Corresponding author: Albert Chakona (
Academic editor: N. Bogutskaya
Chakona A, Swartz ER, Skelton PH (2014) A new species of redfin (Teleostei, Cyprinidae,
Species of the cyprinid genus
The Verlorenvlei and Berg lineages of
Thus,
Institutional abbreviations follow
Two
Meristic and morphological characters were examined following Hubbs and Lagler (1958),
Morphological characters of
Character | Description | Acronym |
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Standard length | Tip of the snout to the point of flecture of the caudal fin | SL |
Pre-dorsal length | Tip of the snout to the origin of the dorsal fin | PDL |
Head length | Tip of the snout to the posterior bony margin of the operculum | HL |
Snout length | Tip of the snout to the anterior bony edge of the orbit | S |
Orbit diameter | The greatest bony diameter of the orbit | OD |
Inter-orbit width | Shortest distance between bony edges of the orbits | IO |
Post-orbit length | Distance between the posterior bony edge of orbit to the posterior bony edge of operculum | PO |
Head depth | Maximum depth measured from the nape | HD |
Body depth | Maximum depth measured from the anterior base of the dorsal fin | BD |
Anterior barbel length | From base to tip of anterior barbel | AB |
Posterior barbel length | From base to tip of posterior barbel | PB |
Dorsal fin base | Distance between origin of dorsal fin and base of last dorsal fin ray | DB |
Dorsal fin height | From anterior base to tip of dorsal fin | DH |
Pectoral fin length | From anterior base to tip of pectoral fin | PtL |
Pelvic fin length | From anterior base to tip of pelvic fin | PvL |
Anal fin base | Distance between origin of anal fin and base of last anal fin | AfB |
Anal fin height | From anterior base to tip of anal fin | AfH |
Caudal peduncle length | Distance from posterior base of anal fin the point of flecture of the caudal fin | CPL |
Caudal peduncle depth | The least depth of the caudal peduncle | CPD |
Pectoral to pelvic fin length | Distance between the posterior margins of the fin bases | PP |
Pelvic to anal fin length | Distance between posterior base of the pelvic fin to anterior base of the anal fin | PA |
Body width | The greatest width just anterior to the origin of the dorsal fin | BW |
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Lateral line scales | Number of scale rows along the lateral line | LL |
Lateral line to dorsal fin origin | Number of scale rows between lateral line scale row (does not include lateral line scale) and anterior base of the dorsal fin | LD |
Lateral line to pelvic fin origin | Number of scale rows between lateral line scale row (does not include lateral line scale) and anterior base of pelvic fin | LP |
Lateral line to anal fin origin | Number of scale rows between lateral line scale row (does not include lateral line scale) and anterior base of the anal fin | LA |
Circumpeduncular scales | Number of scale rows around the caudal peduncle at narrowest portion of caudal peduncle | CP |
Predorsal scales | Number of scale rows between the supraoccipital and the anterior base (origin) of the dorsal fin | PDS |
Unbranched dorsal fin rays | Number of unbranched primary dorsal rays | UdR |
Branched dorsal fin rays | Number of branched dorsal rays; two last branched rays counted as one | BdR |
Anal fin rays | Includes both simple and branched rays; two last rays counted as one | |
Pectoral fin rays | Includes both simple and branched rays | |
Pelvic fin rays | Includes both simple and branched rays | |
Total vertebrae | Total number of vertebrae in vertebral column (including four Weberian vertebrae and a single ural centrum) | TV |
Pre-dorsal vertebrae | Total number of vertebrae in advance of the leading dorsal fin pterigiophore (including the four Weberian vertebrae) | PdV |
Pre-caudal vertebrae | Total number of vertebrae in advance of the vertebrae with haemal arch opposite the leading anal pterygiophore plus the four Weberian vertebrae | PcV |
Pre-anal vertebrae | Total number of vertebrae in advance of the leading anal pterygiophore (including the four Weberian vertebrae) | PaV |
Caudal vertebrae | Total number of vertebrae posterior to (and including) the vertebra with haemal arch opposite the leading anal pterygiophore plus a single ural centrum | CV |
We compared morphological and meristic differences among all double barbeled redfins using raw data from
Statistical analyses were performed with the programs InfoStat (Di Rienzo et al. 2012), PAST and STATISTICA 12. Prior to analyses, morphometric data were normalised using procedures described by
Principal component analysis (PCA) was performed using the correlation matrix to explore the separation of the specimens based on the normalised morphometric data (
Discriminant Function Analysis (DFA) was performed to visualise the degree of morphological separation among the species and to identify the most important characters that contribute to the differentiation. DFA also provides jacknifed measurements of re-classification success of individuals to their original group, as well as identifying the group to which individuals were assigned if misclassified. Separate DFAs were performed for the morphometric and meristic characters, as well as for these two data sets combined.
Figure
Bayesian phylogenetic tree showing genetic distances between
Principal components analysis (PCA) of normalised morphometric and raw meristic characters shows
Scatter plot of PC1 against PC2 for a PCA carried out on 15 normalised morphometric and seven raw meristic characters for all examined specimens (
Factor loadings for the first three principal component (PC) axes of a PCA carried out on morphometric and meristic characters of double barbeled
Character | PCI | PCII | PCIII |
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Head length | -0.769 | 0.408 | -0.077 |
Head depth | 0.669 | 0.194 | -0.276 |
Inter orbit | 0.514 | 0.297 | -0.421 |
Snout length | 0.205 | -0.727 | 0.040 |
Post orbit | 0.325 | -0.387 | -0.067 |
Predorsal length | -0.721 | 0.418 | 0.218 |
Dorsal fin base | 0.507 | -0.115 | 0.296 |
Body depth | 0.148 | 0.796 | 0.269 |
Body width | -0.463 | 0.153 | 0.620 |
Caudal peduncle length | 0.478 | -0.431 | 0.107 |
Caudal peduncle depth | -0.335 | 0.178 | 0.752 |
Anterior barbel | -0.405 | -0.729 | 0.418 |
Posterior barbel | -0.250 | -0.477 | 0.520 |
Unbranched dorsal fin rays | -0.215 | -0.477 | 0.111 |
Lateral line scales | -0.650 | -0.340 | -0.153 |
Scale rows between lateral line and dorsal fin | -0.853 | 0.024 | -0.100 |
Scale rows between lateral line and pelvic fin | -0.575 | 0.248 | -0.191 |
Scale rows between lateral line and anal fin | -0.180 | 0.165 | 0.114 |
Scale rows around caudal peduncle | -0.687 | -0.103 | -0.194 |
Predorsal scale rows | -0.758 | -0.352 | 0.095 |
The Discriminant Function Analysis (DFA) performed using combined morphometric and meristic characters correctly classified all individuals of the new species (Table
Classification results of discriminant function analysis using (a) combined morphometric and meristic characters, (b) morphometric characters and (c) meristic characters of double barbeled
Species | Predicted count | Total | Error (%) | ||||
---|---|---|---|---|---|---|---|
1 | 2 | 3 | 4 | ||||
|
1. |
128 | 0 | 0 | 0 | 128 | 0.00 |
2. |
0 | 66 | 0 | 0 | 66 | 0.00 | |
3. |
0 | 0 | 24 | 1 | 25 | 4.00 | |
4. |
0 | 0 | 0 | 47 | 47 | 0.00 | |
|
1. |
124 | 1 | 3 | 0 | 128 | 3.13 |
2. |
0 | 66 | 0 | 0 | 66 | 0.00 | |
3. |
1 | 0 | 22 | 2 | 25 | 12.00 | |
4. |
0 | 0 | 0 | 47 | 47 | 0.00 | |
|
1. |
99 | 9 | 5 | 15 | 128 | 22.66 |
2. |
0 | 65 | 0 | 1 | 66 | 1.52 | |
3. |
0 | 0 | 25 | 0 | 25 | 0.00 | |
4. |
1 | 2 | 0 | 44 | 47 | 12.41 |
The DFA using morphometric measurements revealed morphological shape differences between the new species and the other previously described species of
In contrast, the DFA using meristic characters showed poor classification of individuals of the four species, with three individuals of the new species, 29 individuals of
Based on the deep genetic and significant morphological divergence between individuals from the Verlorenvlei River system and other members of the double barbeled redfin group, the Verlorenvlei population represents a new species.
Verlorenvlei redfin (English), Verlorenvlei rooivlerkie (Afrikaans).
South Africa: Western Cape Province: SAIAB186092, mature male, 70.8 mm standard length (SL), collected from the Verlorenvlei River, 20 m upstream from railway at the Het Kruis bridge on R365 (
(
The new species can be distinguished from its congeners by distinct linear speckles above and below the lateral line, anterior barbels minute and much smaller than eye diameter, lips unretracted, and a cartilaginous fig absent.
Proportional measurements and meristic characters are presented in Table
Comparisons of the morphometric measurements and meristic counts of
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Holotype | Paratypes | ||||
No. of specimens | 1 | 46 | 66 | 25 | 128 |
Standard length (SL) (mm) | 70.0 | 34.0–70.8 | 42.0–109.0 | 28.5–163.4 | 30.0–151.7 |
Head length (HL) (mm) | 18.9 | 10.0–19.6 | 10.9–25.0 | 8.6–51.7 | 8.1–45.8 |
Percentage of SL (%) | |||||
Head length | 27.0 | 28.1±0.2 | 24.9±0.1 | 30.5±1.5 | 26.8± 1.0 |
Predorsal length | 54.0 | 53.1±0.4 | 47.6±0.1 | 53.3± 1.7 | 51.2± 1.4 |
Dorsal fin base | 12.9 | 13.2±0.1 | 13.7±0.1 | 12.0± 0.8 | 13.4± 0.7 |
Dorsal fin height | 25.3 | 25.6±0.3 | 24.2±0.1 | 21.2± 1.8 | 22.9± 1.4 |
Body depth | 27.6 | 29.1±0.2 | 26.9±0.3 | 25.6± 1.3 | 25.6± 1.6 |
Body width | 16.1 | 16.4±0.2 | 15.2±0.3 | 17.9± 1.5 | 17.1± 1.6 |
Caudal peduncle length | 20.9 | 23.5±0.2 | 25.7±0.1 | 22.4± 0.8 | 25.0± 1.2 |
Percentage of HL (%) | |||||
Head depth | 72.5 | 73.0±0.4 | 74.1± 0.3 | 64.2± 3.1 | 70.1± 2.6 |
Inter-obit | 36.0 | 34.4±0.3 | 33.1±0.3 | 28.1± 2.1 | 31.3± 2.1 |
Snout length | 31.2 | 31.0±0.4 | 36.6±0.4 | 38.0± 2.2 | 36.5± 1.9 |
Post orbit | 46.0 | 47.6±0.5 | 46.4±0.3 | 45.4± 1.8 | 45.2± 1.9 |
Anterior barbel length | 6.3 | 3.3±0.3 | 5.1±0.2 | 20.3± 9.5 | 16.5± 4.3 |
Posterior barbel length | 30.7 | 20.8±0.8 | 19.3±0.3 | 27.5± 11.9 | 28.4± 5.6 |
Orbit diameter | 31.2 | 31.5±0.4 | 28.8±0.4 | 21.5± 4.4 | 27.7± 2.8 |
Percentage of caudal peduncle length (%) | |||||
Caudal peduncle depth | 61.0 | 52.6±0.8 | 46.9± 4.0 | 53.4± 3.6 | 49.4± 3.5 |
Unbranched dorsal fin rays | iv | iii (iii-v) | iii (iii–iv) | iii (iii–iv) | iv (iii–iv) |
Branched dorsal fin rays | 7 | 7 (7–8) | 7 (6–7) | 7 (7–8) | 7 (6–8) |
Unbranched anal fin rays | iii | iii (iii-iv) | iii (ii-iv) | iii (iii-iv) | iii (iii-iv) |
Branched anal fin rays | 5 | 5 | 5 (5–6) | 5 (4–5) | 5 (4–6) |
Pectoral fin rays | 14 | 15 (13–16) | 14 (13–16) | 13 (13–16) | 14 (13–16) |
Pelvic fin rays | 8 | 8 (7–9) | 8 (8–9) | 8 (7–8) | 8 (7–8) |
Lateral line scales | 33 | 32 (29–36) | 32 (28–37) | 38 (36–39) | 35 (29–37) |
Scale rows between lateral line and dorsal fin | 6 | 6 (5–6) | 5 (4–6) | 7 (6–7) | 6 (5–7) |
Scale rows between lateral line and pelvic fin | 4 | 5 (4–5) | 4 (3–5) | 5 (5–7) | 4 (4–5) |
Scale rows between lateral line and anal fin | 4 | 4 (4–5) | 4 (3–4) | 5 (4–6) | 4 (4–6) |
Caudal peduncle scale rows | 12 | 12 (12–16) | 12 (12–13) | 16 (15–18) | 12 (12–16) |
Predorsal scale rows | 16 | 16 (13–18) | 15 (12–16) | 19 (17–21) | 17 (14–22) |
Total vertebrae | 36 | 36 (34–37) | 37 (35–38) | 37 (36–38) | 36 (35–37) |
Pre-caudal vertebrae | 19 | 19 (18–21) | 19 (18–20) | 20 (19–21) | 19 (17–20) |
Caudal vertebrae | 17 | 17 (16–19) | 18 (16–19) | 17 (16–18) | 18 (17–20) |
Predorsal vertebrae | 11 | 11 (10–13) | 11 (10–12) | 12 | 12 (11–13) |
Pre-anal vertebrae | 20 | 20 (19–21) | 20 (19–22) | 21 (20–22) | 19 (18–21) |
Counts for the holotype are given in a separate column in Table
Scales moderately large; lateral line complete, majority of specimens have 32 scales along lateral line (range 29–36); 5–6 (mode 6) scale rows between dorsal fin origin and lateral line; 4–5 (mode 5) scale rows between pelvic fin origin and lateral line; 4–5 (mode 4) scale rows between lateral line and anal fin origin; 12–16 (mode 12) circumpeduncular scale rows. Predorsal scale rows 13–18 (mode 16), embedded in skin, smaller than flank scales. Patch between head and posterior base of pectoral fins naked; scales between posterior base of pectoral fins and anterior base of pelvic fins smaller than flank scales and embedded. Pelvic fins lack prominent or elongate axillary scales. Scales radiately striated.
Nuptial tubercles have only been observed in one individual of
Live colouration is golden-tan dorsally and laterally, becoming lighter and more silver ventrally (Figure
Lateral and dorsal view of
Radiographs of the holotype (SAIAB 186092) and paratypes show that the species has osteology typical of all
SAIAB59813, juveniles (n=68, 13.5–28.4 mm SL) and adults (n=3, 59.3–64.6 mm SL), collected from the Verlorenvlei River, near Grootfontein farm (
Unknown, but spawning possibly occurs around October-December, based on the general patterns of congeners.
Map of a part of the west coast of South Africa. The map shows the likely present distribution of
The species is named after the Verlorenvlei River system to which it is now confined.
The Verlorenvlei redfin was listed as Endangered during the most recent IUCN assessment by
1 | Mouth terminal, 36–39 lateral scale series |
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– | Mouth sub-terminal |
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2 | Lower lip unretractable, cartilaginous fig absent, conspicuous linear speckles above and below lateral line |
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– | Lower lip retractable, cartilaginous fig present |
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3 | Anterior barbels less than 30% of orbit diameter |
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– | Anterior barbels more than 30% of orbit diameter |
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Specimens of
Phylogenetic results from the present study,
Reduced tubercle occurrence in
This work was supported by the National Research Foundation (NRF) of South Africa under the Foundational Biodiversity Information Programme: Biodiversity surveys in priority inland areas (IBIP) grants (grant reference no. IBIP-BS13100251309). We thank Roger Bills, Daksha Naran and Whit Bronaugh for field collections, Rose Palmer for the illustration of