#ZooKeys ZooKeys 1241: 261-290 (2025) DOI: 10.3897/zookeys.1241.138917 Research Article Description of two new species of Chiloglanis (Teleostei, Mochokidae) from the Eastern Zimbabwe Highlands freshwater ecoregion: an overlooked hotspot of rheophilic fishes Tadiwa |. Mutizwa'”®, Taurai Bere®®, Wilbert T. Kadye’”®, Pedro H. N. Braganca’®, Albert Chakona’?"© Pe wo ND NRF-South African Institute for Aquatic Biodiversity (NRF-SAIAB), P Bag 1015, Makhanda (Grahamstown) 6140, South Africa Department of Ichthyology and Fisheries Science, Rhodes University, PO Box 94, Makhanda (Grahamstown), 6140, South Africa School of Wildlife, Ecology and Conservation, Chinhoyi University of Technology, Private Bag 7724, Chinhoyi, Zimbabwe Department of Ichthyology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA Corresponding author: Tadiwa I. Mutizwa (T. Mutizwa@saiab.nrf.ac.za) OPEN Qrceess Academic editor: Yahui Zhao Received: 9 October 2024 Accepted: 12 March 2025 Published: 16 June 2025 ZooBank: https://zoobank. org/8F4F2E53-1A28-4778-BABD- 782F1421C165 Citation: Mutizwa Tl, Bere T, Kadye WT, Braganca PHN, Chakona A (2025) Description of two new species of Chiloglanis (Teleostei, Mochokidae) from the Eastern Zimbabwe Highlands freshwater ecoregion: an overlooked hotspot of rheophilic fishes. Zookeys 1241: 261-290. https://doi. org/10.3897/zookeys.1241.138917 Copyright: © Tadiwa |. Mutizwa et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Growing evidence indicates that species diversity within the genus Chiloglanis Peters 1868 is poorly resolved and major taxonomic revisions are required. By integrating genetic and morphological analyses, this study describes two new Chiloglanis spe- cies from the Eastern Zimbabwe Highlands (EZH) freshwater ecoregion, a region that, until only recently, had been poorly-explored in terms of its ichthyological diversity. Chiloglanis asperocutis Mutizwa, Braganca & Chakona, sp. nov. is distinguished from other southern African congeners by a combination of characters, including ridge- like tubercles distributed on the dorsal and lateral surfaces of the head and body giving the skin a conspicuously rough texture, ten closely packed mandibular teeth, deeply forked caudal fin and a high number of primary premaxillary teeth (68-128). Chiloglanis compactus Mutizwa, Bragancga & Chakona, sp. nov., attains the smallest size (> 46 mm SL) of all currently known congeners in southern Africa. It is distin- guished from all the other congeners from the region by a combination of characters; the possession of seven pectoral fin rays, conical tubercles distributed across the dorsal and lateral surface of the head and body, eight widely spaced mandibular teeth, a shallow forked caudal fin with rounded lobes, a low number of primary premaxillary teeth (31-53) and fewer dorsal fin rays (5). These two species are distributed in both the Buzi and Pungwe River systems. The study is the first in a series of publications that will provide formal descriptions of a number of deeply divergent lineages (candi- date species) identified in previous studies from southern Africa. The persistence of the unique riverine fauna in the EZH is threatened by multiple impacts that are altering the hydrological regime of the rivers and streams as well as habitat degradation and excessive sedimentation from gold panning and agricultural activities. Key words: Afromontane streams, diversity, ecological impacts, integrative taxonomy, southern Africa, suckermouth catfishes * These authors contributed equally to this work. 261 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Introduction Previous and ongoing studies indicate that many of the currently recognised freshwater fish species in southern African harbour hidden diversity (e.g., Swartz et al. 2009; Chakona and Swartz 2013; Chakona et al. 2015, 2018; Kam- bikambi et al. 2021; Mazungula and Chakona 2021; Sithole et al. 2023; Mutizwa et al. 2024; Scheepers et al. 2024). Recently, Chakona et al. (2018) provided the first evidence of undocumented diversity and taxonomic conflicts in fish- es from high attitude streams in the Eastern Zimbabwe Highlands freshwater ecoregion (EZH), one of the poorly explored regions in southern Africa. Similar findings were reported from a number of studies on species from other high altitude streams in east and west Africa (e.g., Friel and Vigliotta 2011; Schmidt et al. 2015, 2016, 2017, 2023; Morris et al. 2016; Schmidt and Barrientos 2019; Schedel et al. 2024). This indicates that, despite being historically assumed to have a depauperate fish fauna, these high altitude freshwater ecosystems rep- resent previously overlooked hotspots of diversity and endemism (e.g., Schmidt et al. 2017). Growing evidence shows that many groups of fishes inhabiting high altitude streams in Africa are in need of major taxonomic reassessments (e.g., Friel and Vigliotta 2011; Schmidt et al. 2015, 2016, 2017, 2023; Morris et al. 2016; Schmidt and Barrientos 2019). The aim of the present study was to investigate the taxonomic status of suckermouth catfishes of the genus Chiloglanis Peters, 1868 from the EZH. The first published detailed checklist of freshwater fishes of Zimbabwe by Jubb (1961) contained one species of suckermouth catfish, Chiloglanis neumanni Boulenger, 1911, from the EZH. For reasons that remain unclear in the literature, subsequent publications such as Bell-Cross (1976) listed two species, Chilogla- nis emarginatus Jubb and Le Roux, 1969 and C. neumanni, whereas Skelton (2001) recognised three species, C. emarginatus, C. neumanni and Chilogla- nis pretoriae Van der Horst, 1931 from the same region. Marshall (2011), who summarised the existing knowledge of the fishes of Zimbabwe, concluded that the Limpopo River system represented the northern-most distribution limit for C. pretoriae, and questioned the taxonomic assignment of the EZH sucker- mouth catfishes to C. neumanni, a species that was described from the Bubu River, a tributary of the Great Ruaha River in Tanzania. Marshall (2011) further commented that C. neumanni and C. emarginatus are unlikely to occur in Zim- babwe and suggested that specimens that were previously assigned to these two species likely represented several undescribed species whose taxonomic identity required further investigation. Marshall’s (2011) tentative proposition was supported by subsequent results from DNA barcoding studies that showed considerable genetic di- vergence of suckermouth catfishes of the EZH from currently described species of Chiloglanis from southern Africa (Chakona Et Al. 2018; Mutizwa et al. 2024) and specimens from the Great Ruaha River system in Tanzania (Day Et Al. 2023). Chakona et al. (2018) identified at least three candidate species of Chiloglanis from the Pungwe and Buzi River systems in the EZH. Here, we apply an integrative taxonomic approach combining genetic and morphological analyses to provide evidence supporting the recognition of two of these lineages tentatively named Chiloglanis sp. “rough skin” and Chiloglanis sp. “dwarf” in Chakona et al. (2018), as distinct species. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 262 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Materials and methods Sample collection Voucher specimens and tissue samples were collected during surveys of the EZH conducted in 2013, 2014 and 2022 in the Buzi and Pungwe river systems (Fig. 1). Sampling was done using a Samus-725M electrofisher with a seine net placed downstream to capture immobilised fish in fast-flowing current. Species were iden- tified using regional identification guides by Skelton (2001) and Marshall (2011). Captured fishes were euthanized with clove oil, some specimens were digitally photographed, and a small piece of muscle tissue was dissected from the right side of each specimen and preserved in 95% ethanol. Tissue samples were stored at -80 °C at the National Research Foundation-South African Institute for Aquatic Biodiversity (NRF-SAIAB), Makhanda. Voucher specimens were fixed in 10% for- malin in the field. They were then transferred through 10% and 50% to 70% ethanol for long-term storage. All voucher specimens were deposited into the fish collec- tion facility at the NRF-SAIAB as reference material. In addition, the study included specimens collected from the Pungwe and Buzi Rivers prior to the 2013 that were lodged at the National Fish Collection at NRF-SAIAB (see Material examined). DNA extraction, amplification, and sequencing This study used sequences generated by Chakona et al. (2018) and Mutiz- wa et al. (2024) deposited in GenBank (accession numbers: MH432018- MH432062, PP156890-PP156895). An additional eight COl sequences were generated for this study using specimens collected from tributaries of the Pungwe River in Gorongosa National Park, Mozambique in surveys conducted between July and August 2022 (GenBank accession numbers: PQ424602-PQ424609). Preparation and sequencing of genetic material was done in the Aquatic Genomics Research Platform at the NRF-SAIAB. Axe. compactus specimens only Ac compactus genetic sample + specimens (IC. asperocutis specimens only EC. asperocutis genetic sample + specimens ras Chiloglanis sp. 'Pungwe' genetic sample + specime ===National borders 0 50 100 km A mae Continental boundary Towns Figure 1. Collection sites of Chiloglanis specimens from the Pungwe and Buzi river systems. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 263 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands DNA extraction, amplification, and sequencing followed Mutizwa et al. (2024). Genomic DNA was extracted from preserved tissues using the salt- ing-out method (Sunnucks and Hales 1996). The mitochondrial DNA cyto- chrome c oxidase subunit | (COI) gene was amplified by polymerase chain reaction (PCR) using the universal fish DNA barcoding primer set FishF1 and FishR1 (Ward et al. 2005). PCRs were performed with a Veriti 96 well thermal cycler (Applied Biosystems, USA) and each reaction mixture (25 uL) contained 50-100 ng) of template DNA, 6.5 uL of water, 0.5 uL of each primer (10 uM), and 12.5 uL Taq DNA polymerase 2 x master mix red (Am- plicon PCR enzymes and reagents, Denmark). The PCR amplification profile had an initial denaturation step of 3 min at 94 °C followed by 38 cycles of 30 sec at 94 °C, annealing at 55 °C for 30 sec, and extension at 72 °C for 50 sec, and final extension at 72 °C for 7 min. The amplicons were pu- rified using an Exonuclease I-Shrimp Alkaline Phosphate (Exo/SAP, Ther- mo Fisher Scientific, USA) protocol (Werle et al. 1994), sequenced using standard fluorescent BigDye v. 3.1 (Applied Biosystems, USA) terminator chemistry in the forward direction, and analysed on a 3500 Genetic Analyser (Applied Biosystems, USA) at the NRF-SAIAB. Locality details for Chiloglanis sp. “dwarf”, Chiloglanis sp. “rough skin” and comparative sequences from southern African Chiloglanis species and outgroup species are given in the Suppl. material 1: table $1). Phylogenetic analyses Phylogenetic relationships among Chiloglanis species from southern Africa were inferred using Bayesian and maximum likelihood approaches. Mito- chondrial DNA sequences were edited, aligned, and trimmed in MEGA-X (Ku- mar et al. 2018). The sequences were translated into amino acid sequences in MEGA-X to check for stop codons and gaps to ensure that they were copies of functional mitochondrial protein coding sequences. Haplotype groups were identified using DNASP 6 (Rozas et al. 2017). Bayesian analy- sis was performed using MrBayes 3.1.2 (Ronquist et al. 2012). The dataset was partitioned by codon position and sampled using a reversible-jump Mar- kov chain Monte Carlo (RJ-MCMC), with time-reversible substitution models and gamma distributed rate heterogeneity (Huelsenbeck et al. 2004). Two parallel analyses of four Markov chains were ran for 10 million generations, sampling of trees was done every 1000 generations discarding the first 25% of trees as burn-in. An average standard deviation of split frequencies of < 0.05 indicated the convergence of the two runs in MrBayes. Effective Sample Size (ESS) values and the potential-scale reduction factor for all pa- rameters examined were > 100 and 1.0, respectively. Resulting trees were vi- sualized in FigTree, v. 1.4.3 (Rambaut 2016). Maximum likelihood (ML) anal- ysis of the same dataset was performed in IQ-TREE (Nguyen et al. 2015) through the webserver at http://iqtree.cibiv.univie.ac.at/ (Trifinopoulos et al. 2016). A total of 1000 Ultrafast bootstraps were performed (Minh et al. 2013). Bootstrap values equal to or higher than 70% (Hillis and Bull 1993), and posterior probability values at 0.95 or higher (Alfaro and Holder 2006) were considered to indicate well supported nodes. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 264 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Morphological analysis The study examined 110 specimens of Chiloglanis sp. “rough skin’, 55 spec- imens of Chiloglanis sp. “dwarf”, and five specimens of Chiloglanis sp. “Pun- gwe” from the EZH. These specimens were compared to their southern Af- rican congeners using raw data from Mutizwa et al. (2024). A total of ten counts and 43 measurements (Fig. 2) were considered for this study follow- ing Friel and Vigliotta (2008) and Skelton and White (1990). The morphomet- ric characters are defined in Suppl. material 1: table S2). Measurements were taken to the nearest 0.1 mm using digital callipers. Postcranial meristics (i.e., total vertebrae, abdominal vertebrae, and caudal vertebrae) were taken from radiographs following criteria described by Skelton and White (1990). Verte- brae counts excluded the Weberian vertebrae. The ural centrum was counted as a single element. Abdominal vertebrae included vertebrae anterior to the leading anal-fin pterygiophore. The caudal vertebrae included all vertebrae posterior to the leading anal-fin pterygiophore. The specimens were sexed by external examination. Males possess an elongated and pointed genital papilla, while females are distinguished by possession of longitudinal invagi- nation between the anus and the uro-genital papilla (Skelton 2001). Figure 2. Illustrations of linear measurements recorded from Chiloglanis specimens A lateral view B ventral view of the body C ventral view of the oral disc D dorsal view of the head. Character abbreviations are explained in the morphological analysis section. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 265 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Results Molecular analysis The Maximum Likelihood analysis based on 97 sequences (525 base pairs) recovered 14 distinct clades (Fig. 3). These clades corresponded to the eight species from the genus Chiloglanis that are currently recognised in southern Africa. The remaining clades correspond to the six candidate species that were identified in previous studies and include the two taxa, Chiloglanis sp. ‘rough skin’ and Chiloglanis sp. ‘dwarf’ whose taxonomic integrity is the subject of investigation for the present study (Fig. 3). A similar number of clades was recovered from the Bayesian Inference analysis (Fig. 4). The two lineages, Chiloglanis sp ‘rough skin’ and Chiloglanis sp. ‘dwarf’ were recovered as well-supported monophyletic clades (Figs 3, 4) that are deeply divergent (10.1-18.5%) from all the formally described congeneric species PP156895 PP156891 PP156893 SB9361 SB9364 SB9367 sees? | C. carnatus SB9365 PP156892 PP156894 ANGFW133 ANGFW077 Ic. fasciatus ANGFW131 0Q308636 0Q308634 0Q308632 i casoses2 | C. emarginatus 0Q308635 OL311818 LN610341 poetics ic. pretoriae 0Q308614. LN610269 LN610272 LN610271 oases | C. anoterus LN610270 0Q308613 0Q308616 0Q308615 , OL31 1816 Ic. bifurcus MH432062 MH432061 MH432030 MH432046 MH432057 MH432031 MH432048 MH432027 MH432044, wiasz026 |) CO. COMpactus Sp. nov. MH432019 MH432054. MH432047 MH432018 MH432032 MH432042 MH432026 MH432033 MH432020 " 7 MHaso022 Ic. sp. "Nyangombe MH432021 0Q308696 0Q308695 ogsosee4 |C. swierstrai 0Q308687 W H W MAFW119 HC. Sp. Shire 0Q308653 MH432045 S B9389 SB9355 SB9387 MH432036 MH432060 MH432043, MH432051 MH432037, MH432034, MH432085 - wHasooeo. «= YC. aSperocutis sp. nov. MH432038 MH432053 MH432056 MH432059 MH432055 MH432058 MH432052 MH432041 SB9354 SB9376 SB9373 SB9357 SB9388 O0Q308652 MH432024 0Q308651 =. [c sp. "Zambezi" ANGFW078 ANGFW015 NAB AR MH432029 Ic. Sp. "Pungwe" MH432040 SB8459 wpumaozs fC. paratus KT192823 Euchilichthys royauxi HM418085 Euchilichthys boulengeri MKO073984 Atopochilus savorgnani MKO073983 Atopochilus savorgnani 0.04. Figure 3. Maximum likelihood tree of the species and lineages of the genus Chiloglanis found in southern African based on cytochrome oxidase | (COI). The grey bars represent sequences belonging to the same species. Node colours indicate bootstrap values: black circle = 70; red circle < 70. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 266 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands PP156895 PP 156894 PP 156893 PP156892 PP156891 C. carnatus SB9361 SB9364 SB9365 SB9366 SB9367 ANGFW133 ee IIc. fasciatus ANGFW131 OQ308696 0Q308695 . i 0a308694 [c. swierstrai 0Q308693 OQ308687 " “ " warwie WC. Sp. "Shire 0Q308653 MH432045 MH432036 SB9355 SB9387 SB9389 MH432060 MH432059 MH432055 MH432058 MH432052 MH432041 icine C. asperocutis sp. nov. MH432053 MH432051 MH432050 MH432043 MH432038 MH432037 MH432035 MH432034 PQ424602 PQ424604 PQ424605 PQ424606 PQ424608 0Q308652 0Q308651 MH432024 MH432023 W “WH pasewort |: SP. "Zambezi ANGFW078 ANGFW015 MH432049 MH432029 ‘4 " MH432028 Ic sp. "Pungwe MH432040 0Q308636 0Q308635 casosss4 | C. emarginatus 0Q308633 0Q308632 OL311818 : ann, Ic. pretoriae LN610341 0Q308616 0Q308615 , OL311816 [c. bifurcus MH432062 0Q308614 0Q308613 0Q308612 ineio272 CO. anoterus LN610271 LN610270 LN610269 MH432061 MH432046 MH432030 MH432057 MH432054 MH432048 MH438048 mHasc0aa =| C. COMpactus Sp. nov. MH432042 MH432026 MH432032 MH432031 MH432027 MH432025 MH432019 MH432033 MH432022 MH432021 Ic. sp. "Nyangombe" MH432020 SB8459 MPUMA025 fic. paratus KT192823_Euchilichthys_royauxi HM418085_Euchilichthys_boulengeri MK073984_Atopochilus_savorgnani MK073983_Atopochilus_savorgnani 0.03 Figure 4. Bayesian inference tree of the species and lineages of the genus Chiloglanis found in southern African based on cytochrome oxidase | (COI). The grey bars represent sequences belonging to the same species. Node colours indicate Bayesian posterior probabilities: black circle = 0.95; red circle < 0.95.. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 267 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands in southern Africa (Table 1). Chiloglanis sp. “dwarf” was deeply divergent from Chiloglanis sp. “rough skin” (11.8-13.5%, Table 1). Intraspecific genetic divergence was low in Chiloglanis sp. “dwarf” (0-0.8%) and Chiloglanis sp. “rough skin” (0-1.7%). Chiloglanis sp. “dwarf” was recovered as sister to an undescribed lineage from the Zambezi River, Chiloglanis sp. “Nyangombe’” with a genetic divergence of 4.1-4.8%. Chiloglanis sp. “rough skin” sp. nov. was recovered as sister to a clade with undescribed lineages from the Pungwe River, Chiloglanis sp. “Pungwe”, and the neighbouring Zambezi River, Chiloglanis sp. “Zambezi”. There was relatively low genetic divergence between Chiloglanis sp. “rough skin”, Chiloglanis sp. “Pungwe” (1.7-3.3%) and Chiloglanis sp. “Zambezi” (2.7-4.1%). However, the low genetic divergence between Chiloglanis sp. “rough skin’, Chiloglanis sp. “Pungwe”, and Chiloglanis sp. “Zambezi” was in some cases higher than the interspecific genetic divergence found between some valid southern African Chiloglanis species. For example, C. anoterus Crass, 1960 and C. bifurcus Jubb and Le Roux, 1969 were separated by 2.5-2.9% genetic divergence while C. emarginatus and C. pretoriae were separated by 3.7% (Table 1). The clade formed by Chiloglanis sp. “rough skin’, Chiloglanis sp. “Pungwe’” and Chiloglanis sp. “Zambezi” was separated from Chiloglanis sp. “Shire” a lineage from the Shire River by a genetic divergence of 4.7-6.2%. Both Chiloglanis sp. “rough skin” and Chiloglanis sp. “dwarf” occurred in sympatry within the Buzi River system. Within the Pungwe River, Chiloglanis sp. “rough skin” and Chiloglanis sp. “dwarf” co-occurred with Chiloglanis sp. “Pungwe” (Fig. 1). These results further corroborate the presence of undescribed Chiloglanis lineages, such as Chiloglanis sp. “Nyangombe” from the Nyangombe River (Zambezi River system), Chiloglanis sp. “Shire” from the Shire River (Zambezi River system), Chiloglanis sp. “Pungwe” from the Pungwe River and Chiloglanis sp. “Zambezi” from the Zambezi River that were identified in previous studies. Morphological analysis Detailed morphological examination revealed a number of phenotypic charac- ters that consistently separate Chiloglanis sp ‘rough skin’ and Chiloglanis sp. ‘dwarf’ from all the eight formally described and allopatrically distributed spe- cies from southern Africa. These characters include the caudal peduncle depth, dorsal-fin base length, lower lip length, mandibular tooth row width, upper lip length, number of pectoral-fin rays, number of dorsal-fin rays, number of man- dibular teeth, number of primary premaxillary teeth, shape and depth of the caudal fin and the shape of the tubercles (Fig. 5, Table 2). Taxonomic decisions Based on the genetic and morphological divergence of Chiloglanis sp. ‘rough skin’ and Chiloglanis sp. ‘dwarf’ from all the formally described con- geners from southern Africa and the other candidate species identified from the Eastern Zimbabwe Highlands freshwater ecoregion, below we pro- vide formal descriptions of these two species as Chiloglanis asperocutis sp. nov. and Chiloglanis compactus sp. nov., respectively. The other lineages ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 268 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands “Trz] zt osresr|ruen|eore sa zocoaileoselssirsi| se |rocesleses| 6 |reeajeseuleesa| —pnetor 3] a OEL-B°CL | BLL-OLL | L9L-G9OL Vel-eLL SOL L8L-G9OL | O8L-8ZL | CSL-6V7L | ELL-ELL | OZL-L ZL | OLZL-O'ZL | CLL-OLL | O9L-L9OL | O8L-GZL | O8L-O'ZLL wabuajnog "7 OL — TL-0 | v61-€81 OZL-S9OL | 68L—-OZL | COL-ESL | COL-GZL | OGL-S'SL | SLZL-L'ZL | GZL-@LL | LBL-SZL | L8L-8ZL | BZL-EZLL | S6L-CEL | ESL-S'ZL Qol-78L, ——ueub ones y | SL ,quobuekn,, c 0-0 teeta SEL-B LL | SSlL-29L | OCL-VeL| ELI-9-OL | OCL-S°EL.| SOL-LOL | SEL-GSEL | SCL-SLL | S*ZL-9OL | eal 6Z1L-2£LL |S LL-¢ LE| C@EL-SCL| OSL-S LL |Z UL-S LL) SSI-EPE| CEL-E CL) SPL-ZEL 6 ul ZOQUEZ,, ° ‘aou ‘ds sinoosedse siuesbojiy9 ‘aou ‘ds snyoedwoo sjuesboyiy9 a ee } — 90-00 SLL SSL-9PL| se nisiomms “9 a ee ee eee Se ajo} se | wm |e | aw |] mw | m | 6 | e@ | e« | os | os fe fee |e te | ‘Apnys Juasaid au} ul papnjoul saioads sjuejbojiyD ay} uaamiaq (%) Saduejsip 1}9UaB (|OD) | asepixo awWOJYDOJAD Jo sabuey “1 ajqeL 269 ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Figure 5. Some of the readily identifiable diagnostic characters separating Chiloglanis asperocutis from C. compactus. Ridge-like tubercles on the dorsum of a C. asperocutis and conical tubercles on b C. compactus. Deeply forked caudal peduncle with pointed lobes in c C. asperocutis and shallow fork with rounded lobes in d C. compactus. Table 2. Morphological characters used to distinguish Chiloglanis compactus sp. nov. and Chiloglanis asperocutis sp. nov. from the valid Chiloglanis species found in southern Africa. ; Collogiauts. | Chuteg rans Chiloglanis | Chiloglanis | Chiloglanis | Chiloglanis | Chiloglanis | Chiloglanis | Chiloglanis | Chiloglanis Species compactus | asperocutis i : ‘ ; ; : carnatus __ pretoriae | anoterus bifurcus | emarginatus | fasciatus paratus __swierstrai Sp. nov. Sp. nov. Number of 55 110 19 16 1.0 10 10 10 2 F specimens Total length 28.9-57.2 | .32:3-87:6: |45.3-62:2 | 31:7-67.1 80.1 68.7-84.9 | 50.2-66.6 | 37.7-53.3 | 44-51.9 | 45.2-65.7 Standard length 26.0-45.9 | 33.9-66.6 | 35.5-48.9 | 26.5-54.6 61.7 51.4-63.9 | 40.3-55.6 | 30.3-41.7 | 35.8-42.4 | 34.9-51.9 Head length 7.1-14.4 | 11.2-22.8 | 12.1-15.6 | 8.8-19.4 20.3 1S719.5. | 123=15:7 | 100-137 | 111-1328 | .9.2=13:6 % Standard length Caudal peduncle 10.0-12.4 | 7.5-10.0 | 11.3-13.2 | 11.0-13.8 122 11.1-14.1 | 10.2-11.9 7.5-8.8 9.6-9.9 7.2-8.7 depth Dorsal-fin base length | 9.9-14.2 | 7.9-13.4 | 10.7-14.1 | 12.8-18.0 8.6 95=13:2 8.8-12.9 | 10.4-13.7 | 12.8-14.6 | 7.5-9.6 % Head length Lower lip length 1#25=22.6- | -18:4-27.4 | 18.3-26:6: | -22.A=27°7 251 V7 725-6 | 23:5528.8: 0 || 18.8-24:9°) 22.8-27:6. | $9:2—26:0 Mandibular tooth row | 5.6-10.4 36-72 4.6-8.1 16.0-25.6 10:5 10:4=17.3-| -9:6-13:5 4.8-6.6 F2Z-TF, | V0.0-16:6 width Upper lip length 6.9-17.1 | 11.4-22.2 | 11.1-16.2 | 11.7-18.8 16.7 8.4-12.3 6.6-10.6 8.8-15.5 | 12.3-12.4 | 7.0-10.5 Meristics Pectoral-fin rays 7 8 8 (6-8) 8 8.0 8 (7-8) 7 8 8 8 Dorsal-fin rays 5 6 (6-7) 6 (5-7) 6 5.0 5 (5-6) 6 (5-6) 6 (5-6) 5 5 (5-6) Mandibular teeth 8 (6-8) 10 (8-10) 10 12 12.0 8 8 (6-8) 8 12 11 (11-14) Primary premaxillary | 38(31-53) | 80(68- | 60(43-69)| 51-59 86.0 54 (50-64) 36-54 64 (51-65) | 39-51 50 (34-59) teeth 128) ZooKeys 1241: 261-290 (2025), DOI: 10.38 270 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands identified from the previous and present study will be considered for de- scription in a future study once we have gathered additional material for detailed morphological examination. Taxonomic accounts Chiloglanis asperocutis Mutizwa, Braganga & Chakona, sp. nov. https://zoobank.org/ODDBD844-CDOF-432C-9AD0-E11FE7C7095A Figs 6, 7 Chiloglanis sp. “rough skin’: Chakona et al. 2018: 76-79. Type material. Holotype. » SAIAB 246255, female specimen 66.0 mm SL, Honde River, Bridge on road to Honde Mission, Pungwe River system, Manicaland Prov- ince, Zimbabwe, 18.5438°S, 32.8044°E, 13 December 2014, Albert Chakona, Wilbert Kadye and Taurai Bere, genseq-1 COI MH432036. Paratypes. » SAIAB 201026, 14 male, 29 female specimens (32.3-77.5 mm SL), Honde River, Pungwe River system, Bridge on road to Honde Mission, Manicaland Province, Zimbabwe, 18.5438°S, 32.8044°E, 13 December 2014, Albert Chakona, Wilbert Kadye and Taurai Bere. Other material examined. Specimens detailed in Suppl. material 1. Diagnosis. A higher number of primary premaxillary teeth (68-128) readily distinguishes C. asperocutis sp. nov. from all its congeners from southern Af- rican that consistently have fewer than 68 primary premaxillary teeth except for C. anoterus and C. carnatus Mutizwa et al., 2024. Possession of ten closely packed mandibular teeth further distinguishes C. asperocutis from C. fasciatus Pellegrin, 1936 that has eight closely packed mandibular teeth; C. compactus sp. nov., C. bifurcus and C. emarginatus that have eight widely spaced mandibular teeth; C. anoterus, C. paratus Crass, 1960, and C. pretoriae that have 12 close- ly packed mandibular teeth; and C. swierstrai Van der Horst, 1931 that has 14 closely packed mandibular teeth. Chiloglanis asperocutis sp. nov. possesses more dorsal fin rays (6-7) compared to C. compactus sp. nov. (5 rays). Posses- sion of eight pectoral fin rays distinguishes C. asperocutis from C. compactus sp. nov. and C. emarginatus that possess seven pectoral fin rays. A deeply forked caudal fin readily separates C. asperocutis sp. nov. from C. compactus sp. nov., C. pretoriae, C. paratus, and C. swierstrai that have shallower fork depths in the caudal fin; C. emarginatus that has an emarginated caudal fin with a very shallow fork; and from C. anoterus that has a caudal fin with no fork due to extended median rays in males and emarginated in females. A caudal fin with an upper lobe shorter than lower lobe distinguish C. asperocutis sp. nov. from C. bifurcus that has a caudal fin with an upper lobe that is longer than the lower lobe. An oral disc with a well-developed mid-ventral cleft distinguishes C. asperocutis sp. nov. from C. swierstrai that lacks a mid-ventral cleft. Chiloglanis asperocutis sp. nov. has a dorsal spine with gently serrated anterior and posterior margins that distinguish it from C. paratus that has a dorsal spine with a serrated posterior margin. Absence of a fleshy skin on the basal portion of the dorsal fin separates C. asperocutis sp. nov. from C. carnatus, a character present in the latter spe- cies. Chiloglanis asperocutis sp. nov. is further distinguished from its southern African congeners by long ridge-like tubercles distributed across the dorsal and ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 271 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Figure 6. Chiloglanis asperocutis female holotype SAIAB 246255. Scale bars: 2 cm. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 272 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Figure 7. Chiloglanis asperocutis male paratype SAIAB 201026. Scale bars: 2 cm. lateral surfaces of the head and body compared to conical tubercles found in C. compactus sp. nov., C. bifurcus, C. emarginatus, C. fasciatus, C. paratus, C. car- natus, and C. swierstrai; and the lack of conspicuous tubercles in C. anoterus and C. pretoriae. A narrow caudal peduncle depth distinguishes C. asperocutis sp. nov. (7.5-10.0%SL) from C. carnatus (11.3-13.2%SL), C. pretoriae (11.0- 13.8%SL), C. anoterus (12.2%SL), C. bifurcus (11.1-14.1%SL), and C. emargin- atus (10.2-11.9%SL) that have deeper caudal peduncles. A narrow mandibular ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 273 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands tooth row width separates C. asperocutis sp. nov. (3.6-7.2%HL) from C. pretoriae (16.0-25.6%HL), C. anoterus (10.5%HL), C. bifurcus (10.4-17.3%HL), C. emargin- atus (9.6-13.5%HL), C. paratus (7.2-7.7%HL), and C. swierstrai (10.0-16.6%HL) that have wider mandibular tooth row widths. A longer upper lip length further separates C. asperocutis sp. nov. (11.4-22.2%HL) from C. emarginatus (6.6- 10.6%HL) and C. swierstrai (7.0-10.5%HL) that have shorter upper lips. Description. Figs 6, 7 shows general body features of adult male and female C. asperocutis. Morphometric and meristic data for holotype, paratypes, and non-type specimens are presented in Table 3. Where meristics counts have a range, counts from the holotype are given in parentheses. Body elongate. Depressed pre-dorsal region, mid-sections more cylindri- cal, slightly laterally compressed posterior section. Pre-dorsal profile convex, sharply slopping from snout to posterior nostril, gentler slope from poster nos- tril to dorsal-fin origin. Body greatest depth at dorsal-fin insertion. Post-dorsal profile almost straight to adipose fin insertion, gently concave from adipose-fin origin to caudal fin. Ventral profile gently convex from region just posterior of oral disc to anal-fin origin, gently concave from anal-fin origin to caudal fin. Skin numerous tubercles spread across body with exception of ventral sur- face, forming distinct ridge like structures most conspicuous on head dorsum giving skin a distinct “rough” texture. Lateral line complete, originating just above horizontal level of orbit, below anterior dorsal fin origin, gently undulating along midlateral of body to base of caudal fin. Head relatively big; longer than body depth; ~ 2/3 of pre-dorsal length. Eye large and oval, orbit lacks free margin; located dorsally, closer to margin of operculum than tip of snout. Interorbital space slightly wider than space between nares spac- es. Posterior nares slightly closer together than anterior nares. Anterior nares with short skin flaps on their posterior margins. Posterior nares with short skin flaps along anterior margin. Gill opening restricted, located adjacent to pectoral fin origin. Mouth inferior, projecting below ventral body surface, upper and lower lips combined into a distinct large oral disc. Oral disc width greater than length. Upper and lower lips with pronounced roundish papillae, largest papillae con- centrated around mid-ventral cleft of lower lip. Three unbranched pairs of bar- bels. Maxillary barbel, originating from lateral region of oral disc, extending to posterior region of oral disc. Lateral mandibular barbel longer than medial man- dibular barbel, both integrated into lower lip. Dentation variable number 68-128 (102) of primary pre-maxillary teeth ar- ranged in 3-5 (4) rows on two ovoid tooth patches. Secondary pre-maxillary teeth small and scattered on posterior surface of premaxillae. Tertiary teeth small and needle-like; in a row near midline of dorsal edge of tooth plate. Up to 10 (10) closely packed mandibular teeth; central teeth projecting higher than outer teeth forming a gentle arc; replacement tooth row emerges anteriorly to functional row, number of mandibular teeth may vary due to loss of teeth and their replacements, but the number does not exceed ten. Vertebral counts Total vertebrae 27-29 (29), abdominal vertebrae 11-13 (13), caudal vertebrae 15-17 (16). Urogenital papillae distinctly elongate and sharply pointed in males, reduced to a shallow invagination in females. Pectoral fin ray count 8, origin adjacent to gill opening; pectoral spine slightly curved, anterior margin smooth; gently serrated posterior margin; pectoral spine ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 274 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Table 3. Summary of the Chiloglanis asperocutis sp. nov. morphological characters. All measured values, except stan- dard length (SL) and Head length (HL) are given as percentages of the HL or SL. The numbers in the parentheses repre- sent the mean of the morphometric characters and the mode for the meristic characters. Chiloglanis asperocutis holotype | Chiloglanis asperocutis paratypes | Chiloglanis asperocutis non-types SAIAB catalogue number 201026 201026 - Number of specimens 1 42 67 Total length 81.6 32.3-77.5 43.5-87.6 (65.1) Head length 21.5 17.8-21.5 11.2-22.8 (16.8) Standard length 66 61.4-66 33.9-66.6 (51.7) % standard length Adipose fin to caudal peduncle length 17.6 16-17.6 13.5-21.6 (16.4) Adipose-fin base length 11? 11.7-14.5 10.4-17.9 (13.7) Adipose-fin height 4.7 4.7-4.9 3.9-7 (5.4) Anal-fin base length 9.7 9.7 8.2-13.2 (10.9) Anal-fin length along longest ray 12.1 1271-1631 10.9-18.9 (14) Body depth at anus 12.8 12.8-15.4 12.5-16.9 (14.8) Body depth at dorsal-fin insertion 19.6 18.1-19.6 17.2-22.8 (19.4) Caudal peduncle depth 8.1 8.1-8.4 7.5-10 (8.9) Caudal peduncle length 19.1 18.8-19.1 13.9-21.3 (17.4) Dorsal fin to adipose fin length 26.4 24.9-26.4 18.7-29.2 (24.6) Dorsal-fin base length 12.4 11.6-12.4 7.9-13.4 (10.2) Dorsal-spine length 19.4 16.7-19.4 13.8-25.4 (17.5) Pre-anal length 73.6 70.4-73.6 68.4-77.6 (72.8) Pre-dorsal length 42.5 39.2-42.5 37.6-46.3 (40.9) Pre-pectoral length 29 26.5-29 25.733 (29.7) Pre-pelvic length 57.8 57.1-57.8 53.6-61.8 (58.8) Pectoral-spine length 20.8 20.8-21.3 13.4-26.6 (20.2) Pectoral-fin length 21.8 21.8-25.5 19.4-27.4 (22.7) Pelvic-fin length 14.3 13.2-14.3 1.7175 (14.6) Width at pectoral-fin insertion 23.9 227-239 21-2721 (23:9) Pelvic-fin interspace 4.8 4.8-5.4 2.8-9.2 (5.1) Head length 32.6 29-32.6 28.6-36.3 (32.6) Dorsal-fin length along longest ray 20.5 20.5-20.6 12.3-22.8 (17.8) Caudal fork length 10.3 8.5-10.3 9.4-15.9 (12.5) % Head length Anterior nares interspace 13.9 13.4-13.9 10.3-19.5 (14.4) Eye diameter (vertical axis) 12.5 12.5-13.4 7.9-15.9 (12.7) Lower lip length 21.3 18.7-21.3 18.4-27.4 (22.4) Mandibular tooth row width 6.4 6.4-7.2 3.6-6.9 (5.5) Maxillary barbel length 27.8 23.9-27.8 20.2-38.9 (28.3) Mouth width 25.9 25.9-31 26.6-34.1 (31) Orbital interspace 20.8 20.4-20.8 16.5-22.6 (19.9) Oral disc length 57.2 53.7-57.2 47.1-63.3 (52.6) Oral disc width 51.7 51.7-59 49.6-69.2 (57.9) Premaxillary tooth-patch length 10.5 LOT 10:5 7.6-14 (10.6) Premaxillary tooth-patch width 36.4 36.4-46.8 29.7-51.2 (43.5) Posterior nares interspace 17.3 10.1-17.3 7.5-14.9 (11.3) Snout length 65 60.1-65 54.8-68 (63.9) Upper lip length 13 13-16.8 11.4-22.2 (15.6) Eye diameter (horizontal axis) 18:7 1a7 7 22.3-51.4 (34.2) Occipital shield width 36.1 36.1-40.1 48.4-60 (54.2) Head depth 52 52-558 22.5-89.5 (61.6) Meristics Pelvic fin count Pectoral fin count ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 275 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Chiloglanis asperocutis holotype | Chiloglanis asperocutis paratypes | Chiloglanis asperocutis non-types Primary premaxillary tooth row count 4 3-5 4 (3-5) Primary premaxillary tooth count total 102 74-102 80 (68-128) Mandibular tooth count 10 10 10 (8-10) Dorsal-fin rays 6 6 6 (6-7) Anal-fin rays 10 LOS 18 9 (9-11) Total vertebrate 29 28-29 27 (27-29) Abdominal vertebrae 13 11=13 12°(11=13) Caudal vertebrae 16 15-17 15-17 length ~80% of longest pectoral-fin ray. Dorsal fin originating in anterior 1/3 of body, posterior to pectoral-fin origin, 6-7 (6) branched rays. Dorsal spine, length ~ 80% of longest dorsal-fin ray, gentle serrations on anterior and posterior mar- gins of the terminal 1/4 of dorsal spine. Pelvic fin ray count 7, origin posterior to midpoint between end of dorsal-fin and adipose-fin origin; rounded. Adipose fin relatively small, margin convex, located in posterior 1/3 of body, origin just anterior of anal-fin origin, ridge-like tubercles sometimes present on its surfaces. Anal fin ray count 12-13 (13), origin just posterior to origin of adipose fin; termi- nating just before posterior end of adipose fin; rounded. Caudal fin deeply forked, lobes unequal, lower lobe generally longer, both lobes with gently pointed tips. Coloration in ethanol dorsal and lateral skin generally dark brown becom- ing a lighter shade of brown below mid lateral line. Dark melanophores distrib- uted throughout dorsal and lateral surfaces. Dorsal surface has blotches of lighter shades of brown located between orbits and posterior nostril, at dorsal fin insertion, between dorsal and adipose fin, and posterior to adipose fin, all these blotches extend to lateral surface. Lateral surface with large pale blotch- es such as those extending from dorsal surface. Lateral line clearly visible as a continuous pale brown stripe with very small pale blotches distributed above and below it. Ventral surface pale cream to yellowish in colour with dark melanophores only present at base of pelvic fins. Dorsal and adipose fins dark brown base and middle with hyaline margins. Pectoral fins have dark brown dorsal surfaces and lighter ventral surface, hyaline margins. Pelvic and anal fins pale brown in colour with dark melanophores and hyaline margins. Dark band stretches across middle of upper caudal fin lobe, lower lobe mostly cov- ered by a dark blotch originating from the fin base. Live colouration displayed similar patterns as described in persevered specimens with a few differences. All pale brown sections in live specimens have a golden colour. Live specimens display a range of pigmentation intensity, some specimens are very dark with patterns described in the Coloration in ethanol section being very clear whilst other specimens are lightly coloured (golden) with little pigmentation visible. Reproduction: there are no dedicated studies on the breeding biology of C. asperocutis, but spawning is likely to begin in summer (October-November) based on the general pattern of other congeners (Skelton 2001), and other mo- chokid fishes from this region. Distribution, habitat, and feeding ecology. this species is known from mul- tiple localities in the Pungwe and Buzi River systems. It is a rheophilic species that occurs in rocky habitats with fast flowing water. Macroinvertebrates from the families Simuliidae, Chironomidae, Hydropsychidae and Libellulidae were the dominant prey item for this species with algae forming a smaller compo- nent of the diet of this species (Matomela et al 2018). ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 276 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Etymology. a combination of the Latin words, aspero, meaning rough, and cutis, meaning skin. The name refers to the distinct ridges on the skin (more pronounced on the head dorsum) which is characteristic of this species. Chiloglanis compactus Mutizwa, Braganga & Chakona, sp. nov. https://zoobank.org/3EECOC8D-4992-4A8C-9226-1 BE59BAAB312 Figs 8,9 Chiloglanis sp. “dwarf”: Chakona et al. 2018: 76-79. Type material. Holotype. * SAIAB 246256, male, right middle cut, 44.4 mm SL, Makanga River Bridge on road to Hauna growth point, Pungwe River system, Manicaland Province, Zimbabwe, 18.5438°S, 32.801°E, 16 December 2014, Albert Chakona, Wilbert Kadye and Taurai Bere, genseg-1 COI] MH432044. Paratypes. * SAIAB 210377, 10 males (29.8-37.5 mm SL), 2 females (29.1- 40.3 mm SL), Makanga River bridge on road to Hauna growth point, Pungwe River system, Manicaland Province, Zimbabwe, 18.5438°S, 32.801°E, 16 De- cember 2014, Albert Chakona, Wilbert Kadye and Taurai Bere. Other material examined. Specimens detailed in Suppl. material 1. Diagnosis. Chiloglanis compactus sp. nov. attains the smallest size (< 46 mm SL) for all congeners currently known from southern Africa. Pos- session of seven pectoral fin rays in C. compactus sp. nov. distinguishes this species from C. asperocutis, C. carnatus, C. anoterus, C. fasciatus, C. paratus, C. pretoriae, and C. swierstrai that all have eight pectoral fin rays. Possession of eight widely spaced mandibular teeth further distinguishes C. compactus sp. nov. from C. fasciatus (eight closely packed mandibular teeth); C. asperocutis and C. carnatus (ten closely packed mandibular teeth); C. anoterus, C. paratus, and C. pretoriae (12 closely packed mandibular teeth); and C. swierstrai (14 closely packed mandibular teeth). Fewer primary premaxillary teeth in C. com- pactus sp. nov. (31-53) readily distinguished it from C. asperocutis (68-128) and C. anoterus (86). Fewer dorsal fin rays (5) readily distinguish C. compactus sp. nov. from C. asperocutis (6-7). A relatively shallow forked tail and round- ed caudal fin lobes readily separate C. compactus sp. nov. from C. asperocu- tis, C. carnatus, and C. fasciatus (deeply forked caudal fin with pointed lobes); C. emarginatus (emarginated caudal fin with very shallow fork); and from C. anoterus that has a caudal fin with no fork due to extended median rays in males and emarginated in females. A lower caudal fin lobe longer than the up- per lobe separates C. compactus sp. nov. from C. bifurcus that has a longer up- per caudal fin lobe. A well-developed mid-ventral cleft in the oral disc of C. com- pactus sp. nov. distinguishes it from C. swierstrai that lacks a mid-ventral cleft. Gently serrate anterior and posterior margins of the dorsal spine in C. compac- tus sp. nov. distinguish it from C. paratus that has a dorsal spine serrated only on the posterior margin. Absence of a fleshy skin on the basal portion of the dorsal fin separates C. compactus sp. nov. from C. carnatus, a character pres- ent in the latter species. Conical tubercles distributed across dorsal and lateral surfaces of the head and body distinguish C. compactus sp. nov. from C. asper- ocutis with ridge like tubercles; C. anoterus and C. pretoriae that lack conspic- uous tubercles. A deep caudal peduncle distinguishes C. compactus sp. nov. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 277 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Figure 8. Chiloglanis compactus male holotype SAIAB 246256. Scale bars: 2 cm. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 278 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Figure 9. Chiloglanis compactus female paratype SAIAB 210377. Scale bars:2 cm. (10.0-12.4%SL) from C. asperocutis (7.5-10.0%SL), C. fasciatus (7.5-8.8%SL), C. paratus (9.6-9.9%SL), and C. swierstrai (7.2-8.7%SL) that have narrower caudal peduncles. A longer dorsal-fin base separates C. compactus sp. nov. (9.9-14.2%SL) from C. swierstrai (7.5-9.6%SL) and C. anoterus (8.6%SL) that possess shorter dorsal-fin bases. Shorter lower lips distinguish C. compactus ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 279 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands sp. nov. (17.5-22.6%HL) from C. pretoriae (22.4-27.7%HL), C. anoterus (25.1%HL), C. emarginatus (23.5-28.8%HL), and C. paratus (22.8-27.6%HL) that have longer lower lips. A narrow mandibular tooth row separates C. com- pactus sp. nov. (5.6-10.4%HL) from C. asperocutis (3.6—7.2%HL), C. pretoriae (16.0-25.6%HL), C. bifurcus (10.4-17.3%HL), and C. swierstrai (10.0-16.6%HL) that possess greater mandibular tooth row widths. Description. Figs 8, 9 shows general body features of adult male and female C. compactus. Morphometric and meristic data for holotype, paratypes and non-type specimens are presented in Table 4. Where meristics counts have a range counts from the holotype are given in parentheses. Body short and rotund. Depressed pre-dorsal region, mid-sections more cy- lindrical, slightly laterally compressed posterior section. Pre-dorsal profile con- vex, almost continuous rounded profile from snout to dorsal-fin origin. Body greatest depth at dorsal-fin insertion. Post-dorsal profile convex to adipose fin insertion, becoming gently concave from adipose-fin origin to caudal fin. Ventral profile convex from region just posterior of oral disc to anal-fin origin, becoming gently concave from anal-fin origin to caudal fin. Skin numerous conical tubercles spread across body with exception of ven- tral surface. Lateral line complete, originating above horizontal level of orbit, below anterior dorsal fin origin, almost completely straight along midlateral of body to base of caudal fin. Head relatively big; longer than body depth; ~ 2/3 of pre-dorsal length. Eye large and oval, orbit lacks free margin; located dorsally, closer to margin of operculum than tip of snout. Interorbital space wider than space between nares spaces. Posterior nares slightly closer together than anterior nares. Anterior nares with short skin flaps on posterior margin. Posterior nares with short skin flaps along anterior margin. Gill opening restricted, located adjacent to pectoral fin origin. Mouth inferior, projecting below ventral body surface, upper and lower lips combined into a distinct large oral disc. Oral disc width greater than length. Upper and lower lips with pronounced roundish papillae, largest papillae con- centrated around mid-ventral cleft of lower lip. Three pairs of barbels. Maxil- lary barbel unbranched, originating from lateral region of oral disc, extending to posterior region of oral disc. Lateral mandibular barbel longer than medial mandibular barbel, both integrated into lower lip. Dentation variable number 31-53 (44) of primary pre-maxillary teeth ar- ranged in 2-3 (3) rows on two ovoid tooth patches. Secondary pre-maxillary teeth small and scattered on posterior surface of premaxillae. Tertiary teeth small and needle-like; in a row near midline of dorsal edge of tooth plate. Up to 8 (6) spaced out mandibular teeth; replacement tooth row emerges anteriorly to the functional row, number of mandibular teeth may vary due to loss of teeth and their replacement, but the number hardly ever exceeds 8. Vertebral counts Total vertebrae 27-29 (28), abdominal vertebrae 11-12 (11), caudal vertebrae 17-18 (17) Urogenital papillae distinctly elongate and sharply pointed in males, reduced to a shallow invagination in females. Pectoral fin ray count 7, origin adjacent to gill opening; pectoral spine slightly curved, anterior margin smooth; gently serrated posterior margin; pectoral spine length ~ 80% of longest pectoral-fin ray. Dorsal fin originating in anterior 1/3 of body, posterior to pectoral-fin origin, five branched rays, con- ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 280 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Table 4. Summary of Chiloglanis compactus sp. nov. morphological characters. All measured values, except standard length (SL) and Head length (HL) are given as percentages of the HL or SL. The numbers in the parentheses represent the mean of the morphometric characters and the mode for the meristic characters. % standard length Adipose fin to caudal peduncle length 11.3-19.8 (16) Adipose-fin base length 13:2=19°6:(15.7) Anal-fin base length 10.6-16.4 (13.5) Anal-fin length along longest ray 11.8-20.5 (16) Body depth at anus 14.3-20.8 (17.9) Body depth at dorsal-fin insertion 16.8-25.3 (21) Caudal peduncle depth 10-12.4 (11.4) Caudal peduncle length 15.6-24.5 (19.9) Dorsal fin to adipose fin length 19.5-30.2 (24.2) Dorsalfin base length 9.9-11.8 (11.1) Dorsal-spine length 11.6-21.6 (15.7) Pre-dorsal length 35.4-46.2 (39.7) Pre-pectoral length 24.3-33.6 (29.3) Pre-pelvic length 51.5-59.3 (55.8) Pectoral-spine length 137-21, 17.1) Pectoral-fin length 15.4-25.6 (21.2) Width at pectoral-fin insertion 21.6-27.6 (24.2) Dorsal-fin length along longest ray 7.6-22.4 (15.6) Caudal fork length 12.4-24.5 (17.5) % Head length Anterior nares interspace 10.9-23 (16.4) Eye diameter (vertical axis) 11.7-18 (15) Mandibular tooth row width 5,9-10.4 (8.6) Maxillary barbel length 17.4-45.7 (30.6) Orbital interspace 24,.7-27.5 (25.7) Oral disc length 38.6-54.7 (47.6) Oral disc width 43.8-69.7 (51.7) Premaxillary tooth-patch length a ee ee 5.4-14.3 (8.6) Premaxillary tooth-patch width 29.8-46.8 (38.9) Posterior nares interspace 12.8-20 (15.6) Eye diameter (horizontal axis) 22.9-46 (35.5) Occipital shield width 68.7-83.6 (74.3) Meristics ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 281 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Chiloglanis compactus holotype | Chiloglanis compactus paratypes Chiloglanis compactus non-types Primary premaxillary tooth row count 3 2-3 2: (2-3) Primary premaxillary tooth count total 44 42-47 38 (31-53) Mandibular tooth count 6 6 8 (6-8) Dorsal-fin rays 5 5 5 Anal-fin rays 10 9-10 10 (9-12) Total vertebrate 28 27-29 29 (27-29) Abdominal vertebrae 11 11-12 11 (11-12) Caudal vertebrae 17 17-18 17 (17-18) ical tubercles sometime present on its surfaces. Dorsal spine, length ~ 80% of longest dorsal-fin ray, gentle serrations on anterior and posterior margins of terminal 1/4 of dorsal spine. Pelvic fin ray count 7, origin posterior to mid- point between end of dorsal-fin and adipose-fin origin; rounded. Adipose fin relatively small, margin convex, located in posterior 1/3 of body, origin just anterior of anal-fin origin, conical tubercles sometimes present on its surfac- es. Anal fin ray count 12-13 (13), origin just posterior to origin of adipose fin; terminating just before posterior end of adipose fin; rounded. Caudal fin with shallow fork, lobes unequal with lower slightly longer, both lobes rounded, conical tubercles sometime present on their surfaces. Coloration in ethanol dorsal and lateral skin generally dark brown becoming a lighter shade of brown below mid lateral line. Dark melanophores distribut- ed throughout dorsal and lateral surfaces. Dorsal surface has large blotches of lighter shades of brown located at base of dorsal fin, between dorsal and adipose fin, and posterior end of adipose fin; these blotches extend to lateral surface. Lateral surface with large pale blotches such as those extending from the dorsal surface. Additional large blotches on lateral surface are found below lateral line above pelvic fin, anal fin and caudal peduncle. Approximately seven small circular blotches just above lateral line, two or three similar blotches found above each of them. Approximately four small circular blotches below the lateral line distributed from below dorsal fin to pelvic fin. Lateral line clearly visible as a continuous pale brown stripe. Ventral surface pale cream to yellowish in colour with dark melanophores only present at base of pelvic fins and caudal peduncle. Dorsal and adipose fins brown base and middle with hyaline margins. Pectoral fins have dark brown dorsal surfaces and lighter ventral surface, hyaline mar- gins. Pelvic and anal fins pale brown in colour with hyaline margins. Dark band stretches across middle of upper caudal fin lobe, lower lobe mostly covered by a dark blotch originating from the fin base. Live colouration displayed similar patterns as described in the preserved specimens with a few differences. Live specimens display a range of pigmentation intensity, some specimens are very dark with patterns described in the Coloration in ethanol section being very clear whilst other specimens are lightly coloured with little pigmentation visible. Reproduction: there have been no dedicated studies on the breeding biology of C. compactus, but spawning is likely to begin in summer (October—Novem- ber) based on the general pattern of other congeners (Skelton 2001), and other mochokid fishes from this region. Distribution, habitat, and feeding ecology. this species occurred at multiple localities in the Pungwe and Buzi river systems with the majority of the collec- tions occurring at high elevation. It is a rheophilic species that occurs in rocky habitats with fast flowing water. Its diet was not examined. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 282 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Etymology. the name is drawn from the word compact which is inspired by the short and rotund body shape of this species as well as it being the smallest of all the currently known congeners in southern Africa. Discussion Consistent with Marshall's (2011) proposition regarding the taxonomic uncer- tainties within the genus Chiloglanis in central southern Africa, and the subse- quent work by Chakona et al. (2018), this study identified two new suckermouth catfishes that were previously lumped together into a single species, C. neu- manni. The two newly described species, C. asperocutis and C. compactus, are sympatrically distributed in the Pungwe and Buzi river systems. The description of these new species from the EHZ, together with the recent discovery of C. car- natus in the middle Zambezi River (Mutizwa et al. 2024) and the occurrence of several lineages (e.g., Chiloglanis sp. “Pungwe”, Chiloglanis sp. “Zambezi”, Chiloglanis sp. “Nyangombe” and Chiloglanis sp. “Shire”) highlights this region as a potentially overlooked biodiversity hotspot of rheophilic taxa. This is con- sistent with the presence of several recently discovered rheophilic species in other genera such as Anoplopterus marshalli, A. leopardus, Heteromormyrus sp. “Pungwe’, Heteromormyrus sp. “Buzi”, and Zaireichthys monomotapa that are endemic to the Buzi and Pungwe river systems as well as the tributaries of the Lower Zambezi that drain the EZH ecoregion (Eccles et al. 2011; Mazungula and Chakona 2021; Mutizwa et al. 2021). Preliminary examination of specimens of the undescribed Chiloglanis lineages (e.g., Chiloglanis sp. “Zambezi”, Chiloglanis sp. “Nyangombe”, and Chiloglanis sp. “Shire”) show some consistent characters that distinguish them suggesting that they are distinct species yet to be described. In the present study, we refrained from making taxonomic decisions for these lin- eages because we require additional specimens, particularly mature spec- imens for a thorough morphological examination. Chiloglanis asperocutis morphologically resembles the sympatric Chiloglanis sp. “Pungwe’” lineage. The most conservative decision would be to include Chiloglanis sp. “Pun- gwe’ under C. asperocutis, but these two taxa are genetically divergent. De- tailed morphological examination of a larger sample size of mature speci- mens, as well as additional evaluations, including, for example CT scan or cleared and stained samples may identify diagnostic characters that reli- ably distinguish these two taxa. The discovery of high levels of previously unrecognised diversity of suck- ermouth catfishes with largely endemic distribution patterns, for example C. compactus and C. asperocutis endemic to the Pungwe and Buzi rivers, has created renewed interest in the biogeography and evolutionary history of the freshwater fishes of this region. The genus Chiloglanis is inferred to have originated in the Congo Basin in the mid-Eocene around 47.53 Ma (95% HPD: 44.65-50.8 Ma) and its current wide distribution across the Af- rican continent indicates a complex biogeographic history (Day et al 2023). Available evidence from ancestral range reconstruction for Chiloglanis sug- gests that the clade comprising taxa from the Zambezi, East African and Congo Basin ichthyofaunal provinces (Roberts 1975) underwent diversifica- tion as a result of multiple dispersal and recolonization events among these ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 283 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands regions (Day et al. 2023). This complex biogeographic history is reflected in the close phylogenetic relationships of most southern African species such as C. compactus, C. asperocutis, C. carnatus, C. anoterus, C. pretoriae, C. bifurcus, and C. emarginatus with species and lineages from the Zambezi River, whilst a few species such as C. paratus and C. swierstrai have closer ties to species from the Congo Basin (Day et al. 2023). The polyphyletic re- lationship between the sympatrically distributed C. compactus and C. asper- ocutis also suggests complex speciation process within the EZH potentially influenced by dispersal between the Zambezi and the Pungwe and Buzi riv- ers. These hypotheses need to be tested using comparative biogeograph- ic approaches, particularly integrating phylogenomics to more accurately reconstruct the evolutionary relationships and biogeographic patterns of these species. To achieve this, the resolution of the taxonomic uncertainties and more accurate mapping of species distribution ranges is a fundamental requirement. This study provided evidence of the occurrence of two new species in the EZH, highlighting additional taxonomic inconsistencies and paving the way for future studies. The new species described in this study are restricted to rivers systems (Pungwe and Buzi rivers) that currently face multiple threats from anthropo- genic activities. Rich deposits of precious minerals including diamonds and gold support commercial and artisanal mining activities in the EZH. Alluvial gold extraction, in particular, is carried out mainly along rivers and small streams leading to the destruction of the riparian vegetation, increased sed- iments loads, altered flow regimes and often mercury contamination (Ndun- guru et al. 2006; Clark et al. 2019). Exacerbating the situation is the need for better enforcement of regulations to enable sustainable use of the mineral resources in the EZH (Ndunguru et al. 2006; Spiegel 2015; Clark et al. 2019; Kachena and Spiegel 2019). Additionally, climate change, large scale mono- culture plantations and riverbank cultivation often alter physiochemical pa- rameters of river systems negatively impacting aquatic biodiversity in the EZH (Clark et al. 2017, 2019; Mafuwe and Moyo 2020). Furthermore, rainbow trout Oncorhynchus mykiss (Walbaum 1792) was introduced into the rivers of the EZH for angling and this species negatively influences both the distri- bution and abundance of native fish and macro-invertebrate species in the rivers of this region (Kadye et al. 2013). Many of the threats to the environ- ment in the EZH arise due to socio-economic activities, however, there is need to for balancing them with conservation needs in order to ensure long term sustainability of the unique fauna, flora and landscapes that play an important part in supporting the communities in this region. The EZH is a recognised area of high plant diversity and endemism (van Wyk and Smith 2001; Clark et al. 2017). The present and previous studies have uncovered previously overlooked diversity of endemic rheophilic fish- es at a time when critical instream habitats are being degraded by multiple human impacts. This places responsibility on conservation authorities to implement measures to reduce or halt the ongoing impacts on these frag- ile ecosystems. In addition to its unique biodiversity, this region provides ecosystems services of national importance to both Mozambique and Zim- babwe that include water production, carbon sequestration, tourism, non-for- est timber products, commercial forestry and highland agriculture, historical, ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 284 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands cultural and spiritual value (Clark et al. 2019). Although the benefits from this region are well establish, to date there has been few studies that rig- orously quantify the status of ecosystems in this region and the benefits they provide. To address this, the description of species unique to this re- gion and mapping of their distribution provides a base line for assessing the health of the ecosystems of the EZH. For example, due to their dependence on perennial rheophilic habitats, species from the genus Chiloglanis along with other habitat specific species (e.g., aquatic invertebrates) have been used as biological indicators to study habitat integrity and instream flow re- quirements (Kleynhans 1999; Kadye and Moyo 2008; Rashleigh et al. 2009; Soko and Gyedu-Ababio 2015; Achieng et al. 2021). There is need for raising awareness on the uniqueness of the EZH riverine habitats and their endem- ic biota with emphasis on promoting transdisciplinary approaches involving representatives from local communities, government, industry, academia, NGOs, and conservationists to identify effective strategies to balance so- cio-economic development with sustainable management and protection of the unique biodiversity of this previously overlooked freshwater ecoregion. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement Ethical clearance for the approaches used for sample collection and processing was ap- proved by the National Research Foundation-South African Institute for Aquatic Biodiversi- ty (NRF-SAIAB) Animal Ethics Committee (Ref#: 2014/03 and REF#: 25/4/1/7/5_2022-02). Funding Funding for field surveys and genetic analyses was provided by the National Research Foundation-Foundational Biodiversity Information Program for the REFRESH project (FBIP-211006643719) and the Topotypes project (IBIP-BS 13100251309). Author contributions Conceptualization: TB, AC, WTK. Data curation: TIM, WTK. Formal analysis: TIM. Fund- ing acquisition: TB, WTK, AC. Investigation: TIM. Methodology: TIM. Project administra- tion: AC, TB. Visualization: TIM. Writing - original draft: AC. Writing - review and editing: TIM, AC, WTK, PHNB. Author ORCIDs Tadiwa I. Mutizwa © https://orcid.org/0000-0003-4017-1720 Taurai Bere ® https://orcid.org/0000-0002-8603-5137 Wilbert T. Kadye © https://orcid.org/0000-0002-5273-8360 Pedro H. N. Braganc¢a © https://orcid.org/0000-0002-8357-7010 Albert Chakona © https://orcid.org/0000-0001-6844-7501 Data availability All of the data that support the findings of this study are available in the main text or Supplementary Information. ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 285 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands References Achieng AO, Masese FO, Coffey TJ, Raburu PO, Agembe SW, Febria CM, Kaunda-Arara B (2021) Assessment of the ecological health of Afrotropical rivers using fish assem- blages: A case study of selected rivers in the Lake Victoria Basin, Kenya. Frontiers in Water 2(02): 620704. https://doi.org/10.3389/frwa.2020.620704 Alfaro ME, Holder MT (2006) The posterior and the prior in Bayesian phylogenetics. 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Nucleic Acids Research 22(20): 4354-4355. https://doi.org/10.1093/nar/22.20.4354 ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 289 Tadiwa |. Mutizwa et al.: Two new species of Chiloglanis from the Eastern Zimbabwe Highlands Supplementary material 1 Additional material of new species Authors: Tadiwa |. Mutizwa, Taurai Bere, Wilbert T. Kadye, Pedro H. N. Bragan¢a, Albert Chakona Data type: docx Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1241.13891 7.suppl1 ZooKeys 1241: 261-290 (2025), DOI: 10.3897/zookeys.1241.138917 290