Taxonomic diversity, distribution, and ecology of the freshwater fishes of the Zambezian Lowveld Ecoregion in southern Africa: A systematic review.

The Zambezian Lowveld Ecoregion (ZLE) is one of the 22 freshwater ecoregions covering southern Africa. This ecoregion covers ~520,418 km2 and extends from south of the Zambezi Delta in the north to the uMngeni River basin in the south. This study aimed to compile a comprehensive synthesis of the available information on the diversity and distribution of freshwater fishes in this ecoregion based on natural history collection holdings by reviewing the published scientific literature. In total, 105 native species belonging to 39 genera and 17 families were recorded from the ZLE. An important proportion, 20 (19%), of the species are considered to be narrow-range endemics. Nevertheless, the majority of them (81%) are considered to have broad geographic ranges, with some even extending into adjacent ecoregions. However, recent and ongoing studies indicate that the current taxonomy often underestimates the species diversity and therefore overestimates the distribution ranges of the latter group of freshwater fishes concerned. The present synthesis brings to the fore existing knowledge gaps in species diversity and distribution ranges of freshwater fishes of this ecoregion and highlights the need for the use of integrative approaches to address the prevailing taxonomic conflicts.

Naming the other cousin: A new goldie barb (Cyprinidae: Smiliogastrininae) from the northeast escarpment in South Africa, with proposed taxonomic rearrangement of the goldie barb group in southern Africa.

A growing body of evidence indicates that the global diversity of freshwater fishes has not been fully documented. Studies of freshwater fishes that were previously thought to be morphologically variable have revealed the existence of deeply divergent lineages, with many distinct species. In southern Africa a number of Enteromius species exhibit either exceedingly wide or divided distribution patterns that should be rare for freshwater fishes with limited dispersal opportunities between river systems. One such species is the sidespot barb, Enteromius neefi. As currently defined, E. neefi has a disjunct distribution that is divided between rivers in the northeast escarpment in South Africa and Eswatini, and tributaries of the Upper Zambezi in Zambia and southern Congo in the Democratic Republic of Congo, with a large geographic gap between these two populations. With the use of molecular and morphological methods, the level of divergence between the two populations was examined, and a new species was described from the Steelpoort River in the Limpopo River system of South Africa. Findings from this study provide further evidence for a number of taxonomic problems within the goldie barbs of southern Africa, and some taxonomic rearrangements are proposed for this group.

The Complete Genome Sequences of 12 Species of Enteromius (Cyprinidae, Cypriniformes, Actinopteri).

We present the complete genome sequences of 12 species of Enteromius. Illumina sequencing was performed on genetic material from museum specimens. The reads were assembled using a de novo method followed by a finishing step. The raw and assembled data are publicly available via Genbank.

Hidden in the riffles: A new suckermouth catfish (Mochokidae, Chiloglanis) from the middle Zambezi River system, Zimbabwe.

The recent surge in the discovery of hidden diversity within rheophilic taxa, particularly in West and East Africa, prompted a closer examination of the extent to which the current taxonomy may obscure the diversity of riffle-dwelling suckermouth catfishes in the genus Chiloglanis in southern Africa. Currently, the region comprises eight valid species within this genus. Seven of them have relatively narrow geographic distribution ranges except for C.neumanni, which is considered to be widely distributed, occurring from the Buzi River system in the south, and its northern limit being the eastward draining river systems in Tanzania. Recent surveys of the middle Zambezi River system revealed Chiloglanis specimens that were distinguishable from the known species of the genus from southern Africa. Integration of molecular and morphological data indicated that these specimens from the Mukwadzi River represent a new species to science, herein described as Chiloglaniscarnatus Mutizwa, Bragança & Chakona, sp. nov. This species is readily distinguished from its southern African congeners by the possession of a distinctive extended dermal tissue covering the base of the dorsal fin and the possession of ten mandibular teeth (vs 8, 12, or 14 in the other taxa). Results from this study add to the growing evidence of a high level of undocumented diversity within riffle-dwelling taxa in southern Africa.

A new species of Parauchenoglanis (Auchenoglanididae: Siluriformes) from the Upper Lualaba River (Upper Congo), with further evidence of hidden species diversity within the genus.

Parauchenoglanis zebratus sp. nov. is a new species endemic to the Upper Lualaba in the Upper Congo Basin. It is distinguished from all its congeners known from the Congo Basin and adjacent basins by the presence of (1) distinctive dark-brown or black vertical bars on the lateral side of the body, at least for specimens about ≥120 mm LS , (2) a broad and triangular humeral process embedded under the skin and (3) a well-serrated pectoral-fin spine. Genetic analysis based on mtDNA COI sequences confirmed the genetic distinctiveness (2.8%-13.6% K2P genetic divergence) of P. zebratus sp. nov. from congeners within the Congo and adjacent river basins. The study also revealed additional undocumented diversity within P. ngamensis, P. pantherinus, P. punctatus and P. balayi, indicating the need for further in-depth alpha-taxonomic attention to provide more accurate species delimitations for this genus. The discovery of yet another new species endemic to the Upper Lualaba, and this well outside the currently established protected areas, highlights the critical need for further assessments to accurately document the species diversity to guide freshwater conservation prioritisation and biodiversity management in this region.

Nothobranchius balamaensis (Cyprinodontiformes: Nothobranchiidae), a new species of annual killifish from northern Mozambique.

A new seasonal killifish of the genus Nothobranchius is described from the Montepuez River system in northern Mozambique. The new species, Nothobranchius balamaensis Bragana Chakona, is differentiated from congeners by its characteristic colour pattern and molecular data further support its taxonomic distinctiveness. Phylogenetic results based on two mitochondrial and three nuclear genes confirms N. balamaensis is closely related to N. kirki and N. wattersi; all three belonging to the Coastal-Inland Clade. The new species is most similar in colour pattern to N. kirki sharing the characteristic of a deep red-orange colouration in the basal, proximal, and medial zones of the caudal and anal fins that grades to orange in the distal zone. This is a key feature that distinguishes these two species from all other Nothobranchius. The main distinguishing features between the new species and N. kirki is the presence of a light blue to white band or series of irregular markings in the proximal zone of the anal fin in N. kirki, versus the absence of such a colour pattern element in N. balamaensis, as well as differences in the dorsal fin pattern. When compared to all population groups of N. wattersi, the colour pattern of N. balamaensis is distinctive. Nothobranchius balamaensis is a relatively slender member of the genus, a characteristic that clearly distinguishes it from both N. kirki and N. wattersi. Nothobranchius balamaensis is currently only known from a few specimens from the type locality.

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.

New phylogenetic insights into the African catfish families Mochokidae and Austroglanididae.

Several hundred catfish species (order: Siluriformes) belonging to 11 families inhabit Africa, of which at least six families are endemic to the continent. Although four of those families are well-known to belong to the 'Big-Africa clade', no previous study has addressed the phylogenetic placement of the endemic African catfish family Austroglanididae in a comprehensive framework with molecular data. Furthermore, interrelationships within the 'Big-Africa clade', including the most diverse family Mochokidae, remain unclear. This study was therefore designed to help reconstruct inter- and intrarelationships of all currently valid mochokid genera, to infer their position within the 'Big Africa clade' and to establish a first molecular phylogenetic hypothesis of the relationships of the enigmatic Austroglanididae within the Siluriformes. We assembled a comprehensive mitogenomic dataset comprising all protein coding genes and representing almost all recognized catfish families (N = 33 of 39) with carefully selected species (N = 239). We recovered the monophyly of the previously identified multifamily clades 'Big Asia' and 'Big Africa' and determined Austroglanididae to be closely related to Pangasiidae, Ictaluroidea and Ariidae. Mochokidae was recovered as the sister group to a clade encompassing Auchenoglanididae, Claroteidae, Malapteruridae and the African Schilbeidae, albeit with low statistical support. The two mochokid subfamilies Mochokinae and Chiloglanidinae as well as the chiloglanid tribe Atopochilini were recovered as reciprocally monophyletic. The genus Acanthocleithron forms the sister group of all remaining Mochokinae, although with low support. The genus Atopodontus is the sister group of all remaining Atopochilini. In contrast to morphological reconstructions, the monophyly of the genus Chiloglanis was strongly supported in our analysis, with Chiloglanis macropterus nested within a Chiloglanis sublineage encompassing only other taxa from the Congo drainage. This is an important result because the phylogenetic relationships of C. macropterus have been controversial in the past, and because we and other researchers assumed that this species would be resolved as sister to most or all other members of Chiloglanis. The apparent paraphyly of Synodontis with respect to Microsynodontis provided an additional surprise, with Synodontis punu turning out to be the sister group of the latter genus.

Mitogenome recovered from a 19 th Century holotype by shotgun sequencing supplies a generic name for an orphaned clade of African weakly electric fishes (Osteoglossomorpha, Mormyridae).

Heteromormyrus Steindachner, 1866, a genus of Mormyridae (Teleostei: Osteoglossomorpha), has been monotypic since the description of Heteromormyruspauciradiatus (Steindacher, 1866) from a single specimen. No type locality other than "Angola" was given and almost no specimens have been subsequently identified to this species. In order to investigate the relationship of this taxon to fresh specimens collected in Angola and elsewhere, whole genome paired-end sequencing of DNA extracted from the holotype specimen of Heteromormyruspauciradiatus was performed and a nearly complete mitogenome assembled from the sequences obtained. Comparison of cytochrome oxidase I and cytochrome b sequences from this mitogenome to sequences from recently collected material reveal that Heteromormyruspauciradiatus is closely related to specimens identified as Hippopotamyrusansorgii (Boulenger, 1905), Hippopotamyrusszaboi Kramer, van der Bank & Wink, 2004, Hippopotamyruslongilateralis Kramer & Swartz, 2010, as well as to several undescribed forms from subequatorial Africa collectively referred to in the literature as the "Hippopotamyrusansorgii species complex" and colloquially known as "slender stonebashers." Previous molecular phylogenetic work has shown that these species are not close relatives of Hippopotamyruscastor Pappenheim, 1906, the type species of genus Hippopotamyrus Pappenheim, 1906 from Cameroon, and are thus misclassified. Hippopotamyrusansorgii species complex taxa and another species shown to have been misclassified, Paramormyropstavernei (Poll, 1972), are placed in genus Heteromormyrus and one genetic lineage from the Kwanza and Lucala rivers of Angola are identified as conspecific Heteromormyruspauciradiatus. Three additional new combinations and a synonymy in Mormyridae are introduced. The morphological characteristics and geographical distribution of the genus Heteromormyrus are reviewed. The electric organ discharges (EODs) of Heteromormyrus species are to be treated in a separate study.

Allopatric differentiation in the Enteromius anoplus complex in South Africa, with the revalidation of Enteromius cernuus and Enteromius oraniensis, and description of a new species, Enteromius mandelai (Teleostei: Cyprinidae).

The chubbyhead barb, Enteromius anoplus, as currently described, is the most widely distributed freshwater fish in South Africa. The species occurs in almost all the major river systems across the country, with the exception of the small coastal drainages on the south coast. The use of a comprehensive data set of mitochondrial (mtDNA) cytochrome b (cyt b) sequences uncovered the presence of four distinct lineages or operational taxonomic units (OTUs) within E. anoplus: (a) the Gouritz lineage endemic to the Gouritz River system, (b) the Olifants lineage endemic to the Olifants-Doring River system, (c) the Orange lineage endemic to the Orange River system and (d) the Eastern Cape lineage which is distributed across six river systems from the Great Fish to the Buffalo rivers in the Eastern Cape Province of South Africa. The present study provides a new diagnosis for E. anoplus s.s. (the Gouritz lineage), revalidates Enteromius cernuus (the Olifants lineage) and Enteromius oraniensis (the Orange lineage) as distinct species. The study also provides a description for a new species, Enteromius mandelai sp. nov (the Eastern Cape lineage). E. cernuus and E. mandelai differ from both E. anoplus and E. oraniensis by having a complete lateral line (vs. an incomplete lateral line in the latter species). E. cernuus further differs from the other three species by having long maxillary barbels which reach or exceed the vertical through the middle of the eye and the lowest number of circumpendicular scales (10-12 vs. 12-16 for the other three species). E. oraniensis is distinctive from the other three species by having inconspicuous barbels. These findings add to the growing body of literature that shows that the freshwater fishes of the Cape Fold and adjacent freshwater ecoregions have narrow geographic ranges. This has ramifications for conservation planning and management, as well as the understanding of the evolutionary history of the stream fishes in these global endemic hotspots.

Deep genetic and morphological divergence in the Hippopotamyrus ansorgii species complex (Teleostei: Mormyridae) in southern Africa.

The present study used molecular and morphological approaches to investigate hidden diversity within the Hippopotamyrus ansorgii species complex in southern Africa. Phylogenetic reconstructions and three species delimitation methods based on two mitochondrial markers (cytochrome b and cytochrome oxidase I) and one nuclear marker (S7) revealed 12 Molecular Operational Taxonomic Units (MOTUs), with two of them representing two recently described species, Hippopotamyrus longilateralis and Hippopotamyrus szaboi. The highest diversity occurred in the Kwanza River system, which contained five MOTUs, and the Upper Zambezi River system that had two MOTUs. Five other river systems contained a single MOTU each. A major impediment to the review of this complex has been the uncertainty surrounding the type locality of the specimens that were used for the description of H. ansorgii. The present study has, through a careful examination of published literature and synthesis of information on the travel activities of Dr. William Ansorge who collected the specimens, identified the Kwanza River system as the most plausible source of the syntypes. The resolution of the type locality of H. ansorgii facilitates future work on the review of this complex which is critical for providing reliable biodiversity estimates, identifying effective conservation management strategies and understanding the evolutionary history and biogeographic patterns of the fishes of this region.

An integrative taxonomic review of the Natal mountain catfish, Amphilius natalensis Boulenger 1917 (Siluriformes, Amphiliidae), with description of four new species.

An integrative taxonomic analysis combining mitochondrial cytochrome oxidase subunit I sequences, morphology, colour pattern and two species delimitation approaches revealed the existence of five lineages within the Natal mountain catfish, Amphilius natalensis, in southern Africa. These lineages are separated by substantial genetic divergences (1.6%-9.46%), and they can be consistently distinguished from one another based on a combination of morphology and colour pattern differences. Additionally, the lineages are allopatrically distributed and confined to isolated river systems draining discrete mountain ranges, which makes gene flow among them unlikely. One of these lineages is A. natalensis s.s., which is confined to the uMngeni and Tukela river systems in KwaZulu Natal (KZN) Province in South Africa. The other four lineages represent new species to science which are described as Amphilius zuluorum sp. nov., endemic to the uMkhomazi River system in KZN, Amphilius engelbrechti sp. nov., endemic to the Inkomati River system in Mpumalanga Province in South Africa, Amphilius marshalli sp. nov., endemic to the Pungwe and Lower Zambezi river systems in Zimbabwe and Mozambique, and Amphilius leopardus sp. nov., endemic to the Ruo River in Malawi. The results show that Amphilius laticaudatus which is endemic to the Buzi River system in Zimbabwe and Mozambique, belongs to the A. natalensis s.l. complex. A redescription of A. laticaudatus is presented and an updated identification key for the mountain catfishes of southern Africa is provided.

Distribution patterns and habitat associations of Sandelia bainsii (Teleostei: Anabantidae), a highly threatened narrow-range endemic freshwater fish.

Sandelia bainsii is a range-restricted and highly threatened freshwater fish endemic to South Africa. Recent genetic evidence suggests that this species comprises three allopatrically distributed lineages that have been informally designated as Sandelia sp. "Kowie," Sandelia sp. "Keiskamma" and Sandelia sp. "Buffalo." As these lineages have only been recently identified and are likely to face a high risk of extinction because of restricted distributions, there is a critical need for generating ecological information to guide conservation prioritisation. The present study compared the historical and current distribution patterns, together with the habitat associations of Sandelia sp. "Kowie" in the Koonap and Kat rivers, tributaries of the Great Fish River. This study indicated that this lineage has been extirpated from one of the three localities in the Koonap River where it was historically abundant. In the Kat River, the current distribution of Sandelia sp. "Kowie" was comparable to its historical range, but its future persistence is threatened by the presence of non-native piscivores, instream physical barriers and potential future exploration for shale gas and infrastructure development in the Karoo Basin. A generalised hurdle negative binomial model revealed that although this lineage's probability of occurrence was high in habitats with boulder and sand substrates, and low conductivity, habitat characteristics were poor predictors of its abundance. Thus, it was postulated that the current range of this lineage probably represents the only available habitats for the persistence of different life stages for this taxon. Alternatively, the observed patterns may suggest the possibility of a shift in habitat associations, possibly for optimum utilisation of the remaining refugia within this river system. Immediate conservation measures should focus on preventing the spread on non-native invasive fishes, whereas future studies should evaluate the impacts of population fragmentation and identify appropriate intervention measures to maintain this lineage's long-term adaptive potential.

Integrative taxonomy reveals hidden diversity in the southern African darters genus Nannocharax Günther 1867 (Characiformes: Distichodontidae).

The present study explored the diversity of Nannocharax within southern Africa by implementing three species delimitation methods for a data set consisting of 37 mitochondrial cytochrome oxidase subunit I sequences. Two unilocus coalescent methods, the General Mixed Yule Coalescent (GMYC) and the Bayesian implementation of the Poisson Tree Processes (bPTP), and a genetic distance method, the Automatic Barcode Gap Discovery (ABGD), were applied. Both GMYC and bPTP delimited the same operational taxonomic units (OTUs), revealing a higher diversity for the genus in the region than previously recognised, whereas the ABGD failed to delimit the same candidate species. All methods delimited two species groups, and these are supported based on colouration patterning and morphology; the Nannocharax multifasciatus and the Nannocharax macropterus species groups and the delimited OTUs were assigned to each. Two putative new species were identified, Nannocharax cf. lineostriatus "Okavango" from the Okavango River in Angola and N. cf. lineostriatus "Kwanza" from the Kwanza River system in Angola. The distribution of Nannocharax dageti was confirmed for the Upper Zambezi and extended to the Okavango system, and an identification key for the southern Africa Nannocharax species is provided.

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.

Designation of a neotype for Enteromiuspallidus (Smith, 1841), an endemic cyprinid minnow from the Cape Fold Ecoregion, South Africa.

Enteromiuspallidus was described by Smith in 1841 without a designated type specimen for the species. Herein, we designate a specimen from the Baakens River system as a neotype for E.pallidus and provide a thorough description for this species to facilitate ongoing taxonomic revisions of southern African Enteromius. Enteromiuspallidus can be distinguished from the other minnows in the "goldie barb group" by having an incomplete lateral line, lack of distinct chevron or tubular markings around lateral line pores, absence of a distinct lateral stripe, absence of wavy parallel lines along scale rows and lack of black pigmentation around the borders of the scales. We provide mtDNA COI sequences for the neotype and an additional specimen from the Baakens River as DNA barcodes of types and topotypes are a fundamental requirement for further taxonomic studies.

The influence of diet composition and tissue type on the stable isotope incorporation patterns of a small-bodied southern African minnow Enteromius anoplus (Cypriniformes, Cyprinidae).

RATIONALE: In trophic ecology, the use of stable isotope data relies on the general understanding of isotope turnover rates and diet-to-tissue discrimination factors (DTDFs). Recent studies on the application of stable isotope data have shown that isotope turnover rates and DTDFs can be influenced by many factors, including diet composition and tissue type. This study investigated the influence of diet composition and tissue type on stable isotope incorporation patterns in a small-bodied African minnow, the chubbyhead barb Enteromius anoplus. METHODS: The isotopic incorporation patterns of carbon (δ13 C values) and nitrogen (δ15 N values) into white muscle and caudal fin tissues of the chubbyhead barb were examined using two isotopically different diets. Controlled-diet stable isotope feeding trials using a fishmeal-based diet (diet 1) and a soya-based diet (diet 2) were conducted over a 180-day period for the chubbyhead barb. RESULTS: The two diets had contrasting isotopic incorporation patterns: diet 1 was associated with progressively high δ13 C and δ15 N values, whereas diet 2 was associated with progressively low δ13 C and δ15 N values over time for both muscle and fin tissues. The δ13 C turnover rates were similar for both tissues (56 and 61 days), whereas the δ15 N turnover rates differed between fin and muscle tissue in both diets (diet 1 = 4 and 130 days, and diet 2 = 72 and 300 days, respectively). The DTDFs were similar for both tissues in diet 1 (Δ13 C: -3.96 to -2.62‰, Δ15 N: 1.98 to 2.61‰) and diet 2 (Δ13 C: 4.05 to 5.24‰, Δ15 N: 8.45 to 9.69‰). CONCLUSIONS: These results suggest that fin tissue can potentially be used as an alternative for muscle tissue in food web studies with a reasonable level of error. The isotopic turnover rate and DTDFs estimates for E. anoplus, however, require consideration of diet composition because different diets may differ in their isotopic incorporation patterns.

Evidence of hidden diversity and taxonomic conflicts in five stream fishes from the Eastern Zimbabwe Highlands freshwater ecoregion.

Stream fishes of the Eastern Afromontane region are among the least studied vertebrates in this region, despite the potential for harbouring cryptic diversity. The present study examined mitochondrial cytochrome oxidase subunit I (COI) sequence divergence in 153 specimens of stream fishes belonging to four genera and three families, [(Amphilius and Zaireichthys (Amphiliidae); Chiloglanis (Mochokidae); and Hippopotamyrus (Mormyridae)], in the Eastern Zimbabwe Highlands (EZH) freshwater ecoregion to explore the extent to which the current taxonomy conceals the ichthyofaunal diversity in the region. The General Mixed Yule Coalescent (GMYC) species delineation method identified 14 clusters within five currently recognised 'species' from the EZH ecoregion. Only one of these clusters represents a named species, while 13 of them represent candidate or undescribed species. Our results revealed that effective conservation of this region's unique biota is limited by the incomplete knowledge of taxonomic diversity and inaccurate mapping of species distribution ranges.

A review of the Pseudobarbus afer (Peters, 1864) species complex (Teleostei, Cyprinidae) in the eastern Cape Fold Ecoregion of South Africa.

The Eastern Cape redfin, Pseudobarbus afer, has long been considered to be a single widespread and variable species occurring in multiple isolated river systems in the Cape Fold Ecoregion (CFE) at the southern tip of Africa. Mitochondrial cytochrome b and control region sequence data of individuals from populations currently assigned to Pseudobarbus afer across the species' distribution range revealed existence of four deeply divergent taxonomic units: (i) the Mandela lineage confined to the Sundays, Swartkops and Baakens river systems, (ii) the Krom lineage endemic to the Krom River system, (iii) the St Francis lineage occurring in the Gamtoos and adjacent river systems, and (iv) the Forest lineage occurring in several coastal river systems from the Tsitsikamma to the Klein Brak River system. The Forest lineage is closely related to Pseudobarbus phlegethon from the Olifants River system on the west coast of South Africa, suggesting that it does not belong to Pseudobarbus afer s.l. Herein we focus on the three lineages within the Pseudobarbus afer s.l. complex and provide new diagnosis for Pseudobarbus afer s.s (Mandela lineage), revalidate Pseudobarbus senticeps (Krom lineage) as a distinct species, and describe a new species Pseudobarbus swartzi (St Francis lineage). The three species exhibit subtle differences, which explains why they were previously considered to represent a single variable and widespread species. Pseudobarbus senticeps differs from both Pseudobarbus afer and Pseudobarbus swartzi by having fewer (i.e. larger) scales (25-33, mode 29 lateral line scale series; 10-12, mode 11 circumpeduncular scales) and presence of a lateral stripe which terminates in a conspicuous triangular blotch at the base of the caudal fin. Long barbels which reach or surpass the vertical through the posterior edge of the eye further separate Pseudobarbus senticeps from Pseudobarbus afer s.s. which possesses simple short barbels which do not reach the vertical through the posterior margin of the eye. Pseudobarbus afer s.s differs from Pseudobarbus swartzisp. n. by possession of fewer scale rows along the lateral line (29-35, mode 32 vs 34-37, mode 36 in Pseudobarbus swartzi), fewer scales around the caudal peduncle (12-16, mode 12 vs 13-17, mode 16 in Pseudobarbus swartzi) and a distinct mesh or net-like pigmentation pattern on latero-ventral scales.