Exceptional levels of species discovery ameliorate inferences of the biogeography and diversification of an Afrotropical catfish family.

Endeavours in species discovery, particularly the characterisation of cryptic species, have been greatly aided by the application of DNA molecular sequence data to phylogenetic reconstruction and inference of evolutionary and biogeographic processes. However, the extent of cryptic and undescribed diversity remains unclear in tropical freshwaters, where biodiversity is declining at alarming rates. To investigate how data on previously undiscovered biodiversity impacts inferences of biogeography and diversification dynamics, we generated a densely sampled species-level family tree of Afrotropical Mochokidae catfishes (220 valid species) that was ca. 70 % complete. This was achieved through extensive continental sampling specifically targeting the genus Chiloglanis a specialist of the relatively unexplored fast-flowing lotic habitat. Applying multiple species-delimitation methods, we report exceptional levels of species discovery for a vertebrate genus, conservatively delimiting a staggering ca. 50 putative new Chiloglanis species, resulting in a near 80 % increase in species richness for the genus. Biogeographic reconstructions of the family identified the Congo Basin as a critical region in the generation of mochokid diversity, and further revealed complex scenarios for the build-up of continental assemblages of the two most species rich mochokid genera, Synodontis and Chiloglanis. While Syndontis showed most divergence events within freshwater ecoregions consistent with largely in situ diversification, Chiloglanis showed much less aggregation of freshwater ecoregions, suggesting dispersal as a key diversification process in this older group. Despite the significant increase in mochokid diversity identified here, diversification rates were best supported by a constant rate model consistent with patterns in many other tropical continental radiations. While our findings highlight fast-flowing lotic freshwaters as potential hotspots for undescribed and cryptic species diversity, a third of all freshwater fishes are currently threatened with extinction, signifying an urgent need to increase exploration of tropical freshwaters to better characterise and conserve its biodiversity.

Diversity, distribution and extinction risk of native freshwater fishes of South Africa.

Extinction risk for 101 valid species and 18 unique genetic lineages of native freshwater fishes of South Africa was assessed in 2016 following the IUCN Red List criteria. An additional five species (three new species that were described and two species that were revalidated subsequent to the 2016 assessments) were assessed in the present study. A synthesis of the outcome of the assessments of the 106 valid species and 18 genetic lineages indicates that 45 (36%) of South Africa's freshwater fish taxa are threatened (7 Critically Endangered, 25 Endangered, 13 Vulnerable). Of the remaining taxa, 17 (14%) are listed as Near Threatened, 57 (46%) are Least Concern and five (4%) are Data Deficient. More than 60% of the endemic taxa are threatened. The Cape Fold Ecoregion has the highest proportion of threatened taxa (67%) due to the existence of a unique assemblage of narrow-range endemic species. Galaxias and Pseudobarbus have the highest number of highly threatened taxa as most of the species and lineages in these genera are classified as either CR or EN. Major threats to the native freshwater fishes of the country are invasive fish species, deterioration of water quality, impoundments and excessive water abstraction, land use changes and modification of riverine habitats. Immediate conservation efforts should focus on securing remnant populations of highly threatened taxa and preventing deterioration in threat status, because recovery is rare. Accurate delimitation of species boundaries, mapping their distribution ranges, improved knowledge of pressures and long-term monitoring of population trends need to be prioritised to generate credible data for the 2026 IUCN threat status assessments and designation of important fish areas as part of the National Freshwater Ecosystem Priority Areas (NFEPA) initiative.

Utility of DNA barcoding in native Oreochromis species.

The importance of Oreochromis in worldwide aquaculture and regional fisheries motivates the study of their genetic diversity in their native range. In this article, all mitochondrial cytochrome c oxidase subunit I gene (COI) sequences of Oreochromis species are retrieved from Barcode of Life Data system to quantify the available DNA barcoding information from wild individuals collected within the native ranges of the respective species. It is found that 70% of the known species in the genus still lack a COI barcode, and only 15% of the available sequences are from within the respective native ranges. Many of the available sequences have been produced from specimens acquired from aquaculture and introduced, naturalized populations, making the assessment of variation within the original native range challenging. Analyses of the wild-collected fraction of available sequences indicated the presence of cryptic lineages within Nile tilapia Oreochromis niloticus and O. schwebischi, the occurrence of potential introgressive hybridization between O. niloticus and blue tilapia O. aureus, and potential ancestral polymorphism between Karonga tilapia O. karongae and black tilapia O. placidus. This article also reports a case of misidentification of O. mweruensis as longfin tilapia O. macrochir. These results stress the importance of improving the knowledge of genetic variation within the native ranges of Oreochromis species for better-informed conservation of these natural resources.

Enteromius thespesios (Teleostei: Cyprinidae): a new minnow species with a remarkable sexual dimorphism from the south-eastern part of the Upper Congo River.

A new minnow species, Enteromius thespesios, is described from the south-eastern part of the upper Congo River; that is, the Kalule Nord, the Luvilombo and the Chambeshi Rivers. Enteromius thespesios belongs to the group of the soft-rayed species of Enteromius from the Congo Basin; that is, those with a weakly ossified, flexible last unbranched dorsal-fin ray that lacks serrations along its posterior edge. Within this group, E. thespesios is most similar to E. humeralis, from which it is distinguished by a higher number of circumpeduncular scales and shorter anterior and posterior barbels. Enteromius thespesios is a rheophilic and territorial species. It exhibits a marked sexual dimorphism, with males having: a red band towards the distal edge of dorsal, caudal and, to a lesser degree, anal fin; nuptial tubercles; a longer snout; longer pectoral fins; a shorter anal fin. This study gives extensive consideration to sexual shape differences for a species of Enteromius and also briefly reviews the current knowledge of sexual dimorphism in the species of Enteromius from the Congo Basin. Some conservation issues related to the new species are also highlighted.

Molecular phylogeny of Oreochromis (Cichlidae: Oreochromini) reveals mito-nuclear discordance and multiple colonisation of adverse aquatic environments.

Although the majority of cichlid diversity occurs in the African Great Lakes, these fish have also diversified across the African continent. Such continental radiations, occurring in both rivers and lakes have received far less attention than lacustrine radiations despite some members, such as the oreochromine cichlids (commonly referred to as 'tilapia'), having significant scientific and socio-economic importance both within and beyond their native range. Unique among cichlids, several species of the genus Oreochromis exhibit adaptation to soda conditions (including tolerance to elevated temperatures and salinity), which are of interest from evolutionary biology research and aquaculture perspectives. Questions remain regarding the factors facilitating the diversification of this group, which to date have not been addressed within a phylogenetic framework. Here we present the first comprehensive (32/37 described species) multi-marker molecular phylogeny of Oreochromis and closely related Alcolapia, based on mitochondrial (1583 bp) and nuclear (3092 bp) sequence data. We show widespread discordance between nuclear DNA and mitochondrial DNA trees. This could be the result of incomplete lineage sorting and/or introgression in mitochondrial loci, although we did not find a strong signal for the latter. Based on our nuclear phylogeny we demonstrate that adaptation to adverse conditions (elevated salinity, temperature, or alkalinity) has occurred multiple times within Oreochromis, but that adaptation to extreme (soda) conditions (high salinity, temperature, and alkalinity) has likely arisen once in the lineage leading to O. amphimelas and Alcolapia. We also show Alcolapia is nested within Oreochromis, which is in agreement with previous studies, and here revise the taxonomy to synonymise the genus in Oreochromis, retaining the designation as subgenus Oreochromis (Alcolapia).

Nocturnal claroteine catfishes reveal dual colonisation but a single radiation in Lake Tanganyika.

Lake Tanganyika (LT) is a biodiversity hotspot supporting many endemic radiations that provide comparative systems in which to investigate if there are common factors leading to the build-up of its considerable diversity. Despite LT containing the highest diversity of lacustrine catfishes on Earth, the evolutionary relationships of nocturnal catfishes within the sub-family Claroteinae have not been investigated and it is unknown if its constituent genera have diversified via single or independent colonisation events. We report the first molecular phylogeny of the LT claroteine catfishes based on a multigene dataset (three nuclear markers, two mitochondrial totalling 4227 bp), including 85 samples from LT and outside of the lake basin. These data support LT claroteine monophyly, with the exclusion of the LT endemic Chrysichthys brachynema that independently colonised the lake but has not radiated. Multiple sampling localities from LT and the use of Bayesian species delimitation methods reveal additional locally restricted diversity within the LT Claroteinae clade. Fossil calibrated molecular divergence dates suggest that diversification occurred within full lake conditions as demonstrated in other LT lineages.

Continental diversification of an African catfish radiation (Mochokidae: Synodontis).

Despite African rivers containing high species diversity, continental-scale studies investigating the mechanisms generating biological diversity of African riverine faunas are limited compared with lacustrine systems. To investigate the build-up of diversity in a tropical aquatic continental radiation, we test different models of lineage diversification and reconstruct the biogeographic history in a species-rich siluriform genus, Synodontis (~130 species), with a broad distribution across all major tropical African drainage basins. The resulting robust species-level phylogeny (~60% complete, based on a multigene data set) exhibits a near constant rate of lineage accumulation throughout the mid-Cenozoic to recent, irrespective of missing species and despite the changing environmental conditions that were prevalent during this time period. This pattern contrasts with the findings for species-level diversification of large clades that commonly show an early burst of cladogenesis followed by declining rates through time. The identification of distinct biogeographic clades demonstrates a correlation between river hydrology and cladogenesis, although there is evidence of recent repeat dispersal into the southern range of the focal group. We conclude that diverse freshwater fish radiations with tropical continental distributions represent important organisms to test hypotheses of diversification and investigate the effects of palaeo-landscapes and climates on present day biodiversity.

An extant cichlid fish radiation emerged in an extinct Pleistocene lake.

The haplochromine cichlid fish of the East African Great Lakes represent some of the fastest and most species-rich adaptive radiations known, but rivers in most of Africa accommodate only a few morphologically similar species of haplochromine cichlid fish. This has been explained by the wealth of ecological opportunity in large lakes compared with rivers. It is therefore surprising that the rivers of southern Africa harbour many, ecologically diverse haplochromines. Here we present genetic, morphological and biogeographical evidence suggesting that these riverine cichlids are products of a recent adaptive radiation in a large lake that dried up in the Holocene. Haplochromine species richness peaks steeply in an area for which geological data reveal the historical existence of Lake palaeo-Makgadikgadi. The centre of this extinct lake is now a saltpan north of the Kalahari Desert, but it once hosted a rapidly evolving fish species radiation, comparable in morphological diversity to that in the extant African Great Lakes. Importantly, this lake seeded all major river systems of southern Africa with ecologically diverse cichlids. This discovery reveals how local evolutionary processes operating during a short window of ecological opportunity can have a major and lasting effect on biodiversity on a continental scale.

Four new species of Serranochromis (Teleostei: Cichlidae) from the Cuanza and Okavango river systems in Angola, including a preliminary key for the genus.

The present study describes Serranochromis alvum n. sp., Serranochromis swartzi n. sp., Serranochromis cuanza n. sp., and Serranochromis cacuchi n. sp. from Angolan tributaries of the Cuanza and Okavango systems in Angola. The presence of four or five scale rows between the posterior margin of the orbit and the ascending arm of the preoperculum, the presence of widely set unicuspid teeth on the jaws, widely separated gill rakers, and anal fins with egg ocelli place these four species in Serranochromis. The Serranochromis described herein are distinguishable based on a combination of morphological and meristic characters, as well as pigmentation patterns. The interorbital width (14.3-15.9 % HL) of S. alvum is narrower than that of S. swartzi (17.6-19.8), S. cuanza (16.3-18.0), and S. cacuchi (20.0-21.7). Moreover, the interorbital width of S. cacuchi is greater than the other three described species. Serranochromis swartzi has a smaller preorbital depth (16.2-18.9 % HL) and snout length (29.6-31.9 % HL) than Serranochromis cuanza (PD 19.1-22.2, SNL 35.2-39.6 % HL). Serranochromis alvum is known only from the type locality at Cuito-Cuanavale at the junction of the Cuito and Cuanavale rivers, tributary to the Okavango River in Angola. Serranochromis swartzi is known only from the type locality in the Cuanza River, Angola. Serranochromis cuanza is restricted to the Cuanza River, below Capanda Dam, Angola, while S. cacuchi is known only from the Cacuchi River, a tributary of the Cuchi-Cubango River in Angola. The limited distribution of all four species and the absence of many congeners suggest, that in addition to previous studies that invoked a lacustrine speciation model, vicariance through drainage isolation seems to have played an important role in driving speciation in this group. The minimum polygon clusters that are formed when the first principal components of the meristic data are plotted against the second sheared principal components of the morphometric data show separation of the four new species.

Mastacembelid eels support Lake Tanganyika as an evolutionary hotspot of diversification.

BACKGROUND: Lake Tanganyika (LT) is the oldest of the African Rift Lakes and is one of the richest freshwater ecosystems on Earth, with high levels of faunal diversity and endemism. The endemic species flocks that occur in this lake, such as cichlid fishes, gastropods, catfish and crabs, provide unique comparative systems for the study of patterns and processes of speciation. Mastacembelid eels (Teleostei: Mastacembelidae) are a predominately riverine family of freshwater fish, occurring across Africa and Asia, but which also form a small species flock in LT. METHODS: Including 25 species across Africa, plus Asian representatives as outgroups, we present the first molecular phylogenetic analysis for the group, focusing particularly on the evolutionary history and biodiversity of LT mastacembelid eels. A combined matrix of nuclear and mitochondrial genes based on 3118 bp are analysed implementing different phylogenetic methods, including Bayesian inference and maximum likelihood. RESULTS: LT Mastacembelus are recovered as monophyletic, and analyses reveal the rapid diversification of five main LT lineages. Relaxed molecular clock dates provide age estimates for the LT flock at approximately 7-8 Myr, indicating intralacustrine diversification, with further speciation events coinciding with periods of lower lake level. Our analyses also reveal as yet undescribed diversity of lacustrine and riverine species. A Southern-Eastern African clade, that is younger than the LT flock, is also recovered, while West African taxa are basal members of the African mastacembelid clade. CONCLUSIONS: That the LT species flock of mastacembelid eels appears to have colonised and immediately diversified soon after the formation of the lake, supports the view of LT as an evolutionary hotspot of diversification. We find evidence for biogeographic clades mirroring a similar pattern to other ichthyological faunas. In addition, our analyses also highlight a split of African and Asian mastacembelid eels at approximately 19 Myr that is considerably younger than the split between their associated continents, suggesting a dispersal scenario for their current distribution.

Re-elevation to species level and redescription of Serranochromis jallae and Serranochromis robustus (Teleostei: Cichlidae).

Serranochromis robustus robustus from Lake Malawi and Serranochromis robustus jallae from Zambia were compared using morphological data. We re-elevated S. robustus jallae to species based on the following. Serranochromis robustus generally has a longer lower jaw (50.7-59.6% HL) than S. jallae (49.2-52.7% HL). Serranochromis robustus is not as deep-bodied as S. jallae as evidenced by the distance between the posterior insertion of the dorsal fin and the posterior insertion of the anal fin (13.4-15.1% SL in S. robustus vs. 14.9-18.4% SL in S. jallae). Additionally, S. robustus has a narrower least caudal peduncle depth (10.9-12.8% SL) than S. jallae (11.3-14.2% SL); the least caudal peduncle depth of all S. robustus was less than 12.8% SL while, except for the smallest specimen of S. jallae (88.1 mm SL), the least caudal peduncle depth was greater than 13.2% SL. Adults in breeding color of Serranochromis robustus are blue/green laterally, while adults in breeding color of S. jallae are yellow/green laterally. The marginal bands on the dorsal and caudal fins of S. jallae are bright orange in specimens from the Okavango River system and creamy yellow in Upper Zambezi specimens. In S. robustus, there is a yellow marginal band on the dorsal fin.

Diversity of the southern Africa Lacustricola Myers, 1924 and redescription of Lacustricola johnstoni (Günther, 1894) and Lacustricola myaposae (Boulenger, 1908) (Cyprinodontiformes, Procatopodidae).

Through the analysis of a comprehensive database of COI sequences, with the sequencing of 48 specimens, a first insight into the genetic diversity, distribution and relationships between the southern Africa "Lacustricola" species is presented. Species from "Lacustricola" occur mainly in freshwater systems within the arid savanna, and are considered to be widely distributed in southern Africa, but most of them are data deficient taxa. Two species are redescribed, "Lacustricola" johnstoni (Günther, 1894) and "Lacustricola" myaposae (Boulenger, 1908), based on specimens collected at their respective type localities. Detailed osteological and life colouration information is presented for the first time. "Lacustricola" johnstoni was described from the Upper Shire River in Mangochi, Lake Malawi but is herein considered as widespread in the Okavango, Zambezi, southern Africa east coastal drainages and the Bangweulu in the Congo System. A sympatric similar species occurring in the Okavango is also identified. "Lacustricola" myaposae (Boulenger, 1908), was described from the Nseleni River in KwaZulu-Natal Province, South Africa and is herein considered to be endemic to the small coastal river drainages within this region. Lectotypes for both "L." johnstoni and "L." myaposae are designated. A new species from the Lualaba River in the Congo System, sister to "L." macrurus is identified, and the deep bodied "L." jubbi is considered sister taxon to a clade including "L." johnstoni and "L." myaposae.

Nuclear markers reveal unexpected genetic variation and a Congolese-Nilotic origin of the Lake Victoria cichlid species flock.

Phylogenetic analyses based on mitochondrial (mt) DNA have indicated that the cichlid species flock of the Lake Victoria region is derived from a single ancestral species found in East African rivers, closely related to the ancestor of the Lake Malawi cichlid species flock. The Lake Victoria flock contains ten times less mtDNA variation than the Lake Malawi radiation, consistent with current estimates of the ages of the lakes. We present results of a phylogenetic investigation using nuclear (amplified fragment length polymorphism) markers and a wider coverage of riverine haplochromines. We demonstrate that the Lake Victoria-Edward flock is derived from the morphologically and ecologically diverse cichlid genus Thoracochromis from the Congo and Nile, rather than from the phenotypically conservative East African Astatotilapia. This implies that the ability to express much of the morphological diversity found in the species flock may by far pre-date the origin of the flock. Our data indicate that the nuclear diversity of the Lake Victoria-Edward species flock is similar to that of the Lake Malawi flock, indicating that the genetic diversity is considerably older than the 15 000 years that have passed since the lake began to refill. Most of this variation is manifested in trans-species polymorphisms, indicating very recent cladogenesis from a genetically very diverse founder stock. Our data do not confirm strict monophyly of either of the species flocks, but raise the possibility that these flocks have arisen from hybrid swarms.