Corresponding author: Mark-Oliver Rödel (
Academic editor: Johannes Penner
We present a commented catalogue of the type specimens of the Afro-Malagasy frog family
The amphibian collection of the Museum für Naturkunde Berlin (
The oldest specimens from this family present in the collection are from South Africa. These vouchers, sent by G. L. E. Krebs (1792–1844), are two
During the second half of the 19th century, the collection was enriched by material collected, for example, by R. W. Buchholz (1837–1876) and G. A. E. Reichenow (1847–1941) in Cameroon, by C. C. von der Decken (1833–1865) in East Africa and by H. W. L. Dohrn (1838–1913) in Liberia. Several objects from the collection of J. A. de Oliveira Anchieta (1832–1897) from Angola were received in exchange with J. V. Barbosa du Bocage (1823–1907) from Lisbon. Wilhelm Peters undertook the first scientific work on the Berlin
However, most hyperoliid frogs now present in
Examplary page from the first numerical inventory catalogue of the herpetological collection at
At the end of the 19th century, Gustav Tornier (1859–1938), at the time curator of the amphibian and reptile collection at the
“Farbkleidmuster der Gattung
A decade later, at the beginning of 20th century, large scale research expeditions such as the first and second “Deutsche–Zentral–Afrika–Expedition” [German-Central-African-Expedition] from 1907–08 and 1909–10, provided rich new material. Together with specimens obtained from the German colony Cameroon, further remarkable collections of amphibians and reptiles, including considerable numbers of reed frogs, were sent to
In December 1921, Christoph Gustav Ernst Ahl (1898–1945) started to work as a voluntary scientific assistant in the ichthyological and herpetological department of
Several ‘Ahl species’ were synonymized by
After 2000,
We present a list of existing and so far unlocated type specimens of 146 nominal taxa of the family
From 2017 to 2020, all specimens from the family
Examplary page from the second numerical accession catalogue (= “C” catalogue) of the herpetological collection at
We list all types in alphabetic order, using the original name in the description. The present taxonomic status and generic association is given under ‘present name’ in each species account. Remarks in the individual species accounts contain information on illustrations of the type material, the activities of the collectors, and the collection periods as well as information taken from secondary literature on additional type material that is not housed in the
Recently,
In the chapter “Specimens erroneously marked as types in
It is important here to provide commentary on the publication history of
Titlepages of
However, it needs to be emphasized that, without doubt, Ahl intended to publish the paper “Zur Systematik […]“ (
To finally clarify the history of both publications (
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W. C. H. Peters, a zoologist, anatomist and later director of
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In 1896, the German physician, doctor of tropical medicine, and explorer Fülleborn joined the colonial “Schutztruppe” in German East Africa, where he was active as a government physician until 1901. From April 1897 to the beginning of 1898, he participated in the military campaigns against the Wangoni and Wahehe of the “Ungoni”, “Uhehe” and “Ubena” regions, in present day southern Tanzania. From 1898 to 1899 Fülleborn was stationed in Langenburg [Lumbira, Mbeya Region, Tanzania] in the north of Lake Malawi and undertook numerous excursions in the surrounding area, which took him to the southern end of Lake Malawi, through the “Schire-Hochländer” [Shire Highlands, southern Malawi], and on the Shire and Zambezi River to Quelimane and afterwards to the Island of Mozambique. In 1899 he was commissioned to research the “German-Nyassa” region from a zoological and ethnological-anthropological point of view. Together with W. Goetze he participated in the “Nyassa–See- und Kinga–Gebirgs–Expedition” (
The German explorer, zoologist and anatomist Buchholz went to Equatorial Africa from 1872 to 1875. He was accompanied by the Berlin ornithologist Georg Anton Eugen Reichenow and Reichenow’s friend, fellow student and zoologist Wilhelm Lühder. On 1 June 1872 they set off from Bremerhaven to “Akkrá on the Gold Coast” [Accra, Ghana], which they reached on 29 July 1872. The first collecting tours took place in the surroundings of Accra and Aburi (29 July to 16 October 1872). On 16 October they left Accra for “Camaroons” [today part of present day Douala city] where they stayed until 2 November. Then they travelled to Bimbia, Victoria and Bonjonjo (2 November 1872 to 9 December 1873). On 12 March 1873 Lühder died of malaria in ‘Camaroons’. Reichenow, also suffering from malaria, returned via Gabon to Germany in April 1873.
Buchholz was on his own from then on. He travelled between Victoria and `Camaroons’ with intermediate stops on Fernando Pó to get his collections to Camaroons in early December.
Thereafter he went to Abo (9 December 1873 to 24 March 1874) and from Mungo via Balong he returned again to ‘Camaroons’ (5 April to 11 August 1874). He left `Camaroons’ for a stay in Gabon where he also explored the Rembo River (12 August to 9 November 1874). After his return to the Gabon coast he again explored the area around Mungo and Jenssoki (9 November 1874 to 11 January 1875), and again visited Fernando Pó, the Gabon coast and the Ogowe (or Ogooué) River (11 January to 31 August 1875). On 3 September he started from Gabon on his way back to Greifswald where he arrived during the beginning of November 1875 (
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Then, the caravan turned north, crossed Lake Kivu with a stop at “Kwidschwi” Island [September 1907; Idjwi (Ijwi) Island, Lake Kivu, Democratic Republic of the Congo] and reached “Kissenji” [September 1907; Gisenyi on the northeast shore of Lake Kivu, close to the border of Democratic Republic of the Congo, Rwanda]. From there they went to Rutschurru [December 1907, Rutshuru, North Kivu Province, Democratic Republic of the Congo] and further north to Vitshumbi [December 1907] at the southern tip of Lake Edward. The expedition continued further along the west coast of Lake Albert to reach the Rwenzori Mountains via Kasindi [January 1908]. From Fort Beni [January to February 1908] on the western slopes of the Rwenzori Mountains the expedition went to Kassenje [March 1908] on the southwestern shore of Lake Albert. From here, the expedition turned west. Via Mawambi [April 1908] on the Ituri River and Avakubi [April 1908], it went along the left bank of the Aruwimi River to Basoko [May 1908; Tshopo Province, Democratic Republic of Congo] to the confluence with the Congo River, where the expedition ended in June 1908 (
From 1909 to 1910 a second “Deutsche Zentral–Afrika–Expedition”, also under the leadership of Adolf Friedrich, Duke of Mecklenburg, extended along a main route from Cameroon via Spanish Guinea [Equatorial Guinea], Gabon, the Congo and the Ubangi River up to Fort de Possel [Possel, Central African Republic], and from there further north to Lake Chad and back via North Cameroon to the Niger Delta. Schubotz, who accompanied this expedition, deviated along the Ubangi River eastwards, in order to follow the White Nile in southern Sudan and returned via Khartoum and Egypt to Germany (
The “Africanische Gesellschaft”, or formally “Deutsche Gesellschaft zur Erforschung Aequatorial-Africas”, sponsored the “Loango–Expedition” from 1873–1876 under the leadership of the German geographer and explorer Richard Paul Wilhelm Güssfeldt. The expedition had the task to establish a station at the Loango coast (at Chinchoxo), which was to serve as a depot for the material collected during the expedition. Geographic-topographical explorations into the interior of the African continent were also intended to be carried out. With an interdisciplinary research team, comprising the medical officer and zoologist Julius Falkenstein, the geographer Eduard Pechuël-Loesche, the geodesist von Görschen, Reserve Lieutenant Hans von Hattorf, the mechanic Otto Lindner, the botanist Herman Soyaux, and the topographer Major Alexander von Mechow, Güssfeld travelled for two years, starting in July 1873. They mainly followed the coastal area of Cabinda, on the Kouilo river, the Chiluango river, and on the lower course of the Nyanga river. From March 1874, they turned to Luanda, on the Cuango to Dondo and to the rapids of Cambambe, as well as to Quicombo and Novo Redondo (
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Dr. Hösemann served in the colonial German Schutztruppe as medical officer [Stabsarzt], undertook anthropological studies, and between 1897 and 1907, collected zoological objects on the northeastern shore of Lake Tanganyika (Udjidji), in the Kissaka Region (southeast of Lake Mugesera, Ngoma and Kirehe Districts, Eastern Province, Rwanda] and between Mwanza and Moshi [northern Tanzania] (
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Between 1896 and 1903 the engineer and transport officer of the “Mombasa–Uganda Railway” in British East Africa, Felice (sometimes Felix) Thomas sent several shipments, containing amphibians and reptiles, from the Kenyan coast province (Mombasa and Takanugu) and from Nairobi to
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The natural scientist and cartographer Baumann joined the German Colonial Service in 1893. He worked at Klein Popo [Aného, Lacs Prefecture, Maritime Region, Togo] and was later stationed at Misahöhe [Agou Prefecture, Plateau Region, Togo] where he was deputy station chief from 1894–95. In November 1894, he accompanied the “Togo–Hinterland–Expedition” headed by the colonial officer Hans Gruner along the Volta River to Kete Kratschi [Kete Krachi, Oti Region, Ghana] and returned to Misahöhe. In the hinterland of the station (Agome Region) he collected zoological, botanical and ethnological objects, which were given to the museums in Berlin. In 1895 he returned to Germany where he died on 4 September as a result of malaria that he contracted on his return journey (
see unlocated type specimens’.
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In October 1905 and November 1907, Tiesler sent two shipments, including nearly 300 amphibians and reptiles, to
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Rolle was a well-known dealer of zoological and enthnological objects. He maintained a worldwide network of collectors and suppliers and acquired several important collections. From 1889 onwards, he supplied private collectors as well as important European museums with zoological objects from Berlin. In later years, he traded objects under the name of the natural history institute “Kosmos”.
see “unlocated type specimens”.
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Drawings illustrating the variation of this taxon are given by
The Austrian hunter and Africa explorer Grauer undertook several expeditions to Eastern Africa, e.g. to British East Africa [Uganda] (February to May 1904 and September to November 1905) and to Tanganyika in 1907, where he met the first “Deutsche Zentral–Afrika–Expedition” at Lake Kivu in August. Upon this meeting he handed the zoological material he had collected in the “Zwischenseengebiet” [Region between Lake Victoria, Lake Kivu and Lake Malawi, Tanzania] for
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The English ornithologist and entomologist Rosenberg collected mainly for the British Museum of Natural History (
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Gleim was Deputy Governor of the German Colony of Togo from 1896 to 1898. From 1899, he was sent to São Paolo de Loanda by the “Kolonialabteilung des Auswärtigen Amtes”, where he served as the first professional consul for Angola and French Congo. From 1910 to 1911 he was Governor of Cameroon (
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The German zoologist, cartographer, explorer and colonial official Stuhlmann spent a total of 14 years in East Africa. With the financial support of the Akademie der Wissenschaften [Academy of Sciences] zu Berlin, he investigated the coastal regions of Zanzibar and the adjacent mainland including “Usegúa” and “Ungúu” in present-day Tanzania in the summer of 1888; then, until mid-1889, the area of the Zambezi estuary around Quelimane in Mozambique. From April 1890 to 1892, Stuhlmann participated as a scientist, together with Lieutenant Wilhelm Langheld, on the expedition of Mehmed Emin Pasha [also known as Eduard Karl Oskar Theodor Schnitzer] to the German East African colonial area. The expedition led them from Bagamoyo (26 April 1890) via Tabora (29 July) to Bukoba on Lake Victoria (November 1890). From here, Stuhlmann undertook a trip on Lake Victoria to Murchison Bay in Uganda (December 1890) and reached Mengo Mountain (26 to 29 December) via Manjongo [Rubaga Division, Kampala District, Central Region, Uganda]. After his return to Bukoba, he set off (12 February 1891) towards the west in the Karagwe Region, and after crossing the Kagera River (06 April 1891), the expedition reached the southwestern tip of Lake Edward in early May 1891. The expedition turned west of Lake Edward another 250 km to the north, but was terminated in mid-September 1891 due to insurmountable difficulties. With a group of 27 askaris (local soldiers serving in European colonial armies) and 100 porters Stuhlmann went back to Bukoba, where he arrived on 17 March 1891. Emin Pasha, in contrast decided to stay behind with sick expedition members, turned southwest towards the Congo River and was murdered by Arab slave traders 80 km from this destination at Kinene on Mwiko River on 20 October 1892. Stuhlmann’s herpetological collections from these expeditions were sent to Johann Georg Pfeffer at the Zoologische Museum Hamburg, who published the first results (
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In March 1872, Hildebrandt travelled from Berlin via Egypt to the southwest coast of the Arabian Peninsula and to Aden, where he stayed until the end of 1872. In spring 1873, he went from Zanzibar to Karachi and travelled the Indus upwards. After returning to Zanzibar in July 1873, he travelled the Wami and Kingani (Rufu) Rivers in present-day Tanzania together with the animal trader and director of the Hamburg Zoo, Carl Gottfried Wilhelm Heinrich Hagenbeck, and then visited the southern Somali coast alone. He returned to Europe in August 1874.
In February 1875, Hildebrandt arrived again in Aden and visited the “Serrut Mountains” [Somaliland]. Then he went to Zanzibar and the Comoros (Johanna Island [Anjouan], June to September 1875). Back in Zanzibar he prepared his expedition into the Inner Africa, via Pangani [Tanzania], Lamu, through the South Gala countries up the Tana River. He had to return to Mombasa due to illness in December 1875. In November 1876 he started again from Zanzibar via Mombasa (10 January 1877) in the direction of Mount Kenya. He travelled the Taita, Ukamba and Kitui areas, but had to return to Mombasa without reaching his actual destination Mount Kenya, from which he was only a three days’ march away. He arrived again in Mombasa in August 1877 (
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First Lieutenant May collected between 1901 and 1903 in Luanda and surroundings, e.g. in Mubella near Funda on the Bengo River [Municipality of Cacuaco, Luanda Province, Angola] (
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The Berlin ornithologist Reichenow travelled together with Lühder and Buchholz from spring 1872 on a one year collecting trip to “Akkrá” on the Gold Coast [Accra, Ghana] and the region around “Camaroons” [Douala Region, Cameroon] (
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The Prussian medical officer and discoverer of one of the sources of the Nile, Richard Kandt (who used Kantorowicz until 1894) explored the northwestern part of German East Africa from October 1897 to January 1898, and the region around Lake Kivu between 1898 and 1902 (
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First Lieutenant, later Captain, Köhl served from 1912 on in the colonial “Schutztruppe” of Deutsch Ostafrika at Kissenji, and from 1916 on in various missions under General Paul Emil von Lettow-Vorbeck, e.g. at Taveta [Kenya], Port Amelia [Pemba, Cabo Delgado Province, Mozambique] and Medo [Metoro, Mozambique] (Haup 1988;
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The zoologist and anatomist Böhm, together with the explorer Paul Reichard, travelled on behalf of the “Africanische Gesellschaft” from Zanzibar via Bagamojo [27 July 1880] to Tabora, which they reached two-and-a half months later. From here they turned to Kakoma [southeast of Tabora, Tabora Division, Tanzania], where they stayed for over a year. Then they continued to Jagonda [just northeast of Kakoma]. From Jagonda, Böhm and the topographer Emil Kaiser went on a journey to Lake Tanganyika, lasting several months. They reached Karema on the western shore of the lake [Mpanda District, Katavi Region, Tanzania] and returned to Jagonda on December 23, 1881. In March 1882 they travelled along the Wala River. Dr. Kaiser died during an expedition to Lake Rukwa near Upia on 27 October 1882. Towards the end of December 1882, Böhm and Reichard left Jadonda for Karema, crossed Lake Tanganyika to Mpala (at the mouth of the Lufuku River, Tanganyika Province, Democratic Republic of the Congo) and reached the “Marunga Land” in July 1883. From here they turned southwest and discovered Lake Upemba in the Urua region [Upemba, Bukama Region, Haut-Lomami Provinz, Democratic Republic of the Congo]. On 27 March 1884 Böhm died in southern Urua, south of the Lake Upemba (
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Because of a reading error, assuming
The physician Dr. Buchner arrived in Luanda on 5 December 1878 and travelled via Dondo (20 December 1878) and Malanje (30 January to 22 July 1879) to Mussumba in the Lunda Empire (11 December 1879 to June 1880). He returned to Malanje (28 February 1881) and via Golungo and Cazengo travelled back to Luanda, where he arrived at the end of August 1881. He finally returned to Berlin in January 1882 (
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The German naturalist, botanist and gardener Zenker joined the German colonial service as taxidermist in 1889. He was manager of the colonial station Jaunde (Yaoundé, Mfoundi Department, Centre Region, Cameroon) from 1890–1895 (
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The German physician Dr. Oeser undertook various journeys, e.g. to East Asia and Indonesia (1923 as a ship’s doctor), to the USA (1925), to Central America and northern South America (1931–32) as well as to Cameroon (1936). In spring 1928 he undertook a journey along the West African coast, collecting in Benin, Nigeria, Fernando Pó, Sao Tomé and Principe, Angola, Namibia and Liberia. He collected the type material of
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Currently, the status of this taxon is uncertain. In comparison to species in the
Under the leadership of Adolf Friedrich, Duke of Mecklenburg, the first “Deutsche Zentral–Afrika–Expedition” was carried out from 1907 to 1908, to scientifically investigate the area of the African Rift Valley (see also remarks on
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Five drawings showing the variation of this taxon are provided by
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Two drawings showing the variation of this taxon are illustrated by
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Between December 1896 and June 1899, the German planter Martienssen sent amphibians and reptiles from the German colony in East Africa to
36108, “Mohorro, Deutsch-Ost-Afrika” [Mohoro (Muhoro), Pwanai Region, Tanzania], coll. Karl Grass, 22.II.1901.
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36104, “Klein-Popo, Togo” [Anhéo (Anecho or Popovi), Lacs Prefecture, Maritime Region, Togo], coll. Julius Graf von Zech auf Neuhofen.
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The type material of
Scheffler worked from 1899 to 1900 as a horticulturist on plantations of the German East African Society in Usambara (Nguelo and Derema), in the Useguha region [Tanzania] and from 1905 until his death on 10.VI.1911 as a farm manager under managing director Paul Huebner in Kibwezi, British East Africa (
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The German trader and zoologist Spatz carried out various hunting and collecting expeditions in the northern Sahara during which he also collected ethnological and prehistoric objects. In 1884 he travelled to Tunisia and Algeria for the first time. Further journeys to Tunisia were as follows: in spring 1891, together with Alexander Koenig and Koenig’s wife; in 1893 and from November 1896 to July 1898 together with Carlo von Erlanger; and from 1904 to 1906 together with Otto Eduard Graf von Zedlitz und Trützschler and Alfred Blanchet. In the early 1920s he travelled to Mauritania, to the lower Senegal River and the Spanish colony of Rio de Oro. The latter he visited again with the Berlin taxidermist Fritz Bock in spring 1926. On behalf of
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For activities of Stolz in East Africa, see remarks on
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The Prussian explorer and topographer Major Mechow participated in the first “Loango–Expedition” from 1873–1875 under Paul Güssfeldt (Güssfeld et al. 1879). On a second “Kuango–Expedition” (1878–81), led by himself, he and two companions, the botanist Julius Eduard Teusz and the shipwright of the Imperial Navy Jess Bugslag (or Buslag), travelled from Luanda via Malanje (June 1888) to the confluence of the Luhemba and the Cuango River (November 1880). He returned via Malanje (February 1881) to Luanda and arrived in Berlin in August 1881 (
From 1878 to 1879, the expedition of engineer Schütt was carried out together with the architect Paul Gierow on behalf of the “Afrikanische Gesellschaft in Deutschland”. The expedition aimed at compiling topographic reconnaissance and producing maps. They started on 4 January 1878 in Luanda and reached the lower Luachimo River (3 to 9 February 1879) via Malanje (22 February to 4 July 1878), and Quimbundo (12 November to 1 December 1878). They almost reached Mai Munene. From Quimbundo they turned back through the Lunda area and reached Luanda again via Malanje (12 to 24 May 1879) on 21 June 1879 (
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The holotype was probably collected by Rev. Father Pascalis Boneberg of the Trappist Mariannhill Mission who sent several specimens to
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For Martienssen’s activities in East Africa and locality information and collecting dates, see remarks on
From 1900 onwards, the ornithologists, mammalogists and explorers von Erlanger and Neumann undertook a two-year journey through Somaliland to the south of Ethiopia. They were accompanied by the physician and collector of botanical objects Dr. Hans Ellenbeck, the cartographer Johann Holtermüller and the taxidermist Carl Hilgert. From Zeila at the Gulf of Aden [Zeylac District, Awdal Region, Somaliland] they started their journey on 12 January 1900 and travelled via Djeldessa [Jaldessa, Sitti zone, Somali Region, Ethiopia] (3 March 1900), Harar (1 April 1900) , Biar-Woraba [Bia-Woraba, East Harerge Zone, Oromia Region, Ethiopia] (23 May 1900), the Wabbi River [Webi Shebeli] (passage on 10 June 1900), to Addis Ababa (16 August 1900). During that journey they climbed Abu-el-Kassim [Abul Kasim, Arsi Zone, Oromia Region, Ethiopia] on 16 July 1900. From Addis Ababa Neumann went alone to Shoah [Shewa Kingdom, a region present day in Central Ethiopia] and southern Sudan, then returned to Cairo via Khartoum. Erlanger went on to Lake Turkana (
However, the location “Quillimane” as mentioned in the
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One of the 12 syntypes mentioned by
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Not assigned to a valid name according to
As far as we are aware this nomen was not used again as valid after its introduction by
In a letter sent to Wilhelm Peters, dated 12 June 1869, Bocage announced a shipment containing 30 species of “Reptiles et Batraciens”. He listed under no. 21 a “
Beside the specimen from “Dombe” which Tornier regarded as “Type”, Tornier also mentioned “identical” specimens from “Chinchoxo”,
Our examination of the holotype revealed that it is a member of the ‘sharp-nosed reed frogs’,
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For information on the expedition period and route of Erlanger and Neumann, see remarks on
Unlocated type specimens
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Drawing in Ahl, 1931b: 414, fig. 287.
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The German colonial officer and planter Conradt was stationed at “Lolodorf” [Océan Division, South Province, Cameroon] and worked as station manager at “Johann Albrechtshöhe” [southeast of Lake Barombi Mbo, near Kumba, Southwest Region, Cameroon] from 1895 to 1899 (
The original catalogue entry of
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We are grateful to Annemarie Ohler (
Documents kept in the Department of Historical Research (Historische Arbeitsstelle) of the Museum für Naturkunde Berlin and consulted ZMB register (accession catalogues):
Lichtenstein, H. M. Eingangsjournal über den Zugang von Naturalien des Königlichen Zoologischen Museums zu Berlin von 1816 bis 1857 (2 vols.).
Zoologisches Museum, Signature S II, “Barboza du Bocage”.
Zoologisches Museum, Signature S II, “Brühl, Ludwig”.
Zoologisches Museum, Signature S III, “Fülleborn, F.”.
Zoologisches Museum, Signature S III, “Küttner, Otto, Dr.”.
Zoologisches Museum, Signature S II “Museum Hamburg“.
Zoologisches Museum, Signature S II “Martienssen, G.”.
Zoologisches Museum, Signature S II, “Mechow, v. – Major”.
Zoologisches Museum, Signature S III, “Rolle, H.”.
Zoologisches Museum, Signature S II “Stuhlmann, F.”.
Zoologisches Museum, Signature S I “Temminck, C. J.” (incl. correspondence of H. Schlegel and H. Boie to H. Lichtenstein).
Zoologisches Museum, Signature S II, “Thomas, Fe.”.