Research Article |
Corresponding author: Matheus V. Volcan ( matheusvolcan@hotmail.com ) Academic editor: Peter Bartsch
© 2018 Matheus V. Volcan, Francisco Severo-Neto, Luis Esteban K. Lanés.
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:
Volcan MV, Severo-Neto F, Lanés LEK (2018) Unrecognized biodiversity in a world’s hotspot: three new species of Melanorivulus (Cyprinodontiformes: Rivulidae) from tributaries of the right bank of the Rio Paraná basin, Brazilian Cerrado. Zoosystematics and Evolution 94(2): 263-280. https://doi.org/10.3897/zse.94.24406
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The genus Melanorivulus presents a wide geographical distribution in the Neotropical region. Among Melanorivulus, the M. pictus species group has currently 18 species distributed in the Brazilian Cerrado, mainly in the upper Rio Paraná basin, with some species occurrences in the upper Rio Araguaia and Tocantins. In the present study, we describe three new Melanorivulus species, belonging to the M. pictus species group from different drainages of the right bank of the Rio Paraná basin in Mato Grosso do Sul state, Brazil. These new species are easily distinguished from the others by their unique colour patterns. Melanorivulus interruptus is distinguished from all species of the M. pictus group by the presence of oblique chevron-like red bars interrupted, mainly on the midline of the flank in males; while M. ivinhemensis by the yellow colouration of the caudal fin with thin red bars arranged only in the median region of the fin in males. Melanorivulus amambaiensis is distinguished from all species of the M. pictus species group by having an orange anal fin or sometimes more reddish-orange with distal margin grey or dark grey and chevron-like bars along the body, distinctly branched ventrally forming an inverted Y-shape in males. The high diversity of the Melanorivulus species with high levels of endemism demands the development of conservation strategies to avoid the loss of their vulnerable habitats in the Cerrado biome. We expect presence of more species of the M. pictus species group also along the lower reaches of the tributaries of the Rio Paraná. Additionally a dichotomic identification key of the M. pictus species group is provided.
Cerrado, killifish, conservation, diversity, taxonomy
Savanna biomes are distributed across tropical zones around the world. In South America, the Cerrado is the largest savanna biome, the second largest biome in Brazil (smaller only than the Amazon), and represents about 23 % of Brazilian territory (
The rivulid genus Melanorivulus is a typical component of the Cerrado fauna, but presents a wide geographical distribution in the Neotropical region, being also found in areas under the influence of the Chaco, Pantanal, Atlantic Forest and Amazon (e.g.
Among Melanorivulus species, the M. pictus species group is distinguished from all other congeners, by the presence of a vestigial ventral process of the angulo-articular (vs. process well-developed), curved first epibranchial (vs. approximately straight) and intense greenish blue or greenish golden to purplish blue flank above anal-fin base in males (vs. never similar colour pattern) (
A previous analysis indicates that the most recent common ancestor of Melanorivulus probably occupied a region comprising the eastern Amazon savanna and the ecotone Amazon-Cerrado, and the current distribution is the result of a series of dispersal and vicariance events during the evolutionary history of the genus (
In December 2016, a 12-day collection campaign was conducted to sample potential habitats for the occurrence of killifish species in the Mato Grosso do Sul state, Brazil, which included the largest river basins associated with Rio Paraná. In order to define the sampling strategy, we analysed satellite images from Google Earth (earth.google.com) as well as locations of specimens vouched in the Coleção Zoológica da Universidade Federal de Mato Grosso do Sul. Field works were undertaken travelling the main highways and roads to reach access to previously selected areas; at every site we performed an active search for fishes.
Fish samples were taken with a dip-net (D-shaped hand net, 60 cm × 40 cm, 2 mm mesh size), and then were euthanized with clove oil, fixed in situ with 4% formaldehyde, and later transferred into 70% ethanol. The material was collected under IBAMA/ICMBio authorization (process number 56894-1).
Morphological characters were obtained from specimens fixed in formalin after collection, and subsequently transferred to 70% ethanol. Fish measurements were taken point-to-point with digital calipers to the nearest 0.1 mm on the left side of the specimen following
Descriptions of colour patterns were based on photographs of both sides of live specimens photographed in the field, which were fixed in situ after photos, and individuals maintained in aquaria (not preserved). Institutional abreviations are
Melanorivulus interruptus is distinguished from all species of the M. pictus group, except M. nigromarginatus, by the presence of oblique chevron-like bars interrupted, mainly on the midline of the flank in males (vs. bars continuous, vestigial, irregular or presence of longitudinal rows of red dots on the side of the body; never bars interrupted on the midline of the body). It is distinguished from all other species in the M. pictus species group, except M. planaltinus, M. leali and M. pictus, by the presence of a rounded dorsal fin (vs. slight pointed or pointed dorsal fin). In addition, M. interruptus differs from M. amambaiensis, M. apiamici, M. egens, M. faucireticulatus, M. ivinhemensis and M. leali by a higher caudal fin ray count (32–33 vs. 28–31); from M. egens, M. faucireticulatus, M. leali, M. litteratus, M. rutilicaudus and M. formosensis by the position of the anal-fin origin between the pleural ribs of 15th and 16th vertebrae (vs. 13th and 15th vertebrae); from M. apiamici, M. egens, M. faucireticulatus, M. pictus, M. polychromus and M. proximus by the dorsal-fin origin at a vertical through the base of the 8th anal-fin ray (vs. vertical through base of 7th or 9th-10th anal-fin ray); from M. amambaiensis, M. egens, M. faucireticulatus, M. ivinhemensis, M. planaltinus, M. polychromus, M. nigropunctatus, M. ofaie and M. formosensis by the dorsal-fin origin between neural spines of 18th and 19th vertebrae (vs. 19th and 22th); from M. egens, M. faucireticulatus, M. leali, M. litteratus, M. scalaris, M. vittatus, M. linearis and M. proximus by the tip of pelvic fin reaching the urogenital papilla in males (vs. tip of pelvic fin reaching 1st to 4th anal fin ray). Another interesting diagnostic character is the presence of 8 rays in the pelvic fin in about 35% of the analysed specimens (vs. 5–7 rays, never 8 rays present in the pelvic fin).
Morphometric data are presented in Table
Morphometric data for the holotype and paratypes of Melanorivulus interruptus sp. n.
Holotype | Males (4) | Females (7) | |
---|---|---|---|
Standard length | 32.3 | 21.6–31.0 | 22.3–29.0 |
Percentages of standard length | |||
Body depth | 25.6 | 20.8–24.5 | 21.3–26.0 |
Caudal peduncle depth | 14.4 | 13.3–14.8 | 11.4–14.8 |
Predorsal length | 73.3 | 75.0–77.1 | 73.3–77.9 |
Prepelvic length | 54.6 | 52.6–53.9 | 51.8–56.3 |
Dorsal fin base length | 11.8 | 11.9–13.7 | 10.7–12.7 |
Anal fin base length | 21.2 | 18.4–22.5 | 17.8–19.4 |
Caudal fin length | 28.0 | 26.6–32.2 | 26.8–29.4 |
Pectoral fin length | 18.3 | 19.0–20.6 | 17.7–20.2 |
Pelvic fin length | 12.1 | 12.6–13.9 | 8.5–10.4 |
Head length | 26.2 | 26.8–27.5 | 24.9–27.7 |
Percentages of head length | |||
Head depth | 77.9 | 66.7–78.0 | 66.1–76.2 |
Head width | 76.8 | 66.7–73.6 | 74.7–82.1 |
Snout length | 18.5 | 12.6–17.7 | 15.1–19.2 |
Lower jaw length | 19.0 | 16.4–21.7 | 18.7–21.6 |
Eye diameter | 28.2 | 28.6–31.9 | 31.9–35.6 |
Short dorsal and anal fins. Dorsal-fin rays 9–10. Dorsal fin rounded in males and females. Dorsal-fin origin on vertical through base of 8th anal-fin ray, and between neural spines of 18th and 19th vertebrae. Anal-fin rays 12–13. Anal fin slightly pointed in males and females. Origin of anal fin at vertical through pleural ribs of 15th-16th vertebrae. Caudal fin rounded 32–33 rays. Pectoral fin rays 14. Pectoral fins rounded, with posterior margin reaching vertical at about 60–90% of length between pectoral-fin and pelvic-fin bases. Pelvic-fin rays 7–8. Pelvic-fin posterior tip reaching vertical at anus to 3rd anal-fin ray. Pelvic-fin bases in close proximity.
Scales cycloid. Body and head entirely scaled, except anterior ventral surface of head. Body squamation extending over anterior 15–25% of caudal-fin base. No scales on dorsal and anal-fin bases. Frontal squamation E-patterned; E-scales not overlapping medially; scales arranged in regular circular pattern around A-scale without exposed margins; transverse row of scales anterior to H-scale. Longitudinal series of scales 30–32; transverse series of scales 8–9; scale rows around caudal peduncle 16. No contact organs on flank and fins.
Cephalic neuromasts: supraorbital 3+3, parietal 1, anterior rostral 1, posterior rostral 1, infraorbital 1+9-11+1, preorbital 2, otic 1, postotic 1-2, supratemporal 1, median opercular 1, ventral opercular 1, preopercular 2+4, mandibular 2-3+1, lateral mandibular 1-2, paramandibular 1. Two neuromasts on caudal-fin base.
Six branchiostegal rays. Gill rakers on first branchial arch 1+7. First epibranchial slightly curved. Total number of vertebrae 29–30, 13–14 precaudal vertebrae, 16–17 caudal vertebrae. Ventral process of angulo-articular short, pointed. Vomerine teeth 1–3. Dermosphenotic present. Basihyal sub-triangular, greatest width 45–50% of length; basihyal cartilage 20–25% of total basihyal length. Second pharyngobranchial teeth absent.
Males (Figs
Females (Fig.
Melanorivulus interruptus is only known so far from a small stream, in the headwaters of the Rio Pardo, a tributary of the right margin of Rio Paraná, state of Mato Grosso do Sul, Brazil (Fig.
The name interruptus, from the Latin adjective interrupted, referring to the presence of conspicuous oblique red bars forming chevron-like marks usually disrupted in the midline of the body in males.
The species was recorded in a typical Cerrado area, inserted into the urban area of Campo Grande town at an altitude of 594 m a.s.l.. It was recorded in a small first order creek, but was only captured in border areas with dense aquatic vegetation, mainly grasses (Fig.
There are imminent threats to the population of M. interruptus: the stream is located in the Campo Grande urban area, which is developing rapidly around the area of the new species; there is a large real estate development alongside the stream that can cause significant impacts on the environment quality; the stream is crossed by a high traffic road and it is subject to accidents with potentially dangerous loads; besides, there are agricultural activities over its complete extension. Although larger collection efforts are likely to be undertaken in the area to better understand the species distribution pattern, M. interruptus appears to have a very restricted distribution. In the adjacent watersheds distinct species occur (e.g. M. apiamici in the mouth of the Rio Pardo, M. egens in the upper Rio São Domingos and M. nigropunctatus and M. ofaie in the Rio Verde – see Figure
Melanorivulus ivinhemensis is distinguished from all other species of the M. pictus species group by having a yellow caudal fin with thin red bars arranged only in the median region of the fin in males (vs. red bars absent or red bars extending from the ventral or median region to the dorsal region of the caudal fin), except in M. rutilicaudus. Besides, it is distinguished from all M. pictus species by the origin of the anal fin at a vertical through pleural ribs of 15th-18th vertebrae (vs. 13–15 vertebrae), except in M. amambaiensis, M. planaltinus, M. scalaris, M. polychromus, M. ofaie, M. interruptus and M. nigropunctatus; and by the greater snout length in males (16.1–21.3 vs. 11.4–16.3), except in M. amambaiensis, M. apiamici, M. interruptus, M. vittatus, M. polychromus and M. nigropunctatus. Distinguished from M. planaltinus, M. rutilicaudus, M. interruptus by a lower caudal fin ray count (29–31 vs. 32–34) and from M. amambaiensis by a higher caudal fin ray count (29–31 vs. 25–28). Distinguished from M. faucireticulatus, M. giarettai, M. interruptus, M. leali, M. vittatus and M. formosensis by a short basihyal cartilage 10–15% of total basihyal length (vs. 20–25%). From M. interruptus, M. vittatus, M. ofaie, M. nigropunctatus, M. proximus, M. linearis and M. nigromarginatus it is distinguished by a smaller dorsal fin base length in females (8.9–10.4% vs. 10.5–14.8%). Distinguished from M. giarettai, M. proximus, M. linearis, M. nigromarginatus and M. planaltinus by lower number of gill rakers on the first branchial arch (1+7 vs 1-2+8), and from the M. pictus by the dorsal-fin origin on a vertical through base of 8th or 9th anal-fin ray (vs. on vertical through base of 7th anal-fin ray).
Morphometric data are presented in Table
Morphometric data for the holotype and paratypes of Melanorivulus ivinhemensis sp. n.
Holotype | Males (n=9) | Females (n=9) | |
Standard length | 30.1 | 18.6–29.5 | 18.5–30.4 |
Percentages of standard length | |||
Body depth | 23.4 | 19.3–23.5 | 19.1–23.1 |
Caudal peduncle depth | 14.3 | 12.4–14.3 | 11.7–14.0 |
Predorsal length | 75.6 | 74.0–79.0 | 75.1–79.4 |
Prepelvic length | 54.9 | 51.8–55.5 | 53.3–58.5 |
Dorsal fin base length | 9.4 | 9.3–12.4 | 8.9–10.4 |
Anal fin base length | 19.1 | 17.2–23.3 | 16.3–20.9 |
Caudal fin length | 29.2 | 24.9–31.0 | 26.6–31.6 |
Pectoral fin length | 17.1 | 18.4–20.8 | 17.5–20.7 |
Pelvic fin length | 11.4 | 9.1–12.2 | 8.5–10.4 |
Head length | 25.0 | 25.1–28.0 | 23.8–28.2 |
Percentages of head length | |||
Head depth | 71.6 | 60.5–73.6 | 61.7–70.8 |
Head width | 81.1 | 70.5–76.2 | 72.3–82.0 |
Snout length | 17.3 | 16.1–21.3 | 12.9–20.6 |
Lower jaw length | 20.6 | 18.3–22.0 | 16.8–21.6 |
Eye diameter | 33.0 | 29.6–37.0 | 29.1–34.4 |
Short dorsal and anal fins. Dorsal-fin rays 7–9. Dorsal fin rounded or slightly pointed in males, rounded in females. Dorsal-fin origin on a vertical through base of 8th or 9th anal-fin ray, and between neural spines of 20th and 22th vertebrae. Anal-fin rays 12–14. Anal fin slightly pointed in males and females. Origin of anal fin at a vertical through pleural ribs of 15th-18th vertebrae. Caudal fin oval shaped, deeper than long, 29–31 rays. Pectoral fin rays 11–13. Pectoral fins rounded, with posterior margin reaching vertical at about 60–90% of the length between pectoral-fin and pelvic-fin bases. Pelvic-fin rays 6–7, one individual with one single pelvic fin. Posterior tip of pelvic fin reaching vertical between anus to 2nd anal-fin ray. Position of pelvic-fin bases variable, in close proximity, or with bases separated by a great distance, similar to the size of the base of the pelvic fin.
Scales small, cycloid. Body and head entirely scaled, except anterior ventral surface of head. Body squamation extending over anterior 15–25% of caudal-fin base. No scales on dorsal and anal-fin bases. Frontal squamation E and F-patterned or with two patterns, one on each side in one specimen. E-scales generally not overlapping medially. In one specimen E-scales marginally overlapped. Transverse row of scales anterior to H-scale; scales arranged in regular circular pattern around A-scale. A-scale usually without exposed margins, four specimens with posterior margin exposed, overlapping the B-scale. Longitudinal series of scales 29–31; transverse series of scales 8–9; scale rows around caudal peduncle 16. No contact organs on flank and fins.
Cephalic neuromasts: supraorbital 3+3, parietal 1, anterior rostral 1, posterior rostral 1, infraorbital 1+11–12+1, preorbital 2, otic 1, postotic 1–2, supratemporal 1, median opercular 1, ventral opercular 1–2, preopercular 2+4–6, mandibular 3–4+1, lateral mandibular 1–2, paramandibular 1. Two neuromasts on caudal-fin base.
Six branchiostegal rays. Gill rakers on first branchial arch 1+7. First epibranchial slightly curved. Total number of vertebrae 29–31, 14 precaudal vertebrae, 15–17 caudal vertebrae. Ventral process of angulo-articular short, pointed. Vomerine teeth 2–5. Dermosphenotic present. Basihyal sub-triangular, greatest width 45–50% of length; basihyal cartilage 10–15% of total basihyal length. Second pharyngobranchial teeth absent.
Males (Fig.
Females (Fig.
Melanorivulus ivinhemensis is so far only known from first order streams and small wetlands associated with both margins of the lower course of the Rio Ivinhema, Rio Paraná basin (Figure
The name ivinhemensis is a reference to the occurrence of the new species in the Rio Ivinhema basin.
Melanorivulus ivinhemensis was recorded in marginal areas of small first order streams and in wetlands completely exposed to the sun on a slightly undulating terrain at altitudes ranging from 267 to 315 m a.s.l. (Fig.
The new species is endemic of Rio Ivinhema and was recorded in four different areas, in a range of about 25 km along both sides of the river and there is a large density of potential environments for its occurrence along this stretch. It has been recorded in areas with no major impacts – although under intense agricultural activity – and therefore there is no evidence that the species is threatened with extinction.
The presence of an orange or orange-red anal fin with grey or dark grey distal margin (vs. no similar colour pattern) and chevron-like red bars in inverted Y-shape in the flank of males (vs. no similar pattern) distinguishes the M. amambaiensis from all other species of the M. pictus species group. Additionally, the new species is distinguished by the lower caudal fin ray count (25–28 vs. 29–34), except M. faucireticulatus; by the lower body depth in males (29.4–21.6 mm SL vs. 21.8–26.5), except from M. vittatus, M. polychromus, M. nigropunctatus, M. interruptus and M. ivinhemensis; by a lower caudal peduncle depth in males (12.4–13.5 mm SL vs. 13.5–16.8 mm SL), except in M. polychromus, M. nigropunctatus, M. interruptus and M. ivinhemensis. Females are distinguished from M. apiamici, M. faucireticulatus, M. giarettai, M. planaltinus, M. rutilicaudus, M. scalaris, M. nigropunctatus, M. proximus, M. linearis, M. nigromarginatus and M. formosensis by the lower body depth (19.1–21.5 mm SL vs. 21.6–26.0 mm SL); and from M. apiamici, M. egens, M. faucireticulatus, M. giarettai, M. planaltinus, M. nigropunctatus, M. linearis, M. nigromarginatus and M. ofaie by a shorter pre-dorsal length (74.5–76.6 mm SL vs. 76.4–83.3 mm SL).
Morphometric data are presented in Table
Morphometric data for the holotype and paratypes of Melanorivulus amambaiensis sp. n.
Holotype | Males (n=5) | Females (n=11) | |
---|---|---|---|
Standard length | 27.9 | 20.9–25.9 | 24.0–28.7 |
Percentages of standard length | |||
Body depth | 21.6 | 20.4–20.9 | 19.1–21.5 |
Caudal peduncle depth | 12.9 | 12.4–13.5 | 11.7–13.7 |
Predorsal length | 76.1 | 74.9–79.1 | 74.5–76.4 |
Prepelvic length | 56.8 | 53.6–55.3 | 52.0–54.5 |
Dorsal fin base length | 12.3 | 10.1–12.0 | 9.3–11.4 |
Anal fin base length | 21.4 | 19.1–21.4 | 18.5–20.7 |
Caudal fin length | 27.5 | 28.1–30.7 | 26.5–28.6 |
Pectoral fin length | 17.1 | 17.9–21.0 | 17.3–20.1 |
Pelvic fin length | 11.3 | 9.4–12.0 | 8.5–11.6 |
Head length | 26.5 | 26.2–26.9 | 24.6–27.6 |
Percentages of head length | |||
Head depth | 69.9 | 61.6–69.5 | 62.4–71.4 |
Head width | 75.7 | 70.5–72.4 | 70.1–79.3 |
Snout length | 15.2 | 16.7–17.8 | 15.2–20.6 |
Lower jaw length | 20.6 | 18.3–21.3 | 18.0–21.0 |
Eye diameter | 28.7 | 28.5–32.1 | 28.7–33.1 |
Short dorsal and anal fins. Dorsal-fin rays 8–9. Dorsal fin slightly pointed in males, rounded in females. Dorsal-fin origin on a vertical through base of 8th or 9th anal-fin ray, and between neural spines of the 20th and 21th vertebrae. Anal-fin rays 12–14. Anal fin slightly pointed in males and females. Origin of anal fin at a vertical through pleural ribs of 15th–16th vertebrae. Caudal fin oval shaped, longer than deep, 25–28 rays. Pectoral fin rays 11–13. Pectoral fins rounded, with posterior margin reaching vertical at about 70–90% of length between pectoral-fin and pelvic-fin bases. Pelvic-fin rays 6–7. Posterior tip of pelvic fin reaching a vertical at slightly anterior to the anus to 1st anal-fin ray. Pelvic-fin bases in close proximity.
Scales small, cycloid. Body and head entirely scaled, except anterior ventral surface of the head. Body squamation extending over anterior 15–20% of caudal-fin base. No scales on dorsal and anal-fin bases. Frontal squamation E and F-patterned or with two patterns, one on each side (present in one specimen). E-scales not overlapping medially. Transverse row of scales anterior to H-scale; scales arranged in regular circular pattern around A-scale. A-scale without exposed margins. Longitudinal series of scales 27–31; transverse series of scales 8–9; scale rows around caudal peduncle 16. No contact organs on flank and fins.
Cephalic neuromasts: supraorbital 3+3, parietal 1, anterior rostral 1, posterior rostral 1, infraorbital 1+9–11+1–2, preorbital 2, otic 1, postotic 1–2, supratemporal 1, median opercular 1, ventral opercular 1–2, preopercular 2+4–5, mandibular 2–3+1, lateral mandibular 1–2, paramandibular 1. Two neuromasts on caudal-fin base.
Six branchiostegal rays. Gill rakers on first branchial arch 1+7–8. First epibranchial slightly curved. Total number of vertebrae 30–31, 13–14 precaudal vertebrae, 17 caudal vertebrae. Ventral process of angulo-articular short, pointed. Vomerine teeth 2–3. Dermosphenotic present. Second pharyngobranchial teeth absent.
Males (Fig.
Females (Fig.
The species is only known from its type locality, a small first order drainage and direct tributary of Rio Amambaí, state of Mato Grosso do Sul, Brazil (Fig.
The name amambaiensis is a reference to the occurrence of the new species in the Rio Amambaí basin.
Melanorivulus amambaiensis was recorded in a small natural drainage, direct tributary of the Rio Amambaí, with depth not exceeding 50 cm and with low flow in an area totally exposed to the sun, parallel to BR 487 highway (Fig.
The species is known only from its type locality. It seems to be a micro endemic of the Rio Amambaí basin, since other species occur in the adjacent basins. Part of its original area was fragmented by BR 487 and there is high frequency traffic of vehicles on this highway. Consequently, the species is subject to accidents with potentially dangerous loads, which present a threat to the population of M. amambaiensis. In addition, the surrounding region is largely degraded by agricultural activities that also threaten its habitat quality. Thus, M. amambaiensis was accounted as “Vulnerable” according to the
1 | Red pigmentation forming chevron-like bars in the body of males | 2 |
– | Red pigmentation in the flank arranged in irregular lines to form vermiculate color pattern in males | Melanorivulus giarettai |
2 | Pelvic fin present, 1–5 vomerine teeth | 3 |
– | Absence or extreme reduction of pelvic fin, 5–7 vomerine teeth | Melanorivulus planaltinus |
3 | 5 rays in the pelvic fin, 10 rays in the pectoral fin, 27 vertebrae | Melanorivulus apiamici |
– | 6–8 rays in the pelvic fin, 11–14 rays in the pectoral fin, 29–32 vertebrae | 4 |
4 | Red bars present in the caudal fin | 5 |
– | Red bars absence in the caudal fin | Melanorivulus egens |
5 | 25–28 caudal fin ray, anal fin orange or orange-red in males | Melanorivulus amambaiensis |
– | 28–34 caudal fin ray, anal fin yellow, light yellow to orange or yellowish-hyaline in males | 6 |
6 | 7 scales in transversal series, oblique rows of red dots on flank, more concentrated on caudal peduncle, forming chevron-like marks | Melanorivulus leali |
– | 8–10 scales in transversal series, oblique red bars forming chevron-like bars along the body, with forward-pointing vertex, generally in the posterior portion of the body | 7 |
7 | Chevron-like red bars interrupted on midline of flank, fragmented or vestigial in the body of adult males | 8 |
– | Chevron-like red bars well defined, generally not interrupted in the body of males | 10 |
8 | Dorsal fin rounded in males, 14 pectoral fin rays | Melanorivulus interruptus |
– | Dorsal fin pointed or slightly pointed in males, 12–13 pectoral fin rays | 9 |
9 | Frontal squamation E-patterned, pectoral fin hyaline and pelvic fin light yellow in males | Melanorivulus nigromarginatus |
– | Frontal squamation F-patterned, pectoral and pelvic fin orange in males | Melanorivulus nigropunctatus |
10 | Red bars present only on middle of caudal fin in males | 11 |
– | Caudal fin with dorsal and middle portions presenting vertical red bars | 12 |
11 | 10–12 dorsal fin rays, 32–34 caudal fin rays, 34–35 scales in longitudinal series, 10 scales in transversal series | Melanorivulus rutilicaudus |
– | 7–9 dorsal fin rays, 29–31 caudal fin rays, 29–31 scales in longitudinal series, 8–9 scales in transversal series | Melanorivulus ivinhemensis |
12 | Dorsal fin origin on vertical of 7th ray of anal fin, 11–12 anal fin ray, dorsal fin rounded | Melanorivulus pictus |
– | Dorsal fin origin on vertical of 7–10th ray of anal fin, 12–14 anal fin ray, dorsal fin pointed or slightly pointed | 13 |
13 | 6 pelvic fin rays, dorsal fin origin between neural spines of vertebrae 21th-22th, absence of red color in dorsal fin | Melanorivulus polychromus |
– | 6–7 pelvic fin rays, dorsal fin origin between neural spines of vertebrae 18th-21th, red pigmentation present in dorsal fin | 14 |
14 | Frontal squamation E-patterned | 15 |
– | Frontal squamation F-patterned | Melanorivulus litteratus |
15 | Broad sub-basal red strip on dorsal fin in males, black reticulate pattern in ventral portion of the head of females, 28–30 caudal fin rays | Melanorivulus faucireticulatus |
– | No distinctive broad sub-basal red strip on dorsal fin in males, sometimes black pigmentation present in ventral portion of the head of females never forming an reticulate pattern, 30–34 caudal fin rays | 16 |
16 | One gill raker on upper limb of first branchial arch | 17 |
– | Two gill rakers on upper limb of first branchial arch | Melanorivulus proximus |
17 | Red dots present on the anteroventral portion of flank, distinctive dark marks on humeral region in males | 18 |
– | Red dots absent on the anteroventral portion of flank, absence of dark marks on humeral region in males | Melanorivulus linearis |
18 | Flank greenish blue to metallic blue, with oblique red bars forming narrow chevron-like bars, posterior tip of pelvic-fin reaching vertical at anus to 1st anal-fin ray | 19 |
– | Flank intense metallic blue with broad red chevron-like oblique bars, posterior tip of pelvic-fin reaching vertical at 1st to 4th anal-fin ray | 20 |
19 | 8 scales in transversal series, origin of anal fin at vertical through pleural ribs of 14th–15th vertebrae, chevron-like red bars with a branching on the upper portion forming Y-shaped red marks in males | Melanorivulus formosensis |
– | 9–10 scales in transversal series, origin of anal fin at vertical through pleural ribs of 15th-16th vertebrae, chevron-like red bars in males never forming Y-shaped red marks | Melanorivulus ofaie |
20 | Small red squares in the flank aligned to form broad red chevron-like oblique bars in males, 12–13 pectoral fin rays, 2 vomerine teeth | Melanorivulus scalaris |
– | Broad chevron-like oblique red bars in males not forming squares, 14 pectoral fin rays, 3–4 vomerine teeth | Melanorivulus vittatus |
Relationships among species belonging to the M. pictus group are not clear. Usually there is a high overlap in meristic, morphometric and osteological characters among congeners, however with well defined, homogeneous and structured colour patterns in each species (Fig.
The three new species present distinct and unique colour patterns, mainly in relation to the pigment and arrangement of the chevron-like bars along the body, the colour pattern of the fins, as well as the position and shape of the bars in the caudal fin. Males of M. ivinhemensis, for example, have a yellow caudal fin, with vertical bars arranged only in the median portion of the fin, M. interruptus has a yellow caudal fin with diagonal red bars, sometimes vestigial and inconspicuous, extending from the median to the dorsal region, while males of M. amambaiensis have a hyaline or yellowish-hyaline caudal fin with yellowish-orange ventral portion and irregular brownish-red bars. This is always combined with an orange or orange-red anal fin with a dark grey margin (Fig.
It is hypothesized that M. interruptus is closely related to M. linearis as judged by the absence of distinctive dark marks on the humeral region, and the absence of red dots on the anteroventral portion of the flank. With M. egens it shares the E-pattern of the frontal squamation, the yellow unpaired fins and the absence or reduction of red bars or red pigmentation at the base of the anal fin in males. Females of these three species also present all unpaired fins yellow with a dark grey distal margin as well as hyaline pectoral fins.
Melanorivulus ivinhemensis shares some characteristics with M. rutilicaudus and seems to be more closely related to this species. Mainly this is because both have the unpaired fins coloured yellow and red bars are present only in the middle of the caudal fin in males, a unique pattern among the species of the M. pictus species group. On the other hand, relationships between M. amambaiensis and the other species of the M. pictus species group are not clear because of the unique colour pattern. Chevron-like red bars with inverted Y-shaped red marks and an orange or orange-red anal fin with a dark grey distal margin in males are not encountered in any other species of the group.
Species of the M. pictus group generally present a high degree of endemism, occurring in small stretches of basins that drain the upper Rio Paraná and the upper Rio Araguaia and Tocantins (See Figure
Most species of the M. pictus group are reported for highlands, mainly in the headwaters of the main tributary drainage on the right bank of the Rio Paraná (
All from Brazil, Mato Grosso do Sul state. Melanorivulus nigropunctatus: Holotype.
This study was supported by Instituto Chico Mendes de Proteção da Biodiversidade (ICMBio) and funded by Fundação Grupo Boticário de Proteção à Natureza as part of the project “Peixes Anuais dos Campos Sulinos”. The manuscript benefited from corrections and suggestions made by Wilson Costa, Marcelo Loureiro and Dalton Nielsen.