kafuensis: (Latinised adjective) = belonging to or originating from the Kafue River [Zambia] system from where all presently known populations have originated from.

Wildekamp, R. H. & J. Rosenstock (1989) "Anmerkungen zu den Nothobranchius-Arten Sambias, mit der Beschreibung von Nothobranchius kafuensis spec. nov. (Cyprinodontiformes, Nothobranchiinae)" - DATZ 42(7): 413-419; 11 figs, map.

The Terra Typica of the type material is a temporary pool in the Kafue National Park, 5-km south of Chunga Camp, 100-m of the Kafue river, Central Zambia, (26°00'E.-15°03'S).

The, in Zambia located, various collecting localities of this species comprise: Chunga Camp, Kafue National Park, Luansanza, Munkeya, Mazabuka, Nega Nega, Lochinvar Game Reserve, and Kayuni State Farm {10-km north of Monze and 180-km south of Lusaka}. Over the years, some of these populations were introduced into the hobby by Danish hobbyist John Rosenstock during his missions through Zambia.

The species' first description by Wildekamp & Rosenstock in 1989 gave following species' characteristics: dorsal fin having 16 to 18 rays, anal fin 16 to 19 rays, 26 to 31 scales in the longitudinal line series, 2 to 3 scales on the basis of the caudal peduncle, 11 to 12 scales in front of the anal fin around the body. Males possess ctenoïd hooks on the rays of dorsal and anal fins. In both sexes, the beginning of the dorsal fin lies clearly in front of the anal fin origin. The first branchial arch carries 14 branchial rays. In males, the branchiostegal membrane reaches beyond the border of the operculum. Mouth opening directed upwards with prominent lower jaw. Female have a more triangular caudal fin, in which rays 4 to 8 are the longest.

Wildekamp & Rosenstock (1989) studied the morpho-metrical characteristics of several populations. The recorded values as reported by Wildekamp & Rosenstock (1989) are presented in the final table.

According to Wildekamp & Rosenstock (1989), following synonyms can be found in literature:

N. taeniopygus non Hilgendorf, 1891; Bell Cross 1965; Jubb 1981
N. species "Chisanga Loup" Sainthouse, 1985
N. species "Chunga" Sainthouse, 1985
N. species "Kafue River" Sainthouse, 1985
N. species "Kayuni State Farm" Rosenstock, 1981
N. species "Lochinvar" Sainthouse, 1985
N. species "Nega Nega" Rosenstock, 1983
N. species "ZMTW/94 Nanzhila River" (Wood & Ipple, 1994)

All above-mentioned populations belong to the same species-floc, namely N.kafuensis. This was confirmed through crossbreeding experiments, as far back as the fourth generation, descendants continued to produce fertile offspring and by investigations on the karyotype.

Wildekamp & Rosenstock, 1989 consider N.kafuensis as a species closely related to N.polli from Katanga/Shaba province in Zaire/Democratic Republic of Congo. And to an as yet not yet described species originating from the Caprivi Strip [Namibia] (in process by Skelton as reported by Wildekamp & Rosenstock, 1989). They form a closely related group with possibly a common ancestor. They are also related, but to a much lesser extent with N.brieni, N.malaissei from Katanga/Shaba and N.taeniopygus from central Tanzania.

Although N.taeniopygus occurrs on the eastern side of the western arm of the Great Rift, it is the view of Wildekamp & Rosenstock (1989) that a connection could have existed, before the onset of the great East African Rift, over the river systems of the Chambezi, Ruaha, Saisi, Momba and possibly Lake Rukwa and the Luapula system. The existence of a yet undescribed species in the Usangu River [Tanzania], which presents a close relation to N.polli, N.kafuensis and N.spec "Chobe" tends to underline this hypothesis. According to Wildekamp & Rosenstock (1989), the 3 species [N.polli, N.kafuensis and N. spec. "Chobe"] live presently in separated river systems, but their present distribution can be explained with a common ancestor and through an ancient connection between these river systems. In addition, there are indications that the source region of the Upper Lualaba and that of the Upper Zambezi River could have been interconnected over the vast swampy region in Mwinilunga district [around 10° 57' S - 24° 08' E], in northern Zambia (Bell-Cross, 1963; Poll, 1976).

Although one suspects the origin of the Nothobranchius species to lie somewhere in the Upper Nile system, one does not find them in the central Congo River system, where they are replaced by the genus Aphyosemion Myers, 1924. Their occurrence in the Upper Lualaba-Congo basin could possibly be explained by ancient connections between the Upper Zambezi and the Lualaba systems, justifying the relationship between N.polli, N.kafuensis, N. spec "Chobe" and N.taeniopygus.

According to Wildekamp & Rosenstock (1989), there is thus far no proof for a such a recent connection between the Upper Lufira and the Upper Kafue River systems (Bell-Cross, 1968). There are also no indications of a recent connection between the middle Kafue River (along the Kafue Flats) and the Upper Zambezi. An ancient connection between the Kafue Flats and the southern Barotse floodplains, between Namwala [15° 43' S - 26° 32' E] and Sesheke [17° 30' S - 24° 30' E] was reported by Trapnell & Clothier (1937) over the Machili drainage system and its tributary, the Simatanga. The dispersal of Nothobranchius species could possibly have taken place over this ancient connection (Wildekamp & Rosenstock, 1989).

According to Wildekamp (Simoens, 1982), N.polli, N.kafuensis and N. species "Chobe River" would form a series originating from a single evolutive lineage and one could, eventually, arrive at the distinction of three separate sub-species within N.polli.

Nothobranchius sp ZMTW-94 "Nanzhila river" probably also belongs to the same species-floc as they were collected in the same river drainage system. Bowden (1995) is convinced of this thesis after having reproduced several generations of this form.

• ZFMK 13951 - Male (59.9-mm TL; 50.2-mm SL) collected by K. Oestergaard on February 20^th 1983, in a temporary pool in the Kafue National Park, 5-Km. South of Chunga Camp, 100-m of the Kafue River [15° 03' S - 26° 00' E], Zambia.

• ZFMK 13952: Female (45.4-mm TL; 36.0-mm SL) collected together with the holotype;
• MRAC 89-07-P-1: Male (52.4-mm TL; 43.8-mm SL) collected together with the holotype;
• AM PF 1542: Five males (40.7-32.9-mm TL; 34.8-27.9-mm SL) and three females (35.0-34.8-mm TL; 30.0-29.4-mm SL) collected by G. Bell-Cross in May 1964, in a temporary pool in the Kafue River system, Kafue National Park [16° 10' S - 25° 45' E], Zambia, preserved at the Grahamstown Museum Albany, South Africa;

• MRAC 89-07-P-2-5: 3 males (59.9-38.9-mm TL; 50.2-31.6-mm SL) and one female (47.2-mm TL; 39.5-mm SL), collected by J. Rosenstock and K. Oestergaard on December 15^th, 1982, in a temporary pool near Nega Nega, along a sideroad on the right of the main road from Lusaka to Maramba, 18-km from the bridge over the Kafue River in the direction of Mazabuka and 2100 m from the main road.
• MRAC 89-07-P-6-8; 1 male (43.0-mm TL; 35.3-mm SL) and two females (39.7-37.0-mm TL; 32.6-31.3-mm SL), F₁- aquarium generation bred by H. Bitsch from specimens which J. Rosenstock had collected in December 1979 from the Kayuni State Farm, 10-km north of Monze (Zambia).
• MRAC 89-07-P-9; 1 male (43.5-mm TL; 35.2-mm SL), Lochinvar Game Reserve, Zambia, collected in December 1985 by J. Rosenstock.

An administrative map of Zambia is presented in Figure 2. The average meteorological profile in the Lusaka-area where N.kafuensis strives is presented in Figure 3. The dry season extends from Mai to October, the rainy season from November through to April. The hottest season of the year covers the months of October-November, at the beginning of the rainy season. The average drought period in the natural environment lasts on the average for about 5 months. In Zambia, the main crops zones as reflected by the average meteorological profile is presented in Figure 4.

Different populations, which all belong to the same species, are presently known: Chisanga Loup, Chunga Camp, Kafue River [= Kafue National Park], Kayuni State Farm, Lochinvar Game Reserve, Nega Nega, Luasanza, Munkeya, Mazabuka, all presently known collecting sites are located in the Kafue drainage system in Zambia.

They appear in temporary pools, marshes and ditches within the drainage and flood basin of the Kafue River system and incidentally also in that of the upper Zambezi River, Central Zambia (Wildekamp, 1991).

According to Simoens (1982), N. species Kayuni has been found 10-km north of Monze en 180-km south of Lusaka, along the highway towards the Livingstone-Victoria falls, in a temporary pool of 40 x 20-m. In January, the surface reached about 160 a. The pool dries-up during 2-3 months per year. It possibly could have connections with the Magoye River during the rainy season.

According to Bowden (1995), the original collecting site of "Kayuni State Farm" would have been recently destroyed as a consequence of the construction of a dam at the site. Watters (1998) reported that when he visited the area again in 1997, he managed to re-collect this population in three different locations nearby the road. He therefore assumed that there must be many other pools in the area hosting N.kafuensis "Kayuni State Farm".

This is a middle-sized, robust Nothobranchius with several different populations existing in the wild. In the male population two different colour forms [chromo-phenotypes] exist, a red form {Chunga and Nega Nega populations} and a blue phenotype {Kayuni State Farm and Lochinvar Game Reserve populations} [].

Populations originating from the mid-Kafue River system differ slightly from above colour description in so far that their reticulation pattern over the body is more irregular. In these populations, the orange colour on the base of the caudal fin is more dark-brown whilst the reticulation pattern on the base of the anal fin displays more deep-orange and light blue drawings. The orange stripes in anal and caudal fins are quite noticeable.

Females of the various populations can not easily be distinguished from one another. Alive they are lightly colored in yellowish-gray-brown, a little darker over the back, just as is known in most other Nothobranchius species. Scales on the anterior part of the body and on the operculum show a center with sliver reflections. All fins are colorless, at times somewhat brownish and transparent. No special markings are present over the body.

N. kafuensis Chunga (J. Rosenstock, 1985)
N. kafuensis Kayuni State Farm (J. Rosenstock, 1985)
N. kafuensis Lochinvar National Park (J. Rosenstock, 1985)
N. kafuensis Nanzhila River ZMTW/94 (T. Wood & Ippel, 1994)
N. kafuensis Nega Nega ZAM 92/1 (Watters, 1992)
N. kafuensis Kayuni ZAM 97/1 (Watters, 1997)
N. kafuensis Kayuni ZAM 97/9 (Watters, 1997)

Most probably the first discovered Nothobranchius species in Zambia originates from the Kafue River swamps, between the Kafue Bridge and Mazabuka [15° 51' S - 27° 46' E]. According to Wildekamp & Rosenstock (1989), these specimens were collected in March 1955 by P. Evans and were identified by Jubb (1969) as N.taeniopygus. Jubb (1969) gave also a good colour description of the male, which was mainly orange-red. Body scales had a turquoise blue center with a large red-orange marginal band. Anal and caudal fins were also red-orange and presented an orange stripe running over them and a black marginal band. According to Sainthouse (in Wildekamp & Rosenstock, 1989) information kept at the National Museum of Bulawayo, where some specimens from this collection are preserved, indicates that they are considered as descendants of the N.taeniopygus Old Shinyanga-population which had apparently been introduced for malaria control.

About 300-km more to the north-west, a similar male chromo-phenotype was collected in May 1962 by P. B. J. Jackson and G. Bell-Cross in a seasonal pool in a Dembo overflowing into the Munkea (also Munkuye) rivulet [about 14° 53' S - 25° 22' E], not far from the road from Mankoya to Mumba. This rivulet is a tributary of the Upper Zambezi River.

During the same month of May 1962, J. B. Shenton collected a closely related form in the vicinity of the spring of the Luansanza River [15° 30' S - 26° 00' E], in the Kafue River basin, in the Kafue National Park. Since the Shenton collections, P. J. Wright collected on several occasions at the same location. Bell-Cross identified all these Nothobranchius from these collections as N.taeniopygus (Bell-Cross 1965, 1968 and 1974) and regarded them as descendants of the introduced N.taeniopygus Old-Shinyanga populations (Jubb, 1981; Wildekamp 1980).

In 1982, Rosenstock (Rosenstock, 1991) received from Kaj Oestergaard a Nothobranchius population, which looked very similar to the Chunga population. It originated from the Kafue National Park, from a several kilometer-wide area named Chisanga Loup (also called Shishamba Loop). When Rosenstock (1991) visited Zambia in December 1989, the rainy season was delayed and he could not find the exact locality again. On February 20^th 1983, K. Oestergaard collected the species again in a temporary pool, 5-km south of Chunga [15° 03' S - 26° 00' E], about 100-m of the Kafue River and in December 1983 in Chisanga Loup [15° 04' S -26° 00' E], a death arm of the Kafue River, about 12-km of Chunga-Lager.

This chromatic form was again collected near Mazabuka in December 1982 by J. Rosenstock and in 1983 by K. Oestergaard, Danish Development Agency (DANIDA). This new collecting site was a temporary pool in the vicinity of the Kafue River not far away from Nega Nega [15° 45' S - 28° 03' E], about 10-km from the River itself (Rosenstock, 1983).

The form was again collected in Nega Nega in December 1985 but this time in two locations, separated from each other by about 100 -m. The present specimens in the hobby originating from Nega Nega are descendants of the 1982 collection (Wildekamp & Rosenstock, 1989).

In December 1989, Rosenstock (Rosenstock, 1991) re-visited Nega Nega, which he had already visited previously in 1982 and 1985. This locality can easily be found about 80-km south of Lusaka, of which 78-km on good tarmac road, and, 18-km passed the Kafue bridge in the direction of Mazabuka, on the right hand-side along a secondary road for 1 or 2-km. But because of the delayed rains he could not find any Nothobranchius.

All above-mentioned forms constitute in reality two bio-geographical entities: a northern Kafue River system-group (Chunga Lager, Munkeya, Luansanza, Kafue National Park, Chisanga Loup) and a southern Kafue River system-group (Mazabuka and Nega Nega). Cross-breeding experiments between these two geographically distant populations showed that they were not yet genetically isolated and that hybrids of the Chunga x Nega Nega and Chisanga Loup x Chunga populations produced fertile descendants up to the 4^th generation (Wildekamp & Rosenstock, 1989).

At a certain point in time, most authors (Jubb, 1981; Wildekamp, 1980; Sainthouse, 1985) saw in nearly every Nothobranchius species from this part of Africa descendants from the N.taeniopygus "Old Shinyanga" [central Tanzania] population said to have been introduced into several locations during the second World War for malaria control. Therefore Wildekamp & Rosenstock (1989) compared whether N.kafuensis could have close ties with descendants of the N.taeniopygus "Old Shinyanga" population of central Tanzania. They compared the Vanderplank material, collected in Old Shinyanga and preserved in the British Museum for Natural History with specimens from Kayuni State Farm, Nega Nega and Chunga populations. Differences in interorbital width, snout length, caudal peduncle depth and average number of anal and dorsal fin rays was observed besides from differences in caudal fin drawing and colour pattern.

According to Wildekamp and Rosenstock (1989), the initial erroneous identification of this species as N.taeniopygus bij Bell-Cross, Jubb and Wildekamp was mainly based on the male colour pattern of the Old Shinyanga population as presented in Vanderplank's 1940 description. The colour description is however unclear as two different caudal fin colour patterns are presented. In one, it presents a wide orange, in the other a red-brown band running in front of a black marginal border. A later publication by Vanderplank (Vanderplank, 1941) probably provides the correct description of the Old Shinyanga population's caudal fin colour pattern. This colour pattern differs from above mentioned N.kafuensis populations. Following the discovery of N taeniopygus specimens close to the type locality of this species in central Tanzania, Lake Chaya, near Kazi Kazi, by Wildekamp, it became possible to carry out a comparative investigation of both species. As could be expected, both species are very similar and do not present striking morphological differences. However, the difference in male colour pattern is so obvious that Wildekamp and Rosenstock (1989) considered them to be two distinct species. On these grounds, together with the fact that over a period of only 50 years no new species can possibly evolve, it thus appeared that both species could not be considered as closely related or could not have evolved from one another. Therefore Wildekamp & Rosenstock (1989) decided that N.kafuensis constituted a biologically valid new species.

Jubb drew Wildekamp's attention (Wildekamp & Rosenstock, 1989) to a photograph of another phenotype, which had been collected in 1970 bij D. J. Stewart in the Lochinvar Game Reserve, about 60-km west of Mazabuka. Jubb had identified this new colour form as N.taeniopygus (Jubb, 1981). It differed remarkably from the previously discussed colour form. Males from this locality showed a marmorated, at times with transversal stripes, body and fin pattern. Caudal and anal fins presented a wide light blue to white band in front of a marginal black border band. This colour pattern showed some close similarities with a species known from the Upper Lufira system in Katanga, Congo, namely N.polli. Rosenstock re-collected this blue-form in December 1985 in Lochinvar Game Reserve. He had already brought back a similar form in 1979, from the Kayuni State Farm location, 10-km north of Monze [about 27° 30' E - 16° 15' S]. This author provided besides from photographs also a reasonable description of the male colour pattern as well as some ecological information's: the collected specimens appeared in two different colour forms; one showed more blue over the body and in the caudal fin, the second was more red (Rosenstock, 1981).

Another chromo-phenotype having a male colour pattern similar to the above discussed blue form was collected on several occasions, between 1973 and 1977 by B. van der Waal in Chobe and in the floodplains of the Upper Zambezi River, in the eastern Namibian Caprivi Strip. In general these specimens are lighter and their stripes in anal and caudal fins are more light and yellowish. According to Wildekamp & Rosenstock (1989), Jubb identified these last specimens as N.brieni (Jubb, 1976; Van der Waal, 1976), but after the re-description of N.brieni by Wildekamp in 1978, these specimens were identified as N. spec. "Chobe" (Wildekamp, 1978; Jubb, 1981; Van der Waal & Skelton 1984; Skelton, 1987). In the "Catalogue of Freshwater Fishes of Africa - Cyprinodontidae" from Wildekamp, Romand and Scheel, published in 1986, the blue N.kafuensis population of Kayuni State Farm and Chobe River are considered as populations of N.polli.

Rosenstock expedited specimens of the F₁ and F₂ aquarium generations of the Nega Nega and Chunga populations as well as specimens of the Kayuni State Farm and Lochinvar populations to J. J. Scheel for analysis of the karyotypes. In the mean time, eggs of the Kayuni State Farm, Nega Nega and Chunga populations were send by Wildekamp to E. Krysanov, Russia. Both investigators came to the same conclusion: the studied populations presented the same karyotype of n=18 haploid ([diploid] 2n=36) metacentric chromosomes. This showed that both colour forms were closely related.

As both phenotypes had similar karyotypes, Wildekamp (1989) initiated crossbreeding experiments with the assistance of Théo Steinfort in order to discover possible genetic differences between the blue and red forms. Hybrids of the crossbreeding experiment between Kayuni State Farm x Chunga population were fertile over 3 generations, indicating the absence of any genetic shift between both forms. Wildekamp & Rosenstock (1989) report that males of the F₁ crossbred generation displayed an apparently uniform body pattern, whereby the general body colour corresponded to the Kayuni State Farm form whilst the colour pattern of anal and caudal fins belonged to the Chunga population. Males of the F₂ and F₃ crossbred generations displayed a clear differentiation in red and blue chromo-phenotypes. In general, males of the red phenotypes corresponded to males of the original Chunga form, whilst males of the blue phenotype corresponded to males of the Kayuni State Farm form. Also from these crossbreeding experiments and from the karyotype investigations, Wildekamp & Rosenstock (1989) concluded that both forms represented in fact one single species, with two distinct chromo-phenotypes as could be observed in N.eggersi, N.jubbi, N.ugandensis, and N.korthausae.

In May-end 1994, Trevor Wood and Johann Ipple (Wood, 1994) traveled from South Africa through Botswana to Zambia in search of N. spec. "Chobe" and N. spec. "Caprivi". At the level of Kalomo in Zambia, they turned northwest towards the Kafue National Park. It took them more than 7 hours to travel the 190-km on a two-wheel track through the bush. The night temperatures were chilly, about 5°C, rising to about 28°C at noon, with water temperature being also low 18°C or less (Wood, 1994). At a point, approximately 20-km inside the Kafue National Park, they found 2 adjacent ponds on the left-side of the track. One pond on the right had dried-out. In these ponds they found Barbs and Cichlids but also nine pairs and a few spare females of a new Nothobranchius form [N. spec. ZMTW/94 Nanzhila River]. As it became drier more north, they eventually decided to turn back.

According to Trevor Wood (1994), a dry riverbed was crossed 10-km north of the biotope. This was a tributary of the Nanzhila River which itself flows into the Kafue River, 90-km more to the north. It would then seem that there could be a direct link between N. spec. ZMTW/94 Nanzhila River form and the N.kafuensis species previously mentioned. The discovery of N. spec. ZMTW/94 Nanzhila River added to the knowledge of the genus in this area, as it lies midway between N. spec. Chobe River and N.kafuensis. Since then, the N. spec. ZMTW/94 Nanzhila River form was identified as N.kafuensis ZMTW/94 Nanzhila River.

This last form has in fact a very similar look to N.kafuensis. Compared to N.kafuensis Kayuni State Farm, there is the same typical outer black edge and inner pale blue, almost white, submarginal band on the caudal and anal fins. The dorsal and body have the same reticulated patterns. However, the Kayuni fish has more red to the front half of the body, whereas ZMTW/94 displays little sign of red, being mainly green/blue dependent on the light source (Wood, 1994). The pectorals are clear at their base, suffusing to violet at the outer edge. The ventrals are pale blue. The other significant difference lies in the anal fin in which the Kayuni form exhibits a diffused pale blue horizontal line close to the body. This line is absent in the ZMTW/94 form.

According to Simoens (1982), the breeding and maintenance of this species does not present any particular problems. A 30-litre aquarium is already sufficient for a trio. However, the species appears to be sensitive to Oödinium (velvet), which can be fought with the addition of kitchen salt to the water {30 gr. per 10 liters}. Rannou (1990) mentions the addition of 1 teaspoon of salt per 5 liters of water. Ensuring a continuous gentle water flow in the tank with a small powerhead pump can also combat this disease.

Walker (1995) mentions an incubation period of only 2 months whilst hatching the eggs can easily take-up to 4 hours.

At hatching time, the fry is relatively large and can be fed immediately with freshly hatched Artemia nauplii. Around 1.5 to 2 months of age, one can already identify the sexes.

Some hobbyists mention as problem with breeding this species, and more particularly in raising the fry, that it could happen that suddenly, in one single day, all fry become belly-sliders. This is possibly being caused by a kind of viral or bacterial disease of the swim bladder or by a physiological circumstance possibly related to stress. This problem can be avoided by applying a rigorous hygiene to the tank. When raising Nothobranchius species, one need to change regularly part of the water (some breeders mention up to 20% of the tank content per day - which seems too much to us). Other authors mention sudden fierce waves of panic amongst the fry and sub-adult fishes causing large abrupt mortality levels. For this species one should thus always use tanks with plenty of aquatic plants and located in quiet places and approach the tank with great tranquillity.

As alternative to the addition of kitchen salt to the tank to preventively combat Oödinium (velvet) some authors introduce a fine naked strip of copper of about 10-cm long in the water. In this case, one should surely not add any kitchen salt to the water. The copper strip releases progressively the lethal copper ions into the water. Concentrations remain however very weak thanks to the regular water changes but are nevertheless lethal enough to kill most of the parasites and does not appear to have not negative effects on the fishes (which by the way originate from the so called "Copper Belt" in Zambia). When employing this method, one has however to remain very vigilant because copper still remains very poisonous.

Other authors (Rannou, 1990) place, in case of severe Oödinium attack, (only) adult fish in a temporary bath of copper-sulphate {5 drops per 10 cl. of a 1% copper-sulphate solution during 5 minutes}. The "Oödinium-carpet" can be seen to fall off the bathing fish. Such a drastic treatment can only be done with strong fishes and surely not very often.

According to Trevor Wood, the discoverer of N.spec.ZMTW-94 "Nanzhila River", the males of this population are particularly aggressive toward one another. Bowden (1995) however did not seem to be confronted with this difficulty.

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Table 1: Morphological data expressed in % of Standard Length (Source: Wildekamp & Rosenstock, 1989)