Research Article |
Corresponding author: John Sullivan ( jpsullivan@cornell.edu ) Academic editor: Nina Bogutskaya
© 2016 John Sullivan, Sebastien Lavoue, Carl D. Hopkins.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sullivan JP, Lavoué S, Hopkins CD (2016) Cryptomyrus: a new genus of Mormyridae (Teleostei, Osteoglossomorpha) with two new species from Gabon, West-Central Africa. ZooKeys 561: 117-150. https://doi.org/10.3897/zookeys.561.7137
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We use mitochondrial and nuclear sequence data to show that three weakly electric mormyrid fish specimens collected at three widely separated localities in Gabon, Africa over a 13-year period represent an unrecognized lineage within the subfamily Mormyrinae and determine its phylogenetic position with respect to other taxa. We describe these three specimens as a new genus containing two new species. Cryptomyrus, new genus, is readily distinguished from all other mormyrid genera by a combination of features of squamation, morphometrics, and dental attributes. Cryptomyrus ogoouensis, new species, is differentiated from its single congener, Cryptomyrus ona, new species, by the possession of an anal-fin origin located well in advance of the dorsal fin, a narrow caudal peduncle and caudal-fin lobes nearly as long as the peduncle. In C. ona, the anal-fin origin is located only slightly in advance of the dorsal fin, the caudal peduncle is deep and the caudal-fin lobes considerably shorter than the peduncle. Continued discovery of new taxa within the “Lower Guinea Clade” of Mormyridae highlights the incompleteness of our knowledge of fish diversity in West-Central Africa. We present a revised key to the mormyrid genera of Lower Guinea.
Nous avons démontré avec des marqueurs mitochondriaux et nucléaires que trois spécimens de poissons mormyridés faiblement électriques, collectés à trois localités au Gabon sur une période de 13 ans, forment une lignée inconnue au sein de la sous famille des Mormyrinae dont nous avons déterminé la position phylogénétique. Nous décrivons un nouveau genre et deux nouvelles espèces pour ces trois spécimens. Cryptomyrus, nouveau genre, se distingue des autres genres de mormyridés par une combinaison de caractères de l’écaillure, morphométriques, et dentaires. Cryptomyrus ogoouensis, nouvelle espèce, se distingue de sa seule espèce congénérique par sa nageoire anale qui commence nettement en avant de sa nageoire dorsale, son pédoncule caudal gracile et les lobes de sa nageoire caudale pratiquement aussi longs que son pédoncule caudal. Chez Cryptomyrus ona, nouvelle espèce, la nageoire anale commence légèrement en avant de l’origine de la nageoire dorsale, le pédoncule caudal est large, et les lobes de la nageoire caudale sont plus courts que le pédoncule caudal. La découverte ininterrompue de nouveaux taxa dans le «clade de basse Guinée» des Mormyridae illustre notre connaissance incomplète de la diversité des poissons de l’ouest de l’Afrique centrale. Nous présentons une clé révisée des genres de Mormyridae de basse Guinée.
Weakly electric fish, Mormyrinae , integrative taxonomy, phylogeny, DNA, electric organ discharge, EOD, African freshwater fishes, rarity
Mormyrids are nocturnally active fishes endemic to the continental freshwaters of Africa that produce weak electric impulses from a muscle-derived organ located in the caudal peduncle, anterior to the caudal fin. Using specialized electroreceptors distributed over the skin, mormyrids sense nearby objects and prey organisms as distortions to their self-produced electric field (
Here we describe two new species and a new genus of Mormyrinae, based on only three specimens collected over a period of 13 years at three widely separated localities in Gabon, West Central Africa. Despite significant fish collection effort at a number of sites in Gabon since the late 1990s often specifically targeting mormyrid fishes, we know of no other specimens belonging to this unrecognized mormyrid lineage in museum collections. The fishing effort required to produce these three individuals suggests that these species may be extremely rare in nature or that their precise habitat has yet to be discovered. In either case, we have little confidence that additional material will become available soon and believe description of these taxa should not be further postponed.
In July 2001 the first specimen (
Geographic location of collection sites for the three mormyrid specimens treated in this study. Red star = type locality of Cryptomyrus ogoouensis sp. n. at Doumé, Ogooué River; green star = type locality of Cryptomyrus ona sp. n. at the Moukalaba River; green circle = locality of second specimen of C. ona at the Mabounié River.
Photographs of the collection localities of the three mormyrid specimens treated in this study. A Doumé falls on the Ogooué River, Ogooué-Lolo, Gabon, type locality of Cryptomyrus ogoouensis sp. n. during high water in May 2011 B same locality in low water, September 2014 C Nyanga River at confluence with Moukalaba River near collection site of holotype of Cryptomyrus ona sp. n., July 2001 D Collection site of Cryptomyrus ona specimen
In 2012 a second specimen resembling the Moukalaba River fish was collected in a gill net sample from the Mabounié River, a small, right-bank tributary of the lower Ngounié River (Ogooué basin, Ngounié Province; Figs
Finally, in September 2014 during the course of an ichthyofaunal survey sponsored by The Nature Conservancy in the “Rapids of Mboungou Badouma and Doumé” Ramsar site located along the Ogooué River upstream of Lastoursville (Ogooué-Lolo Province), a single live specimen somewhat resembling the two other fish was collected in an earthworm-baited fish trap at Doumé falls, just beside the village of Doumé (Figs
As detailed above, the three specimens described here were collected on three separate expeditions to Gabon in 2001, 2012 and 2014. Specimens of other species used in this study are listed below in the comparative material examined section. Institutional abbreviations follow
Specimens were collected, handled and euthanized in accordance with guidelines published by the
In order to investigate the phylogenetic relationships of the three novel mormyrid specimens, we sequenced from each the complete cytochrome b (cyt-b) gene and portions of the 12S and 16S genes from the mitochondrial genome, as well as partial rag2 and the complete S7 introns 1 & 2 and the short intervening exon from the nuclear genome. We added these sequences to an existing 4209 bp alignment of these markers from 38 species belonging to 17 mormyrin genera published in
Because the possible relationship of these new taxa to the genus Hippopotamyrus Pappenheim, 1906 was of special interest, we also sequenced all of these markers from a specimen of Hippopotamyrus castor from Cameroon, the type species of this genus, a species not included in
DNA was extracted from fin clips or epaxial muscle tissue preserved in 95% ethanol using a QIAGEN (QIAGEN, Inc., Valencia, CA) DNeasy kit. Primer sequences for cyt-b, 12S, 16S, rag2, S7 and COI are listed in Table
Forward and reverse primers used to amplify six genetic markers used in this study shown in 5’ to 3’ orientation.
cytbF-L14724 | GAC TTG AAA AAC CAC CGT TG |
cytbR-H15915 | CTC CGA TCT CCG GAT TAC AAG AC |
COIF-ZPeng | TCT CAA CCA ACC ATA AAG ACA TTG G |
COIR-ZPeng | TAT ACT TCT GGG TGC CCA AAG AAT CA |
12S-L1067 | AAA CTG GGA TTA GAT ACC CCA CTA T |
12S-H1478 | GAG GGT GAC GGG CGG GCG GTG TGT |
16S-L2510 | CGC CTG TTT ATC AAA AAC AT |
16S-H3080 | CCG GTC TGA ACT CAG ATC ACG T |
rag2F2 | ArA CGC TCm TGT CCm ACT GG |
rag2R6 | TGr TCC ArG CAG AAG TAC TTG |
S7RPEX1F | TGG CCT CTT CCT TGG CCG TC |
S7RPEX3Ralt | ACC TTT GCT GCA GTG ATG TT |
Sequences were edited and combined into contigs for each fragment with Sequencher 4.2 (GeneCodes Corporation, Ann Arbor, MI). Requiring no gaps, alignment of the new coding gene sequences (cyt-b and rag2) to the
We performed a maximum likelihood phylogenetic analysis of the matrix from
Because the three specimens under study do not obviously belong to any one of the 19 described mormyrin genera, choosing mormyrin species for comparison was not straightforward. We collected counts and measurements from seven sympatric taxa with which the new taxa could conceivably be confused in the field and two extralimital taxa with (in our estimation) the most overall resemblance to the three specimens, both of which are type species of genera not known to occur in Gabon. The sympatric taxa used for comparison are Ivindomyrus marchei (Sauvage, 1879), I. opdenboschi Taverne & Géry, 1975, Marcusenius moorii (Günther, 1867), Stomatorhinus walkeri (Günther, 1867), Paramormyrops kingsleyae (Günther, 1896) and two undescribed species of Paramormyrops Taverne, Thys van den Audenaerde & Heymer, 1977. The extralimital taxa used for comparison are Hippopotamyrus castor (from coastal drainages of Cameroon) and Cyphomyrus psittacus (Boulenger, 1897) from the Congo basin. Most of these species are illustrated in Figure
Some mormyrid species used for morphological comparison in this study. A Ivindomyrus marchei female, 135 mm SL,
We took 29 point-to-point measurements with a digital caliper on each of the three specimens under study for comparison to those taken from 115 individuals of nine species listed above. Twenty-seven of these measurements are defined in
To facilitate dorsal- and anal-fin ray comparisons among individuals, we report the number of unbranched rays (3) + number of branched rays. (The first two rays are small, the first often only visible in radiographs.)
Specimens were radiographed using a Faxitron Model LX-60 cabinet x-ray system and Kodak Industrex MX125 film. Film images were scanned on a flatbed scanner.
We follow
Sex of the specimens was determined by presence/absence of an “anal-fin notch”: a dorsally directed indentation along the anterior base of the anal fin present in all mature male mormyrids (
The EOD waveform of the specimen from Doumé is triphasic and very brief (Fig.
A Three superimposed electric organ discharge (EOD) waveform recordings of Cryptomyrus ogoouensis holotype
GenBank numbers for the new sequences generated for this study and their appropriate GenSeq codes (
GenBank numbers and specimen information for DNA sequences generated in this study. All but COI were added to alignment of
Specimen | Catalog no. | COI | cyt-b | 12S | 16S | rag2 | S7 | GenSeq Status |
---|---|---|---|---|---|---|---|---|
Cryptomyrus ogoouensis holotype |
|
KT875221 | KT875226 | KT875213 | KT875217 | KT875230 | KT875233 | genseq-1 COI, cyt-b, 12S, 16S, rag2, S7 |
Cryptomyrus ona holotype |
|
KT875222 | KT875227 | KT875214 | KT875218 | KT875231 | KT875235 | genseq-1 COI, cyt-b, 12S, 16S, rag2, S7 |
Cryptomyrus ona Mabounié specimen |
|
KT875223 | KT875228 | KT875215 | KT875219 | KT875232 | KT875234 | genseq-3 COI, cyt-b, 12S, 16S, rag2, S7 |
Hippopotamyrus castor |
|
KT875220 | KT875224 | KT875212 | KT875216 | KT875229 | KT875236 | genseq-3 COI, cyt-b, 12S, 16S, rag2, S7 |
Hippopotamyrus pictus (White Nile) |
|
KT875225 | genseq-4 cyt-b |
Genetic distances (uncorrected p-distances) between three Cryptomyrus specimens and nearest relatives Boulengeromyrus knoepffleri and Ivindomyrus marchei for mitochondrial and nuclear markers sequenced, shown in order as COI/cyt-b/combined 12S-16S/rag2/S7. Boulengeromyrus knoepffleri
1 | Boulengeromyrus knoepffleri | 1 | 2 | 3 | 4 | 5 |
2 | Ivindomyrus marchei | 0.033/0.051/0.014/0.004/0.011 | • | |||
3 | Cryptomyrus ogoouensis holotype | 0.085/0.065/0.028/0.022/0.021 | 0.083/0.064/0.025/0.017/0.015 | • | ||
4 | Cryptomyrus ona holotype | 0.085/0.063/0.028/0.021/0.021 | 0.080/0.064/0.028/0.017/0.015 | 0.006/0.013/0.006/0.0/0.0 | • | |
5 | Cryptomyrus ona Mabounié specimen | 0.083/0.061/0.027/0.021/0.021 | 0.079/0.064/0.026/0.017/0.015 | 0.008/0.015/0.005/0.0/0.0 | 0.002/0.005/0.001/0.0/0.0 | • |
We note that genetic differentiation observed between the Doumé specimen and the Moukalaba and Mabounié specimens, while small, is within the range seen between some closely related, but distinct mormyrid species. For example cyt-b sequences from the morphologically divergent Paramormyrops gabonensis Taverne, Thys van den Audenaerde & Heymer, 1977 and P. hopkinsi Taverne & Thys van den Audenaerde, 1985 – two species for which there is no evidence of mitochondrial introgression – similarly differ by 1.3%. However, in other cases, populations from different river basins regarded as conspecific can differ by this much. For example, Ivindomyrus marchei from the Ogooué and Ivindo Rivers differ from those in the Nyanga River by just slightly less than 1.3% (
Among the more than 300 mormyrid COI sequences in the Barcode of Life Database (BOLD;
Turning to the nuclear markers, the near identity of the nuclear S7 intron sequences among the Mabounié, Doumé and Moukalaba specimens might be thought surprising if these three specimens represent more than one species (less surprising for the identity of the rag2 sequences, as this marker evolves more slowly). However, distinct mormyrid species having identical or near-identical nuclear S7 intron sequences is not unprecedented, as similarly identical S7 intron sequences were found among three species of Campylomormyrus by
The aligned dataset and tree described here have been archived on TreeBASE and can be accessed at http://purl.org/phylo/treebase/phylows/study/TB2:S18468?format=html.
In the maximum likelihood tree produced in RAxML (Fig.
Maximum likelihood phylogenetic tree calculated in RAxML for 46 mormyrin specimens based on DNA sequences from mitochondrial cytochrome b, 12S, 16S and nuclear rag2 and S7 intron, rooted with Myomyrus macrops. Bootstrap support values at nodes. Some clades collapsed for simplicity. Position of new taxa (magenta) within the “Lower Guinea Clade” (green) and distance from Hippopotamyrus castor, type species of genus Hippopotamyrus (red), highlighted.
To explore the relative contribution of mitochondrial and nuclear data to the phylogenetic result, we ran two additional analyses, one with only mitochondrial data included for the three specimens and the other with only nuclear data included for them. In each case, excluded bases were recoded as missing data, while the full dataset was used for all other specimens. In the nuclear data-only analysis, we used a single OTU for the three specimens since rag2 and S7 sequences were identical among them. In both experimental analyses, the lineage of the three new taxa was resolved as sister to Boulengeromyrus plus Ivindomyrus with a 100% bootstrap proportion, indicating strong signal contributing to this result from both the nuclear and mitochondrial data partitions.
Consistent with the genetic distances reported above, measures and counts of the Moukalaba and Mabounié specimens are more similar to each other than they are to those of the Doumé specimen. In particular, dorsal- and anal-fin bases are short in the former two, with low fin-ray counts, 20/25 and 21/24 total rays respectively, compared to 24/30 in the Doumé specimen, in which the anal-fin origin is situated much further in advance of the dorsal-fin origin. Predorsal distance is nearly equal to preanal distance in the Moukalaba and Mabounié specimens, while it is markedly shorter in the Doumé specimen due to its longer anal fin. Additionally, the caudal peduncle is much deeper and the caudal-fin lobes are shorter in the Moukalaba and Mabounié specimens than in the Doumé specimen. We have never observed a range of four dorsal-fin rays, six anal-fin rays, and differences of 25 percent in anal-fin length relative to standard length among individuals of a single mormyrid species.
While our initial hypothesis that the new taxa are close relatives of Hippopotamyrus castor from Cameroon was ruled out by the molecular result, substantial morphological differences are also obvious between the new taxa and H. castor. The new taxa have fewer scales (44–45 lateral line scales/12 around the caudal peduncle vs. 72–81/16 in H. castor), fewer total vertebrae (40–43 vs. 47), fewer dorsal-fin rays (20–24 vs. 31–33), and a longer anal-fin base (dorsal 78–88% of anal-fin base vs. nearly equal), among other significant differences. Also, while H. castor has elongate paired incisor teeth in the lower jaw (from which its specific epithet derives), their morphology is quite different from the lower teeth of Cryptomyrus. In the three specimens under study, the central incisors are flattened and spatulate. The neighboring pair of teeth are smaller, but similarly spatulate and close or appressed to the inner pair which assume a sort of trowel shape, pointing outward (Fig.
Mouth and dentary teeth in A Cryptomyrus ogoouensis holotype
With its moderately swollen chin and small mouth, the head of Cyphomyrus psittacus appears somewhat similar to those of the new taxa and like them C. psittacus has a diffuse band of dark pigment between the origin of the dorsal and anal fins, but the new taxa differ from C. psittacus in being less deep bodied (BD 21–24% SL vs. 30–34%), having a shallower head (HD close to 79% HL vs. 63–75%), a smaller eye (ED 19–20% HL vs. 24–26%), a dorsal fin somewhat shorter than the anal fin (vs. far longer than the anal fin) and fewer midlateral scales (44–45 vs. 54–56). Unlike the spatulate dentary teeth of the new taxa, dentary teeth in species of Cyphomyrus examined are very small, feebly notched, and embedded in the gums.
The three specimens also differ morphologically from individuals of the two species of Ivindomyrus to which the phylogenetic study shows they are closely related. Compared to Ivindomyrus, all three specimens have shorter, deeper caudal peduncles (CPD 31–35% CPL vs. 21–28%), greater interorbital width compared to head length (IOW 33–35% HL vs. 21–30%), a higher dorsal-fin to anal-fin length ratio (DFL 78–83% AFL vs. 65–76%), and considerably fewer midlateral scales (44–45 vs. 53–58). Like the dentary teeth in the new taxa, those in Ivindomyrus are spatulate, however the central pair of incisors are not noticeably elongate along their inner margins as they are in the new taxa (Fig.
Compared to species of Paramormyrops the new taxa are generally deeper bodied (BD 21–25% SL vs. 15–21%) with deeper heads (HD close to 79% HL vs. 63–75%), larger eyes (ED 19% HL vs. 9–17%) and shorter postorbital head length (POL 59–60% HL vs. 61–70%) in addition to having far fewer midlateral scales (44–45 vs. 56–71). Dentary teeth in all Paramormyrops are bicuspid, not spatulate (Fig.
Compared to Marcusenius moorii, a species with a similarly low midlateral line scale count, the new taxa most notably lack a chin swelling that protrudes forward beyond the tip of the snout, have a much more subdued and diffuse area of pigment between the dorsal-fin and anal-fin origins, a larger eye (ED 19–20% HL vs. 15–18%) and unicuspid (vs. bicuspid or notched) dentary teeth.
Two of the three new taxa are of larger size than the largest Stomatorhinus walkeri examined and all three have a smaller head relative to standard length (HL 22–23% SL vs. 28–31%) a larger eye (ED 19% HL vs. 9–13%), a modest chin swelling (vs. none), and the posterior nares are remote from the rictus of the mouth (vs. in close proximity).
Based on the molecular phylogenetic analysis, we conclude that these three morphologically distinctive specimens represent an undescribed lineage of Mormyrinae sister to the clade formed by Boulengeromyrus plus Ivindomyrus. This phylogenetic finding in combination with the morphological differences that exist between these new taxa and Boulengeromyrus and Ivindomyrus precludes placing them in either genus. Creation of an enlarged Boulengeromyrus Taverne & Géry via synonymy in which to subsume all of these taxa is ruled out by the significant morphological differences between them and lack of known morphological synapomorphies. Hence a new genus is required.
We recognize two species: one for the Doumé specimen and the other comprising the Moukalaba and Mabounié specimens, reflecting the greater morphological and genetic similarity of these latter two and their notable morphological differences from the Doumé specimen, discussed above. We choose the Moukalaba River specimen as type for the second species and treat the Mabounié River specimen as a non-type. We describe these new taxa below.
Cryptomyrus ogoouensis sp. n.
Cryptomyrus ogoouensis sp. n., Cryptomyrus ona sp. n.
Cryptomyrus gen. n. is distinguished from all other mormyrid genera by combination of the following features. Scales large: 44 or 45 along the midlateral line, with about 42 pierced lateral line scales; mouth subinferior; broad but nonprotrusive chin swelling that does not extend beyond snout; snout expansive and rounded in lateral profile with slight inflection point visible above anterior margin of eye in lateral view; snout somewhat v-shaped in dorsal view; eye large, 19–20% HL; middle four teeth on dentary squarish, broad and spatulate, oriented nearly horizontally, central two lower teeth longest along inner edges and in contact with each other, jutting in advance of neighboring pair and forming a trowel-like shape (Fig.
Very few Mormyrinae have so few midlateral scales. Only some of the large-scaled Marcusenius such as Marcusenius moorii (
Gender masculine; from the Greek kryptos meaning secret or hidden referring to the rarity of these fishes in collections and the Greek myros, a kind of fish, a suffix used in the names of many other mormyrid genera.
Cryptomyrus ogoouensis sp. n. is readily differentiated from its sole congener, C. ona sp. n., in the possession of an anal-fin origin located well in advance of the dorsal fin (first dorsal ray above anal-fin ray 7 vs. first dorsal ray above anal-fin ray 3), a narrow caudal peduncle (depth 5.1% SL vs. 6.0–6.8% SL), and lobes of caudal fin nearly as long as caudal peduncle (vs. markedly shorter).
Morphometric and meristic data for holotype (female, 111.6 mm SL) presented in Table
Descriptive morphometrics and meristics. Data shown for holotype of Cryptomyrus ogoouensis sp. n. (
Character |
Cryptomyrus ogoouensis sp. n. Holotype Ogooué River at Doumé |
Cryptomyrus ona sp. n. Holotype Moukalaba River |
Cryptomyrus ona sp. n. Non-type Mabounié River |
---|---|---|---|
Standard length (SL), mm | 111.6 | 107.8 | 98.1 |
Head length (HL), mm | 24.3 | 24.3 | 22.6 |
Percent of SL | |||
Body depth at pelvic fin (BD) | 23.4 | 24.2 | 20.8 |
Head length (membrane) (HL) | 21.8 | 22.6 | 23.0 |
Head length (opercle bone) (HLBO) | 18.1 | 20.3 | 19.6 |
Predorsal distance (PDD) | 62.7 | 62.4 | 63.1 |
Preanal distance (PAD) | 55.0 | 62.3 | 58.7 |
Prepelvic distance (PPLD) | 35.0 | 40.2 | 35.7 |
Prepectoral distance (PPCD) | 21.5 | 24.5 | 21.8 |
Caudal peduncle depth (CPD) | 5.1 | 6.8 | 6.6 |
Caudal peduncle length (CPL) | 16.4 | 18.0 | 18.6 |
Head width (HW) | 10.1 | 10.7 | 11.0 |
Pectoral-fin length (PCFL) | 18.8 | 18.5 | 17.8 |
Pelvic-fin length (PLFL) | 11.6 | 10.9 | 12.7 |
Pelvic-anal-fin distance (DPLAF) | 19.8 | 22.9 | 22.3 |
Pectoral-anal-fin distance (DPCAF) | 33.9 | 37.8 | 37.3 |
Anal-fin base length (AFL) | 29.5 | 22.1 | 24.3 |
Dorsal-fin base length (DFL) | 23.2 | 19.4 | 20.0 |
Percent of HL | |||
Snout length (SNL) | 26.6 | 26.3 | 23.1 |
Post-orbital length (POL) | 59.0 | 59.6 | 59.5 |
Head width (HW) | 46.1 | 47.5 | 47.7 |
Interorbital width (IOW) | 33.2 | 31.9 | 34.5 |
Head depth (HD) | 79.4 | 78.9 | 78.7 |
Eye diameter (ED) | 19.5 | 19.9 | 19.0 |
Inter-nostril distance (DNN) | 4.7 | 4.2 | 5.4 |
Nares-eye distance (DNE) | 7.2 | 5.9 | 3.1 |
Mouth width (MW) | 14.8 | 15.8 | 11.8 |
Ratios and angles | |||
Inter-orbital width as % head width | 72.0 | 67.2 | 72.3 |
Pre-anal distance/pre-dorsal distance | 87.8 | 100.0 | 93.0 |
Inter-nostril width as % interorbital width | 14.2 | 13.1 | 15.6 |
Caudal peduncle depth as % CP length | 31.2 | 37.7 | 35.3 |
Length of dorsal/length of anal | 78.5 | 87.7 | 82.5 |
Counts | |||
Dorsal rays (simple+branched) | 3+21=24 | 3+18=21 | 3+17=20 |
Anal rays (simple +branched) | 3+28=31 | 3+21=24 | 3+22=25 |
Anal rays before dorsal | 7 | 2 | 4 |
Anal rays beyond last dorsal ray | 3 | 2 | 2 |
Pectoral rays | 10 | 10 | 10 |
Pelvic rays | 6 | 6 | 6 |
Total midlateral scales (pierced+unpierced) | 43+2=45 | 42+2=44 | 42+2=44 |
Rows scales above lateral line to dorsal | 9 | 9 | 7 |
Rows scales below lateral line to pelvic | 9 | 9 | 10 |
Caudal peduncular scales | 12 | 12 | 12 |
Teeth upper jaw/lower jaw | 5/6 | 5/6 | 5/6 |
Total vertebrae | 43 | 41 | 40 |
Hypurals 1 & 2 | unfused | unfused | unfused |
In life, a light cinnamon brown along sides, darker along dorsum, reflective coppery pigment on lower half of head and along belly to anal fin, mouth and chin whitish. Eye with golden iris, dark center. Faint, diffuse broad band of pigment between dorsal and anal fin occupying 4–5 scales, darker above. Numerous depigmented spots over electroreceptors conspicuous on snout, top of head, belly and upper back. Fins with very lightly pigmented rays, interradial membranes hyaline. In preservation, light brown.
EOD waveform recorded from the holotype specimen is very short, about 0.55 millisecond total duration (recorded at 23.2 °C) if measured from the onset of the first very weak head-negative phase (P0) and only 0.28 milliseconds if measured from the rising phase of the head-positive phase (P1) to the end of the large head negative phase (P2) using a 2% departure-from-baseline threshold (Fig.
The specific epithet is a Latinized noun in the genitive case and refers to the Ogooué River of Gabon.
Currently known only from the Ogooué River at Doumé falls. At the collection site, we recorded a water temperature of 26.7 °C, a pH of 6.89, water conductivity of 13.8 μS/cm and dissolved oxygen of 84.7%.
Cryptomyrus ona sp. n. is readily differentiated from its sole congener, C. ogoouensis sp. n., in having an anal-fin origin located only just in advance of the dorsal fin (first dorsal ray above anal-fin ray 3 vs. first dorsal-fin ray above anal-fin ray 7), a deep caudal peduncle (6.0–6.8% SL vs. 5.1% SL), and lobes of caudal fin markedly shorter than caudal peduncle (vs. nearly as long as peduncle).
Morphometric and meristic data for holotype (male, 107.8 mm SL) presented in Table
Photo of holotype specimen recovered from gill net, several hours post-mortem, shows a purplish-gray body, darker along dorsum, with whitish marbling/speckling on lower half of head and along belly to anal fin and diffuse band of pigment below dorsal fin occupying 4 scales, darker above, lighter or absent above anal fin. Numerous depigmented spots over electroreceptors conspicuous on snout, top of head, belly and upper back. Fins with darkly pigmented rays, interradial membranes hyaline. In preservation, body yellowish tan.
Unknown.
The specific epithet is a noun in apposition that honors Marc Ona Essangui, Gabonese environmental and civic activist, founder and executive director of the NGO Brainforest and recipient of the 2009 Goldman Environmental Prize, in appreciation for his efforts to protect Gabon’s equatorial forests and wetlands.
Known from the type locality in the Moukalaba River at its confluence with the Nyanga River. A second specimen referred to this species comes from the Mabounié River, a small right-bank affluent of the lower Ngounié River, below Samba Falls. Both specimens were caught by gill net at night. At the type locality we recorded water temperature of 23.6 °C, pH of 8.0, water conductivity of 116.0 μS/cm and dissolved oxygen of 4.98 mg/l. Water conductivity at the Mabounié River at time of collection of that specimen was 48 μS/cm.
Lower Guinea is defined as the Atlantic drainages of Africa from the Cross River of Cameroon in the north to the Chiloango River of Cabinda/D.R. Congo in the south (
1 | Nostrils close to one another and to the eye; mouth inferior, below the level of the eye; body short and rather deep | genus Petrocephalus (subfamily Petrocephalinae; widespread) |
– | Nostrils separated from each other and from the eye; mouth terminal or inferior, in advance of the eye, body deep or elongate | (subfamily Mormyrinae) |
2 | Teeth extending along the entire edge of both jaws in a single series, 10–36 in each jaw; mouth terminal, well in advance the eye; body elongate, depth more than 5.2 times into SL | Mormyrops (widespread) |
– | Teeth restricted to middle of each jaw, 3–10 in each jaw | 3 |
3 | Dorsal fin more than twice the length of anal, originating in advance of pelvic fin insertion | Mormyrus (Cross, Sanaga, Nyong) |
– | Dorsal fin 0.35–1.25 times the length of the anal, originating behind pelvic fin insertion | 4 |
4 | Pelvic fin insertion closer to the anal than to the pectoral fins; body very elongate, at least 8–11 times as long as deep | Isichthys (widespread) |
– | Pelvic fin insertion mid-way between anal and pectoral fins or closer to pectorals; body less elongate to short | 5 |
5 | Posterior nostril located close to the border of the mouth | Stomatorhinus (Ogooué, Ivindo, Kouilou-Niari) |
– | Neither nostril close to the border of the mouth | 6 |
6 | Snout elongated and tubular, its length greater than the post-orbital length of the head; snout turned downward | Campylomormyrus (Sanaga) |
– | Snout non-tubular, its length less than the post-orbital length of the head | 7 |
7 | Prominent tapered cylindrical barbel-like appendage under the chin, extending forward from below lower jaw | Gnathonemus (Cross, Sanaga) |
– | Submental appendage reduced to fleshy swelling or absent altogether | 8 |
8 | Submental appendage prominent, extending slightly beyond the end of the upper jaw, mouth terminal | Marcusenius (widespread) |
– | Submental appendage not extending beyond end of upper jaw or absent altogether, mouth terminal or inferior | 9 |
9 | Scales along midlateral line 44 or 45, broad submental swelling, mouth subinferior | Cryptomyrus (Ogooué, Nyanga) |
– | Scales along midlateral line 47 or more, chin swelling only slightly developed or absent; mouth terminal or inferior | 10 |
10 | Dorsal and anal fins approximately equal in length and originating at the same vertical level, dorsal fin with 31–34 rays, anal fin with 31–35 rays, mouth inferior | Hippopotamyrus (Cross, Sanaga, Wouri, Lokoundjé) |
– | Dorsal fin shorter than anal fin and with fewer than 30 rays | 11 |
11 | Body moderately elongate, depth 18–22% SL | 12 |
– | Body moderately deep, more than 23% SL | 13 |
12 | Anal and dorsal fins terminate at about the same level. Distal tips of last anal and dorsal rays not offset | Paramormyrops (widespread) |
– | Anal fin terminates beyond the end of dorsal. Distal tips of last anal and dorsal fin rays offset | Brienomyrus (widespread) |
13 | Mouth terminal | 14 |
– | Mouth subterminal | 15 |
14 | Snout straight, short and blunt, no darkly pigmented transverse band between dorsal and anal fins | Brevimyrus (Cross) |
– | Snout turned downward, long, conical; darkly pigmented transverse band between dorsal and anal fins | Boulengeromyrus (Ntem, Ivindo) |
15 | Posterior nostril closer to anterior nostril than to eye; darkly pigmented transverse band between dorsal and anal fins | Ivindomyrus (Ntem, Ivindo, Ogooué, Nyanga) |
– | Posterior nostril closer to eye than to anterior nostril; no darkly pigmented transverse band between dorsal and anal fins | Pollimyrus (Cross, Wouri, Kouilou-Niari) |
Cryptomyrus is the first new genus of Mormyridae to be described since Paramormyrops, from the same region of Africa, in the late 1970s (
The Moukalaba-Nyanga system of Gabon, the type locality of Cryptomyrus ona, remains understudied and is likely to produce additional taxonomic novelties for Mormyridae and other groups. Doumé, the Ogooué River collection site for C. ogoouensis, is already an important type locality for fishes. From collections made here in 1876–77 by Alfred Marche, naturalist on the first of Pierre Savorgnan de Brazza’s expeditions that explored the sources of the Ogooué (see
Despite its provenance from a part of the greater Ogooué basin, not the Nyanga basin, we treat the Mabounié River specimen as a non-type specimen of Cryptomyrus ona on the basis of its morphological and genetic similarity to the C. ona type specimen. It is worth noting that headwaters of the Ngounié abut those of the Nyanga and that at least two other fish species, Synodontis ngouniensis De Weirdt, Vreven & Fermonm, 2008 and Aphyosemion primigenium Radda & Huber, 1977 appear to have exclusive distribution in these two river basins, having never been collected elsewhere in the wider Ogooué system. Synodontis punu Vreven & Milondo, 2009 is found in these two river basins plus the Kouilou-Niari basin to the south within the Republic of Congo.
The distinctive morphology of these specimens drew our attention to them, but provided few clues about their affinities to other mormyrid species. While we have an EOD waveform recording only from the Doumé specimen, its uniqueness among those known from Gabon’s mormyrids also helped to highlight its special status. Sequence data were necessary for confirming that these three individuals are indeed closest relatives and for placing them phylogenetically within the Mormyrinae. The combination of these datasets– morphology, electric signals and DNA– provides a practical, integrative, and evolutionary framework in which to evaluate the status of candidate mormyrid species. The utility of this approach has been demonstrated in a number of other recent publications on Mormyridae (e.g.
Morphological synapomorphies remain to be discovered for the “Lower Guinea Clade” of Mormyridae to which Cryptomyrus belongs (Fig.
Biodiversity unknown to science is invisible to conservation efforts. The rivers of Gabon, like others across Africa, are increasingly impacted by logging, road-building, mining, and hydropower dam construction (
Cyphomyrus psittacus: D.R. Congo, Orientale, Wagenia Falls,
Hippopotamyrus castor: Cameroon, Centre, Sanaga R. at Nachtigal Falls,
Ivindomyrus marchei: Gabon, Moyen-Ogooué, Ogooué R. at Lambaréné,
Ivindomyrus opdenboschi: Gabon, Ogooué-Ivindo, Ivindo R. at Loa Loa,
Marcusenius moorii: Gabon, Haut-Ogooué, Ogooué R.,
Paramormyrops kingsleyae: Gabon, Moyen-Ogooué, Lambaréné, Mikouma Creek,
Paramormyrops sp. (undescribed species): Gabon, Haut-Ogooué, Okoloville,
Paramormyrops sp. “SN4”: Gabon, Haut-Ogooué, Ogooué R. at Franceville,
Stomatorhinus walkeri: Gabon, Ogooué-Lolo, Ogooué R., Haut-Ogooué, Ogooué R.,
We gratefully acknowledge D.F. Idiata and A.F. Koumba Pambo of CENAREST (Centre National de la Recherche Scientifique et Technologique) for approving the scientific collection of fishes in Gabon and issuing export permits in 2011 and 2014. For administrative assistance as well as help with fieldwork in Gabon, we thank the personnel of IRAF (Institut de Recherches Agronomiques et Forestières) in particular J.D. Mbega, J.-H. Mve Beh and G. Koudaou. V. Mamonekene of Marien Ngouabi University, Brazzaville, Congo assisted with collection at the Moukalaba site during an expedition funded by WWF US (Central Africa Ecoregion Conservation Program). We express our gratitude to Y. Fermon of the