Emmanuel J. W. M. N. Vreven’s research while affiliated with Royal Museum for Central Africa and other places

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Publications (8)


Map of the Upemba National Park and its surroundings. Sites sampled prior to this study are indicated by black circles and those of this study (2012–2020) by red circles. (1) core zone, (2) annex zone, (3) and (4) buffer zones: Bena Mulumbu Hunting Area (3) and Lubudi-Sampwe Hunting Area (4). Double bars indicate major falls: (a) Kamwanga Falls, (b) Kayo Falls, (c) Luvilombo Falls, (d) Kyubo Falls, and (e) Kalumengongo Falls.
Longitudinal profile, subdivisions, and fish species numbers of the Kalule Nord Basin. Red dots indicate the sampling sites. Waterfalls mark the separation of the five different sections numbered 1 to 5. The blue frame represents the highland zone and the orange frame the lowland zone.
Illustrations of bad practices related to fisheries and agriculture, the latter also impacting river ecosystem health. (a) Channel linking the Upper Lualaba and Lake Mulenda in the Kamalondo Depression (3 September 2015). (b) Cutting grass to facilitate seining, Lake Lukanga (10 September 2015). (c) Fishing dam on the Fungwe River, Kebange Village (21 August 2015). (d) Transport of ichthyotoxic fruits (Amblygonocarpus andongensis, Fabaceae) for fishing in the Mwanza River, Kabweyi Village (20 August 2016). (e) Transport of a seine made of mosquito nets in Lake Upemba, Misebo Village (13 October 2017). (f) A dry stretch of the Luvilombo River due to upstream water diversion for irrigating bean fields (25 July 2019).
The Upemba National Park (Upper Congo Basin, DR Congo): An Updated Checklist Confirming Its Status as an African Fish Biodiversity Hotspot
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August 2023

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549 Reads

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5 Citations

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Jos Snoeks

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Emmanuel J. W. M. N. Vreven

An annotated checklist of the ichthyofauna of the Upemba National Park, draining part of the Upper Lualaba basin and situated in the southern part of the Democratic Republic of the Congo, is presented, based on a literature review, a re-examination of museum collections, and a study of recent collections (2012–2020). In total, 247 native and 1 introduced species, Heterotis niloticus, are reported. The native species belong to 78 genera, 26 families, and 15 orders. Of these, 45 species (18%) are endemic to the park, 35 species (14%) await formal description, and 5 taxa (2%) need further study to clarify their status. With 51 species, the Cyprinidae is by far the most species-rich family, followed by the Mormyridae (26), Mochokidae (26), Alestidae (18), Distichodontidae (18), Amphiliidae (17), and Cichlidae (16). The remaining families are represented by less than 15 species. Comments about the species distribution and the fish fauna shared with adjacent ecoregions are provided. Although the park provides some protection for the fish species living within its borders by limiting human access to the core zone, the annex and buffer zones are both subject to strong anthropogenic pressure. These observations underscore the need for the implementation and further elaboration of fish-related preservation guidelines and plans to enable better protection/conservation of the park’s ichthyofauna.

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Revision of Nannocharax luapulae Boulenger, 1915 (Characiformes: Distichodontidae) from the Upper Congo basin: Evidence for a species pair

June 2023

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500 Reads

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4 Citations

For many decades, Nannocharax luapulae has been considered to be widespread in the southern part of the Upper Congo basin. However, meristic, morphometric and cytochrome c oxidase subunit I (COI) barcoding evidence revealed that its geographical distribution is restricted to the Luapula‐Moero basin. The populations of the Upper Lualaba are assigned to a new species, N. chochamandai. This new species, though highly similar to N. luapulae, can readily be distinguished from it by its lower number of lateral line scales, 41–46 (vs. 49–55), its pectoral fin reaching the pelvic‐fin insertion (vs. not reaching the pelvic‐fin insertion) and its pelvic fin reaching the base of the anal fin (vs. not reaching the base of the anal fin). Specimens of N. chochamandai display thickened pads on the first three pelvic‐fin rays that exhibit intraspecific variation in development, which appears to be related to the flow‐strength of the river in which these Nannocharax specimens occur. Nannocharax luapulae is redescribed and an updated identification key to the Nannocharax species of the Congo basin sensu lato is provided as well. Some fish conservation issues related to N. luapulae and N. chochamandai are also highlighted.


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

April 2023

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215 Reads

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4 Citations

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.


Map of the Kundelungu National Park (KNP, Haut Katanga, Democratic Republic of the Congo) and its surroundings: Falls are indicated by = and letters: (a) Mwadingusha, (b) Koni, (c) Kyubo, (d) Mwena, (e) Lofoi, (f) Kasanga, (g) Luiji, (h) Lutshipuka, (i) Masansa, (j) Luansa and (k) Mambilima. The three lakes sampled are (1) Bwaya, (2) Pungwe and (3) Lubanda. KP= Kundelungu Plateau (~1200 to 1700 m); KbP = Kibara Plateau (~1220 to 1850 m); BP = Biano Plateau (~1230 to 1630 m); CZ = Core Zone; AZ = Annex Zone and dashed line = Buffer Zone as defined in this study. See inserted legend for other details.
The native fish diversity per family and its distribution in the three conservation zones of the KNP. CZ, Core, AZ, Annex, and BZ, Buffer Zones.
Venn diagrams showing the number of shared and non-shared species in (a) the three conservation zones of the KNP; (b) the three parts of the Lufira Basin; and (c) the upper Lualaba Ecoregion which include the middle Lufira Basin and its two nearby ecoregions. E, endemic species; and NE, not evaluated.
The conservation status of the KNP fishes (based on the actual fish diversity) according to the list of threatened species of the International Union for Nature Conservation (IUCN, 2019). NE: Not Evaluated, DD: Data Deficient, LC: Least Concern, VU: Vulnerable, and CR: Critically endangered.
Checklist of the Fishes of the Kundelungu National Park (Upper Congo Basin, DR Congo): Species Diversity and Endemicity of a Poorly Known Ichthyofauna

February 2023

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676 Reads

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5 Citations

The fish diversity of the Kundelungu National Park (KNP), one of the seven national parks of the Democratic Republic of the Congo, has never been thoroughly studied. This first checklist is presented based on a literature compilation and the study of historical (1939–1969) and recent collections (2012–2017). A total of 96 taxa are reported, including 64 native described species, one introduced species (Poecilia reticulata), 13 new species that await formal description and 18 possibly new species that require further investigation to verify their status. These taxa represent 39 genera and 17 families from the KNP including its Buffer Zone (BZ). Only six taxa, including five endemics, are known from the Core Zone on the Kundelungu Plateau (1300–1700 m alt.). At lower altitudes (800–1100 m), in the Annex Zone, 71 taxa, including 17 endemics, were found. Finally, 50 taxa, including 13 endemics and one introduced species, are known from its BZ. The fish fauna of the KNP is threatened by overfishing, destructive fishing practices, and habitat degradation due to mining pollution, and deforestation for agriculture on the river banks. The present study provides the much needed baseline data for the protection and conservation planning of this fish fauna, for which conservation suggestions are formulated.


A new Parakneria Poll 1965 (Gonorhynchiformes: Kneriidae), ‘Mikinkidi’ from the Upper Lufira Basin (Upper Congo: DRC): Evidence from a morphologic and DNA barcoding integrative approach

November 2022

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235 Reads

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2 Citations

A new species, Parakneria alytogrammus, is described from the main stream of the Upper Lufira River. This species is easily distinguished from its congeners from the Congo Basin by its unique colouration, consisting of a low number of transversal bands on each of the caudal‐fin lobes, 2 (vs. 3–5) and the presence of an uninterrupted lateral mid‐longitudinal black band in fresh and preserved specimens (vs. absent). In addition, the new species differs from its Upper Lualaba congeners by the narrow width of its pectoral‐fin base, 4.8–5.6% LS [vs. wider, 8.2–10.1% for P. lufirae, 8.6% LS for P. damasi (holotype), and 7.6–7.9% LS for P. thysi]. Finally, it differs from the only species currently known from the Luapula‐Mweru system, P. malaissei, by having a short post‐dorsal distance, 36.4–36.6% LS (vs. longer, 38.6–41.1% LS) and a short post‐pelvic distance of 40.0–40.6% LS (vs. longer, 41.4–44.1% LS). Mitochondrial DNA‐haplotypes of P. alytogrammus sp. nov. form a clade, which is sister to the P. thysi clade, and from which it diverges by a genetic (Kimura 2‐parameter and uncorrected p) distance of 0.7% in the COI‐barcoding locus. The Upper Lufira, one of the sub‐basins of the Upper Congo Basin, remains poorly explored relative to its fish fauna. In contrast, the region is well explored with regard to its mineral wealth. Unfortunately, mining exploitation is carried out in the region without proper concern for the environment. Thus, the discovery of this new species for science calls for increased protection and aquatic biodiversity exploration in this mining region.


Disentangling the Diversity of the Labeobarbus Taxa (Cypriniformes: Cyprinidae) from the Epulu Basin (DR Congo, Africa)

November 2022

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118 Reads

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5 Citations

In an attempt to disentangle the complex taxonomy of the Labeobarbus species of the Epulu River, a right bank headwater affluent of the Aruwimi, Central Congo basin, a morphological study was undertaken on 221 specimens from the Epulu and 32 type specimens. As a result, five different species have been distinguished, including four so-called rubberlips, L. caudovittatus, L. macroceps, L. mawambiensis, and L. sp. ‘thick lip’, and one chiselmouth, L. longidorsalis. While rubberlips have a curved mouth with well-developed lips and often a mental lobe, chiselmouths have a straight mouth with a keratinised cutting edge on the lower jaw. Among the specimens examined, several presented an intermediate mouth morphology between L. mawambiensis and L. longidorsalis, either with one or two pairs of barbels. One specimen exhibited an intermediate morphology between L. mawambiensis and L. macroceps. This morphological study, complemented with a molecular study of the mitochondrial gene cytochrome b (cyt b), suggests that these intermediates are probably hybrid specimens. The Epulu case is reminiscent to a case of possible hybridisation recently discovered in the Inkisi River (Lower Congo basin), but differs in having a lower relative abundance of hybrid specimens in the population, and in phylogenetic patterns.


ML phylogeny (RAxML) of the order Siluriformes, including 30 catfish families (indicated either in blue or in red = target families), based on all mitochondrial protein coding genes (11,400 bp). Bootstrap proportion based on 1000 BS replicates are indicated at nodes either by numbers or by black dots (BS = 100). Genera including more than one taxon are collapsed (see Supporting Information Figure S1 for the noncollapsed tree). Representative species of most included catfish families/subfamilies are depicted from top to bottom (photographers in brackets, specimen/species not necessarily included in the dataset): Mochokidae; Chiloglanidinae: Chiloglanis sp. Ntumbachushi (F.D.B. Schedel), Mochokidae; Chiloglanidinae: Synodontis petricola (F.D.B. Schedel), Claroteidae: Chrysichthys (F.D.B. Schedel), Auchenoglanididae: Parauchenoglanis cf. monkei (F.D.B. Schedel), Schilbeidae: Schilbe grenfelli (F.D.B. Schedel), Malapteruridae: Malapterurus sp. (F.D.B. Schedel), Amphiliidae: Amphilius sp. (F.D.B. Schedel). Ictaluridae: Ameiurus nebulosus (F.D.B. Schedel), Pangasiidae: Pangasius pangasius (E. Schraml), Austroglanididae: Austroglanis barnardi (Roger Bills), Ariidae: Ariopsis cf. guatemalensis (preserved specimen; F.D.B. Schedel), Siluridae: Silurus glanis (E. Schraml), Pimelodidae: Sorubim lima (E. Schraml), Pseudopimelodidae: Lophiosilurus alexanderi (E. Schraml), Sisoridae: Glyptothorax cf. sinensis (E. Schraml), Sisoridae: Hara jerdoni (E. Schraml), Amblycipitidae: Liobagrus reinii (E. Schraml), Bagridae: Tachysurus fulvidraco (F. Schäfer), African Bagridae: Bagrus sp. (F.D.B. Schedel), Horabagridae: Horabagrus brachysoma (J. Geck), Clariidae: Clarias gariepinus (F.D.B. Schedel), Heteropneustidae: Heteropneustes fossilis (preserved specimen, F.D.B. Schedel), Auchenipteridae: Tetranematichthys wallacei (E. Schraml), Doradidae: Platydoras armatulus (E. Schraml), Aspredinidae: Bunocephalus coracoideus (E. Schraml), Plotosidae: Plotusus lineatus (E. Schraml), Chacidae: Chaca bankanensis (preserved specimen, F.D.B. Schedel), Cetopsidae: Cetopsis coecutiens (preserved specimen, F.D.B. Schedel), Diplomystidae: Diplomystes sp. (preserved specimen, F.D.B. Schedel) Loricariidae: Hypostominae: Hypostomus cf. plecostomus (E. Schraml) Loricariidae: Hypoptopomatinae: Otocinclus sp. (J. Geck), Callichthyidae: Corydoras rabauti (E. Schraml)
Detailed relationships among the ‘Big African clade’ (pruned from the ML phylogeny depicted in Figure 1). Bootstrap proportion based on 1000 BS replicates are indicated at nodes either by numbers or by black dots (BS = 100). Represented mochokid species depicted from top to bottom (photographers and corresponding sample ID in brackets): Chiloglanis macropterus (F.D.B. Schedel, DRC‐2012/3637), Chiloglanis sp. ‘Lufupa’ (F.D.B. Schedel, DRC‐2016/4684), Chiloglanis sp. ‘Ntumbachushi’ (F.D.B. Schedel, DRC‐2012/3568), Euchilichthys sp. Mukuleshi (F.D.B. Schedel, DRC‐2012‐4612), Atopochilus savorgnani (B. Sidlauskas, GAB17‐365), Atopodontus adriaensi (B. Sidlauskas, GAB17‐364), Synodontis petricola (F.D.B. Schedel, P‐AA‐1422), Mochokiella paynei (F.D.B. Schedel, P‐AA‐1344), Microsynodontis cf. batasii (F.D.B. Schedel, P‐AA‐1504), Mochokus brevis (F.D.B. Schedel, SUD‐2016‐847), Acanthocleithron chapini (E.J.W.M.N. Vreven, P‐AA‐1382)
Detailed relationships of the family Austroglanididae and related families (pruned from the ML phylogeny depicted in Figure 1). Bootstrap proportion based on 1000 BS replicates are indicated at nodes either by numbers or by black dots (BS = 100)
ML phylogeny (RAxML) of the order Siluriformes (focal families are indicated by pictures of representative species) based on the ‘nuclear dataset’, including the sequence data of rag1 and rag2 provided by the study of Sullivan et al. (2006) and the newly created sequences of four Austroglanis specimens (117 individuals, 1983 bp). Bootstrap proportions based on 1000 BS replicates are indicated at nodes either by numbers or by black dots (BS = 100)
New phylogenetic insights into the African catfish families Mochokidae and Austroglanididae

March 2022

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496 Reads

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16 Citations

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.


New phylogenetic insights into the African catfish families Mochokidae and Austroglanididae

March 2022

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50 Reads

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1 Citation

Several hundred catfish species (order: Siluriformes) belonging to eleven families inhabit Africa, of which at least six families are endemic to the continent. Though 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.

Citations (7)


... The genus is composed of a particularly diverse set of species with divergent morphological traits in mouth parts, both at the intra-specific and inter-specific levels (Nagelkerke et al. 1994;de Graaf et al. 2010;Vreven et al. 2016;Levin et al. 2019;Kisekelwa et al. 2020). In some cases, mouthpart divergence does not necessarily imply further body transformations (e.g., Kisekelwa et al. 2020;Katemo et al. 2023). Multiple Labeobarbus species can be found in syntopy, sometimes representing highly morphologically diverse assemblages of species, such as in Lake Tana in Ethiopia (Nagelkerke et al. 1994;Nagelkerke and Sibbing 1998;de Graaf et al. 2008). ...

Reference:

Insight Into Trophic Niche Differentiation in Labeobarbus (Cyprinidae) in the Luhoho Basin (Upper Congo Basin)
The Upemba National Park (Upper Congo Basin, DR Congo): An Updated Checklist Confirming Its Status as an African Fish Biodiversity Hotspot

... present in Claroteidae), the anterior nostrils placed on the upper lip (vs. on the snout in Claroteidae), and a rounded caudal fin (vs. deeply forked in Claroteidae) (Mo 1991, Geerinckx et al. 2007, 2013, Sithole et al. 2023. Within the family Auchenoglanididae, Parauchenoglanis Boulenger 1911 is the most diverse genus, comprising 11 valid species (Sithole et al. 2023, Fricke et al. 2024, Modimo et al. 2024. ...

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

... A new Nannocharax species for science has been collected in the Lovoi River [50]. This new species, here named N. sp. ...

Revision of Nannocharax luapulae Boulenger, 1915 (Characiformes: Distichodontidae) from the Upper Congo basin: Evidence for a species pair

... Considering the whole Congo Basin, a total of 13 clariid species are reported [15]; among them, seven species are present in the Upper Congo Basin (C. gariepinus; C. ngamensis; C. buthupogon Sauvage, 1879; C. dumerilii Steindachner, 1866; C. liocephalus Boulenger, 1898; C. stappersii Boulenger, 1915; C. theodorae Weber, 1897) [16,17]. The latter five fish species have never been studied for their monopisthocotylan fauna [18]. ...

Checklist of the Fishes of the Kundelungu National Park (Upper Congo Basin, DR Congo): Species Diversity and Endemicity of a Poorly Known Ichthyofauna

... In contrast to the statement of Giltay (1934) that in Kneria auriculata, the ventral fin is directly below the origin of the dorsal fin, and the statement of Grande and Arratia (2010:56) that in gonorynchiforms, 'the dorsal fin is always anterior to the pelvic fins', this character is in general valid. As Mutambala et al. (2022) recently reviewed this issue, only single specimens may deviate from the genus specific position of the pelvic fin. Position, shape, and number of elements are similar within Gonorynchiformes (Table 3); nevertheless, all taxa show some peculiarities. ...

A new Parakneria Poll 1965 (Gonorhynchiformes: Kneriidae), ‘Mikinkidi’ from the Upper Lufira Basin (Upper Congo: DRC): Evidence from a morphologic and DNA barcoding integrative approach

... Two specimens have thick lips (rubberlips with a developed mental lobe) and have longer intestines, whereas the two others displayed a regular mental lobe and had shorter intestines (400-900 mm vs. 250-300 mm). The morphotype 'thick lip' is probably a new species(Kisekelwa 2019;Decru et al. 2022), confused with the morphotype with a regular mental lobe due to sharing a flexible last unbranched dorsal-fin ray. ...

Disentangling the Diversity of the Labeobarbus Taxa (Cypriniformes: Cyprinidae) from the Epulu Basin (DR Congo, Africa)

... In the Hardhead Sea Catfish, Ariopsis felis (Ariidae), an elongate rosette, similar that of auchenoglanidids, has an array of 40 lamellae, but flow is externally induced primarily by a Pitot mechanism in which water enters through a funnel-like anterior nostril perpendicular to the flow of water traveling over the fish's head, in addition to beating of cilia of the sensory epithelium (Zeiske et al., 1994;Cox, 2008). While only distantly related to auchenoglanidids (Lundberg et al., 2007;Schedel et al., 2022), the Striped Eel Catfish, Plotosus lineatus (Plotosidae), has an olfactory system incorporating an accessory sac. Entrance to the sac is located just distal to a compact olfactory rosette which bears just 10-12 lamellae. ...

New phylogenetic insights into the African catfish families Mochokidae and Austroglanididae