in 1895, his brother having returned to Germany. Note that the Tana region may refer to the Tana river which today lies within the political boundaries of Kenya.

Remarks.
The collector was Leopold Conradt (dates uncertain) who established the Johann-Albrechts-Höhe station in Cameroon during its time as a German colonial possession.
Remarks. The collector was Ernst Baumann (1871-1895) who studied natural history before taking up a post at Missahoe in 1893 through the Colonial Department of the German Foreign Office. He collected zoological, botanical and ethnological material, but contracted malaria and died on his return to Germany. The type locality is a former German 'Kolonialstation' and now falls within the political boundaries of Togo. Note that slide number 2850 could not be found as of 2016.

Remarks.
There is historical to reference to a "Pongoor bei Malakka" which implies a type locality in the Malacca State of Malaysia. It may well be an older name for Pangkor Island, whereby Fedor Jagor (1816-1900) is known to have collected material in "Pongoor" for the Berlin museum in 1873-1876 and again from 1890-1893.
Remarks. The type locality could be a misrepresentation of Bolgoda Lake, south of the capital Colombo, the largest natural lake in Sri Lanka.

Remarks. The collector was Eberhard von Oertzen
hilgendorfi Verhoeff, 1905 Thereuonema hilgendorfi Verhoeff, 1905a (Wood, 1862); synonymized by Würmli (1975a).  was curator of molluscs at the Berlin museum and took part in the 1860 'Thetis' [or Eulenburg] Expedition to the Far East which included a port of call at Yokohama. Franz Hilgendorf (1839Hilgendorf ( -1904 was another Berlin curator (for fish and crustaceans) and a pioneer of zoological research in Japan. He was active here from 1873-1876 as a lecturer at the Imperial Medical Academy in Tokyo. Details about the third collector (Brauns) were not available.

Remarks. Eduard von Martens
insularum Verhoeff, 1905 Podothereua insularum Verhoeff, 1905b Type material. Syntypes; ZMB 3841-3843, 3857, 3858; "Ralum" [Bismarck Archipelago, Papua New Guinea]; leg. F. Dahl. Verhoeff, 1905 Remarks. For notes on the collector see P. dahli. Moritz and Fischer (1979) noted that ZMB 3857 and 3858 could not be found at the time of writing but may have been the source of the Verhoeff slides numbered 2505 and 2533. There are also several additional Verhoeff slides (nrs 2505-33) also from the "Ralum" type locality. They are not explicitly marked as types, but could potentially belong to the type series.
Remarks. The collector was the botanist and explorer Johann Maria Hildebrandt (1847-1881) who was active in Madagascar from 1880 until his death there from fever in 1881. A precise locality for the type material is not available.

Remarks. The collector was the explorer and ethnologist
Max Quedenfeldt (1851-1891) who was active in Morocco (and neighboring lands) and was widely regarded by contemporaries as one of the most knowledgeable people about this country. Material from Tangier suggests a specific collecting trip made in 1885.

Remarks. The collector could be the Berlin-based physician and medical staff officer ('Oberstabsarzt'), Hermann
Rabl-Rückhard (1839-1905), although it's not clear from available biographical details whether he visited South Africa. Moritz and Fischer (1979) noted that the type material was originally numbered ZMB 751, but was inadvertently re-registered as number 3849 after being loaned to Verhoeff.

Remarks.
Type material consists of slide-mounted material only.

Remarks. For notes on the collector Hilgendorf see T.
hilgendorfi.

Present name. Thereuopodina tenuicornis
turkestana Verhoeff, 1905 Thereuonema turkestana Verhoeff, 1905a Type material. Holotype, ♀, ZMB 87 / 87 a-c (Verhoeff slide nrs 1615, 2982-2983); "Buchara" [Bukhara, Uzbekistan]; leg. Eversmann. Verhoeff, 1905. Remarks. The collector was Eduard Friedrich Eversmann (1794-1860) who visited Bukhara in 1820 and became a professor of natural history at the University of Kazan. Eversmann sent specimens to Heinrich Lichtenstein, the then director of the Berlin Museum which had recently been founded (in 1810). Thus ZMB 87, collected only ten years later, is one of the oldest centipedes in the collection. The non-slide material is currently held in the dry collection.

Present name. Thereuonema turkestana
vagans Verhoeff, 1905 Pselliophora vagans Verhoeff, 1905a Type material. Holotype, ♂, ZMB 3893 / 3893a-e (Verhoeff slide nrs 2617-2621); "Chinchoxo" [Cabinda Province, Angola]; leg. Falkenstein, VI.1876. Present name. Junior synonym of Sphendononema rugosa (Newport, 1844); synonymized by Würmli (2005), see also Stoev and Geoffrey (2004). Falkenstein (1842Falkenstein ( -1917, a military doctor who took part in the 'Loango Expedition' of 1873-1876 to the Congo river delta under Paul Güßfeldt. "Chinchoxo", sometimes spelled Tschintschotscho, was a station on this expedition. The site of Chinchoxo is a few km north of the modern locality of Lândana, now in an enclave of Angola known as Cabinda Province, surrounded by both the Republic of the Congo to the north and the Democratic Republic of the Congo to the south.
Remarks. The number in the jar is not a ZMB catalogue number and the locality is, unfortunately, indecipherable. However, the collector could be Julius Gollmer a pharmacist who was active in northern South America (mostly Venezuela) around 1856.

Sphendononema rugosa (Newport, 1844)
Material. 1 dry specimen (as Pselliophora vagans Verhoeff, 1905); "Congo", leg. Heyne. Remarks. The collector of the Sri Lankan material was Johannes Nietner (1828-1874), a gardener in the botanical garden in Paradeniya. Note that the identification of the Sri Lankan specimen should be treated with caution (G. Edgecombe, pers. comm.) as this would represent a considerable range extension both for the genus and the family Pselliodid as a whole.

Remarks.
The "Deutsche Südpolar-Expedition" -sometimes referred to as the "Gauß-Expedition" -ran from 1901-1903 under the leadership of Erich von Drygalski. The Simons Town material was evidently collected during a stopover in South Africa on the return leg of the expedition. Silvestri (1904) described the species shortly afterwards, but the Berlin material does not belong to the type species which is cited as having come from "Zwarsberg" in South Africa with Max C. Weber as the collector. The BIOTA material stems from a major project on the southern African fauna and was collected by MfN staff members Jürgen Deckert, Manfred Uhlig and Peter Giere; with identification by Katrin Vohland. Remarks. The collector of ZMB 3835 could either be an A. H. Zietz who was ca. 1892-1908 Assistant Director of the South Australian Museum, Adelaide, or a Eugene Hintz who collected beetles and other arthropods for the Berlin museum. Much of the material of this species stems from a 1905 expedition to southwest Australia by Wilhelm Michaelsen (1860-1937) and Robert Hartmeyer (1874-1923. The latter was from 1900 an assistant (later from 1908 curator) in Berlin for marine groups like bryozoans and echinoderms.

Ballonema Verhoeff, 1904
Ballonema gracilipes Verhoeff, 1904 Material. 1 specimen, ZMB 5382 "Sattelberg bei Finsch-  who was active in New Guinea from 1908-1910. The collector of ZMB 5383 was Leonhard Schultze (1872-1955 a zoologist and anthropologist who took part in a Dutch-German expedition to New Guinea in 1910. Note that the identity of this material was confirmed in the literature by Butler et al. (2010: 548).

Remarks.
With respect to the dry record from Carolina, this predominantly southern European (but also synan-thropic species) is also known from the USA and several other places, although it is not entirely clear whether S. coleoptrata is native or introduced here. Most of the material is from southern Europe and the Canary Islands. The collectors Dirk Striebing, Sepp Lüdecke and Hella Wendt were all former staff members of the MfN. (Newport, 1844) Material. 1 specimen (as rubrolineata); ZMB 3834; "Ceylon" [Sri Lanka]; collector and date not recorded. ; "Ceylon".

Scutigera rubrilineata
Remarks. The Sri Lankan material here was presumably assigned to Cermatia rubrolineata, described by Newport from 'India Orientali'; the status of which is currently uncertain (P. Stoev pers. comm., 2017). In Chilobase it is suggested that this species might be conspecific with Thereuonema microstoma (Meinert, 1886 Remarks. The Ethiopian material was collected by the ornithologists Carlo von Erlangen  and Oscar Neumann (1867-1946 who were active in East Africa. The Paraguayan material may stem from the botanist Andrés Barbero (1877Barbero ( -1951. Details of the other collectors are uncertain. Two additional dry specimens, ZMB 88 ("Aegypt" [Egypt]; leg. Ehrenberg) and ZMB 89 ("Siber?" [Siberia?]; leg. Pallas are labelled with what we presume to be manuscript names of Scutigera under its junior synonym Cermatia Illiger, 1807. Neither of the species names in the museum catalogue could be found in Chilobase. Ehrenberg as collector is mentioned above, while Pallas is presumably Peter Simon Pallas (1741-1811) who carried out a famous collecting trip to Siberia from 1768 to 1774 at the bequest of the Russian empress Catherine the Great. If this specimen does originate from this journey it would be a strong candidate for being the oldest specimen in the myriapod collection; predating the founding of the museum in 1810. Note that Pallas lived in Berlin shortly before his death in 1811 and it is quite possible that some of his zoological material passed to the then newly founded museum.
Remarks. ZMB 5116 was discovered in a termite mound belonging to the species Odontotermes redemanni (Wasmann): "Im Hügel von Termes redemanni". The collector was the forester and entomologist Karl Escherich  who authored a work about the termites and ants of Sri Lanka. The collector of 13180 was Bernhard Rensch   Cermatia illigeri Koch & Berendt, 1854 Type material. Holotype, MfN palaeontology collection Nr. 7317 (ex Berendt collection); Baltic amber (Eocene: Lutetian).
Remarks. The repository of another amber species described by these authors, Cermatia leachi Koch & Berendt, 1854, unfortunately remains equivocal. It could not be traced in the Berlin palaeontology collection at the time of writing (C. Neumann, pers. comm.).