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Everything began in 1843 when Professor Dr Wilhelm Carl Hartwig Peters, during his scientific research trip in Mozambique, discovered a new and previously unknown fish species …

 

Discovery and Early History of Nothobranchius Fishes

 

 

     Peters, a young German scientist, commissioned by King Friedrich Wilhelm the Fourth of Prussia, aimed to research the lower Zambezi during his travels in Africa from 1842 to 1848. Until that time, the internal regions of the African continent remained virtually unknown to the European empires; they were organized by tribal kingdoms and confusing multitudes of hunting areas. The lower reaches of the Zambezi River are located where today Mozambique is. After the discovery voyage of Vasco da Gamma, its coast became an area under influence of the Portuguese. The dominion of the Portuguese has spread along the Zambezi, until the town called Tete, founded by Portuguese Dominican missionaries in 1531, situated some 400 km from the estuary of the river. Between the Zambezi and Limpopo rivers the Zulu founded their kingdom.

     The then 28-year-old Peters reached its original destination of Mozambique in June 1843, after a few months of detour in Angola. In Mozambique, he carried out his research mainly between Tete and the mouth of the Zambezi. He diligently collected a variety of animals along the river and, in addition to several species of birds and reptiles, he also found and counted forty-three fish species, which he regarded as a relatively high number. In 1844, one of the new species descriptions was published with the name of Cyprinodon orthonotus in a German scientific journal.

     Peters found the new species north of the estuary of the Zambezi, near the town of Quelimane. After his return in Europe, he reported about the results of his research in a four volume publication with the title of 'Naturwissenschaftliche Reise nach Mossambique' in 1868. In this publication he provides some more details about this new species which was discovered by him, and of which name he modified to Nothobranchius orthonotus, by erecting a new genus. Peters became the director of the Zoologische Museum der Universität in Berlin and he published around 400 studies and accounts about a variety of animal species.

     The name Nothobranchius is composed of the Greek words νόθος (nothos = false) and βράγχιος (branchion = gill), and refers as false gills to the open preopercular neuromast system which gives the impression of an additional gill opening. Peters also mentions in his description that the locals called the species as 'anamolugo', and also that 'Gebacken gibt er ein vortreffliches Gericht', which may be surprising to Nothobranchius enthusiast because it means that it provides a delicious food when fried.

Nothobranchius orthonotus illustration from Peters 1868
Nothobranchius nov. gen.

     In the foyer of the British Royal Geographical Society in London there are some very interesting relics that are greeting the visitor. In the elegantly furnished lobby there is a time-worn, carefully guarded treasure: the cap worn by David Livingstone when he travelled in the heart of Africa. The cap seems to embody the courage, tenacity, and determination of the original owner. Livingstone was the pre-eminent missionary explorer of the Victorian era who criss-crossed Africa between 1841 and 1873, including much of what is now form part of the large distribution of Nothobranchius fishes. One of the members of Livingston’s second Zambezi expedition from 1858 to 1864 was John Kirk, a doctor naturalist. Sir John also collected Nothobranchius specimens near Quelimane in June 1863. Kirk carried out the collections when he had to wait in Quelimane for the next ship during a period of about one month, before returning home after the expedition.

     At the time when the explorations of the unknown, interior areas of the African continent began, one of the questions with the greatest interest was to search for the source of the Nile. Even the antique world has always been interested in the origin of that river and the ancient Greek geographers were particularly fascinated by this issue. Ptolemy has already known a legend that the Nile originates in the heart of Africa, from three major lakes at the foot of the mountains of the Moon. And the lakes, from where the Nile originates, supposed to be quite far inland. But neither Ptolemy, nor any other Greek traveller has been there; they probably heard about the story from Arab slave traders. This theory has then been verified later.

     In 1862, Speke found at the northern end of Lake Victoria one of the origins of the Nile. Baker discovered the Lake Albert in 1864, and it was found that this is another source of the Nile. Stanley in 1875 found Lake Edward. Another lake and this is also connected to the Nile through the Semliki River, which flows through the Lake Albert. Stanley then discovered the mysterious Moon Mountains, the Ruwenzori, which remained hidden from the eyes of the travellers for very long time due to the constant fog, resulting from the humid air in the surrounding swampy area. The ancient legend became a reality!

Map of the Nile from Coronelli 1690
Fundulus guentheri in Playfair and Guenther 1866

     Zanzibar… The sonorous name of the archipelago was already mentioned by Marco Polo, who never went there personally, but he had heard about the fabulously rich island during his many travels. The 86 km long and 28 km wide main island of Zanzibar is situated in the Indian Ocean, some 37 km away from the coast of the African continent. During the reign of the Sultan of Oman, who has settled down on the island, the largest slave-trading centre was formed there. Given the convenient location of the island, situated just in front of the mainland, countless expedition started from there during the past centuries. Since 1964, the island is united with Tanganyika, located on the opposite mainland, to form the current Tanzania. In 1857, Sir Robert Lambert Playfair collected Nothobranchius specimens on the island of Zanzibar, near the castle of the Sultan of Oman, as well as near Pangani on the opposite mainland a few years later. These collections were recognized as N. orthonotus, the only known species of the genus at that time. The identification of the new collections and the subsequent descriptions of new species have, however, soon begun.

    It is easy to imagine the difficulty to organize an expedition and the condition of the transportation at that time, if we consider that, for instance, the pneumatic tire was invented by John Dunlop only in 1887.

     Franz Stuhlmann, the German zoologist stayed for a long time as an officer in the German army in Africa, before he was appointed to Director of the Institute of Biology and Agriculture in Hamburg. During his stay on Zanzibar, he also collected Nothobranchius species in the island in in 1888. Georg Pfeffer, who worked in the Zoological Museum in Berlin, has prepared the scientific description and named after Albert Günther as Fundulus guentheri in 1893. Günther, the German zoologist and ichthyologist has formerly issued a publication with Playfair about the fishes of Zanzibar.

     The present-day mainland part of Tanzania, formerly called Tanganyika, together with present-day Rwanda and Burundi was a German colonial territory under the name of German East Africa from 1884 to 1918. At that time, several German expeditions had been organized for research purposes but also to strengthen the German colonial influence. One of these expeditions in German East Africa, organized by Franz Stuhlmann and Emin Pasha, was launched in 1889.

     Emin Pasha was a very interesting personality of the era. Born as a son of a Jewish merchant in Germany under the name of Eduard Schnitzer, he converted to the Muslim, and then used the name Emin Effendi. He worked as a doctor in the empire of the Ottomans, and went through many various adventures in Africa. He was appointed to proconsul and pasha in Equatoria, the present-day South Sudan, but he had to escape because of the Mahdi uprising. In order to rescue him from the trapped situation, Stanley organized an expedition, who then made his famous discoveries in relation to the source of the Nile during this voyage. Stanley successfully found Emin Pasha and brought him back to the coast. After their return to Bagamoyo, they attended a reception of the German governor. Emin Pasha, who had very low vision, had fallen out of the window from the first floor, and was later found with bleeding head. After hospitalization, he gladly accepted the invitation to lead an expedition into the interior of the continent. The unfortunate end of Emin Pasha happened in his tent near the Congo River in 1891, when Arab slave traders surprised him and cut his throat.

Fundulus melanospilus in Pfeffer 1896

     The Emin Pasha - Franz Stuhlmann expedition was, however, successful and among others they discovered two fish species that are especially important to us. One day they camped near Farhani, west of Morogoro, and discussed with the Arabs about the recognition of the German colonial power. The La Longa Catholic Mission was situated near the present-day Ilonga. It was there that they found a new fish species in 1890, which was later described as Fundulus melanospilus by Pfeffer in 1896.

     The Emin Pasha - Franz Stuhlmann expedition reached the area of Lake Chaya, despite some troubles they experienced by the Masai tribes along their route. The lake was almost completely dried up, and the remaining water was difficult to reach because of the dense vegetation of reeds and elephant grass. This was however the type locality of another Nothobranchius species, which was first described as Nothobranchius taeniopygus by Hilgendorf in 1891.

     Stuhlmann later found another new Nothobranchius species near Tschangarra in 1892. We had to wait, however, until 1935 for the scientific description when Ahl named the fish as Nothobranchius robustus.

     In the meantime, the first Nothobranchius species was also found in the area of present-day Kenya. Wilson sent specimens to the British Museum in London in 1891. The late Boulenger, the well-known ichthyologist catalogeud them first as Fundulus guentheri. We now know, however, that N. guentheri is present only on Zanzibar, so the species from Mombasa was probably N. elongatus.

     Oscar Neumann arrived at the coastal area of German East Africa in 1892, with hope of an expedition planned with some other participants but that initial plan failed later. Neumann, however, decided that he would try it alone. During the preparation, he managed to collect some N. guentheri on Zanzibar, and an additional species on the opposite mainland near Pangani. Finally, Neumann left to the interior of the continent in 1893. He experienced various adventures along his way. He got into a revolt and rebels injured him with an arrow. The arrow went through his lower jaw and the tongue. The arrow was fortunately not poisoned, but its removal of the arrowhead which had several barbs, was not an easy task. The injuries forced him to look for medical care in Mpwapwa where a German physician was stationed. From there he headed first south and then back to Kwa Mtoro via Tisso. It was on this way between Kwa Mtoro and Tisso that he found a new fish in an area with acacias on August 27, 1893. The species was named after the collector as Fundulus neumanni by Hilgendorf in 1905.

     Italy had also became involved in the process of the East African colonialization, with the occupation of the city of Obbit on the Somali peninsula in 1889. Later, Italy organized its possessions under the name of Italian-Somaliland. In order to strengthen and expand their influence, several expeditions were organized by the Italians. Two of them were led by Captain Vittorio Bottego, a nature lover and army officer. During the second trip, Bottego followed the old caravan route that led from the coastal town of Brava through dry lands towards the Galla plateau, currently in Ethiopia. Because of the intensive drought the expedition was in need of water, so they went to the foot of the close-by Egertha monolith where there are some sources and temporary ponds. This was the place where Captain Bottego found a new Nothobranchius species on October 31, 1895. The species has been described two years later as Fundulus microlepis by Decio Vinciguerra, researcher of the Natural History Museum in Genoa.

     In 1904, Julius Vosseler collected fish specimens between Tanga and Mombo. At that time, Vosseler worked as a zoologist at the Amani Biological and Agricultural Institution in former German East Africa. After his return in Germany, Vosseler became the director of the Hamburg Zoo. Finally, Ahl described Nothobranchius vosseleri on the basis of a preserved female in 1924.

     The Swedish Yngve Sjöstedt led a large Kilimanjaro - Meru expedition in 1905-1906. Sjöstedt was not very happy that they could only organize 80 people to carry the equipment of the expedition. The zoological collections brought back from the expedition comprised over 4300 different species, many of them new to science. Among others, they also found 28 specimens of a new Nothobranchius species in a cocoa plantation near Tanga on June 5, 1905. The species was named by Lönnberg as Fundulus palmqvisti two years later, in honour of Gustaf Palmqvist, the maecenas who financially funded by the whole expedition.

Male of Fundulus guentheri by Arnold 1913
Female of Fundulus guentheri by Arnold 1914

     Species collected until around the turn of the twentieth century hardly arrived alive in Europe. In the early years of that century it has only occurred on rare occasions that Nothobranchius species survived the vicissitudes of a long travel to Europe. In addition, the breeding of annual fish was randomly successful.

     The first reports about successful breeding attempts were in German aquaristic literature published from about 1914. Nothobranchius guentheri, at that time referred to as Fundulus, were spawned over sand and the eggs were collected and water incubated. Fry appeared after a few weeks and larger proportion of the young fish were belly-sliders. The breeding of N. rachovii was reported by using similar methods up to the early 1940s.

     Some insights about the life cycle and ecology of the Nothobranchius fishes were reported by Vanderplank in 1940, when he described the seasonal nature of the natural habitats in the wild. He also mentions that aquarists attempting to breed these fishes would be advised to imitate its natural conditions.

Pair of Fundulus guentheri by Sonn 1914
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