An annotated list of reptiles and amphibians from the 1905 Hamburg expedition to southwest Australia deposited in the Zoological Museum Hamburg

The herpetological material of the 1905 Hamburg expedition to southwest Australia is redetermined and listed with precise locality data and habitat details. Of this material, 275 specimens of 57 species are still part of the herpetological collection of the Zoological Museum Hamburg (ZMH). A map showing 47 (out of 167) collecting points is provided. Some of the type specimens described in the original material by Prof. Franz Werner have been destroyed. A single paratype of Crinia michaelseni (Werner, 1914) now Geocrinia leai (Flecher, 1898), formerly thought to be lost, was recovered in the ZMH collection. This historical collection is a valuable resource for understanding the composition of the herpetofauna of the previous century.


Introduction
Historical collections are valuable sources of information for investigating a country's past and present species inventory. Herpetological material from Australia is relatively rare in museum collections, and today it is difficult to obtain new material from Australia due to strict export and nature conservation regulations.
In 1883, the collections from a German expedition to South Georgia Island, near Antarctica were given to the Natural History Museum Hamburg (today's ZMH). The collection, mainly of invertebrates, was intended for further study and the Museum planned to make further expeditions to obtain collections from biogeographically adjacent areas. In 1892, Wilhelm Michaelsen, first curator of the Natural History Museum Hamburg, went on an expedition to Patagonia, Chile und Tierra del Fuego (Hamburger Magellanische Sammelreise 1892/93). Once that material had been analyzed, Michaelsen began planning another expedition. His original plans to travel to Africa had to be altered because, at the same time, another scientist (Leonard Schultze from Jena) organized an expedition to southwest Africa. Michaelsen rerouted, and decided to travel to southwest Australia. Michaelsen, together with his colleague Robert Hartmeyer from the Natural History Museum Berlin (today's ZMB), organized and led the expedition to southwest Australia in 1905. As studies of this geographic area were, at the time, very rare, there was a distinct gap in the knowledge regarding its biogeographic distribution of animals: as such, Michaelsen's expedition aimed to close this gap and to provide much needed information and material (Michaelsen and Hartmeyer 1907).
The SW Australian expedition received financial support from both the Natural History Museums represented (Hamburg and Berlin), the Senate of Hamburg, the government of Western Australia and private sponsors (including Strelitz Bros., a company of merchants and shipping agents based in Fremantle in Western Australia, established by two brothers from Hamburg). The expedition group travelled aboard the German mail steamer Karlsruhe and, on the morning of May 5, 1905, they arrived at the port of Fremantle, Western Australia. Along with the expedition participants and material, the ship transported 800 litres of pure ethanol. The zoological base station was in Fremantle. Strelitz Warehouse served as a depot for the ethanol and the hundreds of glass jars for zoological material. The expedition party was permitted to use all train lines free of charge, and many of the collection stations were situated along the railway lines. There were 167 collection points in total (stations 1-167) distributed throughout SW Australia, mainly around Fremantle, Perth, on Rottnest Island, and around Albany in the south (see map). All available animals were collected: terrestrial invertebrates, marine and freshwater invertebrates and terrestrial vertebrates. In this paper, I focus only on the reptile and amphibian collection of the expedition.
Franz Werner, Professor of the University of Vienna and an expert in amphibians and reptiles, was tasked with the determination of the expedition material. He published a detailed compilation of the herpetological material discovered during the SW Australian expedition. His lists comprise three parts: reptiles excluding Gekkonidae and Scincidae (Werner, 1909); reptiles (Gekkonidae and Scincidae) (Werner, 1910); and Amphibia (Werner, 1914).
All material deposited in the ZMH herpetological collection has been checked, redetermined and catalogued. The numerous changes in taxonomy of the Australian fauna since that era necessitate a taxonomic list of the present holdings. Furthermore, not all the specimens documented in Werner's lists (1909Werner's lists ( , 1910Werner's lists ( , 1914 are present. Some of the material was destroyed and lost during WWII, or has been deposited in other herpetological collections, including Hartmeyer's home institution, the Natural History Museum Berlin (pers. com. F. Tillack). Franz Werner was not affiliated with any one museum, but instead worked peripatetically through many European collections. He visited the Zoological Museum Hamburg on several occasions to examine material and sent the collections back to Austria to work on there. Most of the material would then have been returned to Michaelsen and Hartmeyer, who would have split it between their two home institutions. However, Werner presumably retained some of the material, which went to the Natural History Museum Vienna (NHMW) on his death (see Adler 1989).
The catalogues of the herpetological collection of the Zoological Museum Hamburg (ZMH) were all lost during WWII, therefore, the only available information is from the jar labels and literature.

Material and methods
All the jars containing material from the SW Australian expedition deposited in the Hamburg Zoological Museum's herpetological collection were checked and catalogued in a database (FileMaker pro advanced 17). Infor-mation on locality and habitat were noted by Michaelsen (1907) in his appendix, and could easily be assigned to each specimen by the station number attached to the jar. Most of the specimens were specified or described by Werner (1909Werner ( , 1910Werner ( , 1914. Where possible, sex and age were recorded.

Results
List of reptile and amphibian species of the SW Australian expedition of 1905 deposited in the herpetological collection of ZMH.
The list is presented in systematic order, with current specific names, catalog numbers, specific localities and remarks, which address geographic and taxonomical importance. The map shows 47 out of 167 collection points where amphibians and reptiles of the expedition were found. Unless otherwise noted, the jar labels indicate all specimens were collected by Wilhelm Michaelsen.
There are 275 specimens of 57 species and one subspecies still present.

Lost type material
Part of the type material of the following species, described by Werner  Remarks. An additional specimen is located in ZMB (under ZMB 21427) from Australia without specific locality, most likely represents the specimen reported by Werner (1909) (Werner, 1910) ZMH R04404-05 (2 paratypes). Lunenberg, station 138, tall forest, Darling Ranges.

Liopholis pulchra
Remarks. Formerly three paratypes from this locality. The specimen with a complete tail is missing. The holotype from Torbay, station 162, is lost. Werner (1910) described this species in the genus Egernia. (Werner, 1910) ZMH R01851 (paralectotype). Lion Mill, station 99, mountain with tall forest, western hillslope of Darling Ranges.
Remarks. Werner listed two additional specimens listed as Lygosoma (Hemiergis) decresiensis form Donnybrook. These are not present in ZMH, and one from Lunenberg is present (under ZMB 21467). Werner (1910) used these specimens (from South Albany) to erect a replacement name, Lygosoma (Hemiergis) quadridigitatum, for Seps peronii Fitzinger, 1826, due to the homonymy within Lygosoma of Seps peronii with Heteropus peronii (Dumeril & Bibron, 1839). Seps peronii Fitzinger is a nomen nudum, and the name was republished, validated with a description, by Gray, 1831, with Tetradactylus decresiensis (Cuvier, 1829), a homonym within Hemiergis of Tridactylus decresiensis (Cuvier, 1829), in its synonymy. Heteropus Peronii is now Carlia peronii. Werner listed only three specimens from South Albany, but also listed nine specimens from Albany, two from NE of Albany, and additional specimens from Bridgetown (4), Yallingup (1) and Torbay (2). It is possible that some of the specimens from Albany and NE of Albany have become combined with those from South Albany. Remarks. The two specimens from Denham, which were the basis for Werner's Lygosoma bipes var. concolor, are not present in ZMH, but one of them is presumably ZMB 21464. Lygosoma bipes var. concolor is a synonym of Lerista miopus. Remarks. An additional 17 specimens listed by Werner (1914) from Albany (7)  Remarks. The above-listed paratype was only recently detected in the collection of ZMH and was therefore not published in previous type catalogues (Hallermann 1998(Hallermann , 2006. The holotype was collected at Donnybrook on 28 or 29 July 1905, and was apparently deposited in the collection of the Zoological Museum, University of Hamburg (ZMH) but is no longer present there. There is no existing correspondence that documents where the holotype was originally deposited or whether it was exchanged with the American Museum of Natural History.

Lerista kendricki
Paratypes included two specimens from Jarrahdale (one of which is now catalogued as NMW 16383 (Häupl et al. 1994)), one specimen from Lunenberg, five additional specimens from Donnybrook (one of which is now catalogued as AMNH 23451, and was identified as the holotype by Cogger et al. 1983), three specimens from Bunbury (including ZMB 24636), and five specimens from Boyanup (including ZMB 24638) (Bauer et al. 1996). The whereabouts of the remaining specimens listed above without catalog number are unknown, but most were probably in the Hamburg collection and may have been destroyed during WWII.  Remarks. Werner (1914) also recorded specimens of this species from Eradu (n = 6, one under ZMB 24611), Moora (n = 10, two under ZMB 24613, ZMB 24641)), Beverley (n = 3, one under ZMB 24622), Cannington (n = 2), Harvey (n = 3), Brunswick (n = 1) and Northhampton (n = 3), and an additional five specimens from York. None of these are still extant in ZMH. An additional specimen from Cannington, not listed by Werner, is present under ZMB 24635.

Discussion
The state of Western Australia is a large area; it covers 2,529,875 square kilometers -about one third of the whole country of Australia. The Hamburg southwest-Australian expedition of 1905 collected specimens from coastal regions of the southwestern part of WA (map).
As of December, 2018, 560 species of reptiles and 80 species of amphibians are known to occur in Western Australia, with 349 reptile species described after 1905 (Uetz et al. 2018;Frost et al. 2019). At the time of the expedition in 1905, there were 211 reptile species known to occur in Western Australia; 30.8% of this number (65) were found during the expedition and 47 species are now part of the collection in ZMH. There are several reasons for this relatively low number of recorded species: the 1905 expedition only covered a small area of WA, it did not have a herpetological focus and the lack of good transportation at that time made it difficult to collect any specimens a long way from the railway stations. Over land, donkey cart was the only mode of transportation beside the railway. Coastal locality points were probably reached by boat. During the expedition, the time period for collection fell partly in winter (June to August) when reptile activity is low. The Australian landscape is characterized by a dry, hot climate in the interior and a more Mediterranean climate in the southwestern area, with maximum temperatures there of 30 °C in the summer months of January and February. Species occurring in these hostile conditions of the interior landscape must be highly adapted to the hot, dry climate. There are currently 80 amphibian species known from Western Australia, of which 35 occur in the southwestern part of the state (Anstis 2013). At the time of the expedition in 1905, there were 20 amphibian species known to occur in WA, and 13 in the SW part of the state (Werner 1914). Werner (1914) listed ten species and subspecies, of which one specimen of Ranoidea moorei (Copland, 1957) was wrongly determined as Crinia signifera, a species known to occur only in eastern Australia. Ten of the listed species are still present in the ZMH herpetology collection (see above), although identifications of these have changed in some cases. Under the former Heleioporus albopunctatus, four different species have since been newly-discovered and described.