Characidium cricarense, Malanski & Sarmento-Soares & Silva-Malanski & Lopes & Ingenito & Buckup, 2019

Malanski, Evandro, Sarmento-Soares, Luisa Maria, Silva-Malanski, Ana Cecilia Gomes, Lopes, Maridiesse Morais, Ingenito, Leonardo Ferreira da Silva & Buckup, Paulo Andreas, 2019, A new species of Characidium (Characiformes: Crenuchidae) from coastal basins in the Atlantic Rainforest of eastern Brazil, with phylogenetic and phylogeographic insights into the Characidium alipioi species group, Neotropical Ichthyology 17 (2), pp. 1-13 : 4-8

Characidium cricarense , new species

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Fig. 1 , Tabs. 1-2

Holotype. MNRJ 51282 , 49.0 mm SL, Brazil, Espírito Santo, São Mateus municipality, São Mateus basin, rio Cricaré at Cachoeira do Inferno rapids, 18°42’24.5”S 40°16’7.2”W, 18 m above sea level, 15 Aug 2017, P.A. Buckup, D.F. Moraes Junior, E. Malanski & V. de Britto.

Paratypes. All from Brazil, Espírito Santo. Rio Itaúnas basin: MNRJ 50962 , 46 (2 for DNA, 2 CS), 41.3-47.7 mm SL (13, 41.3-47.7 mm SL), Pedro Canário municipality, rio Itaúnas at Cachoeira da Mata waterfall, north of Floresta do Sul , 18°12’11.6”S 40°4’36.3”W, 17 m above sea level, 15 Aug 2017, P.A. Buckup, D.F. Moraes Junior, E. Malanski & V. de Britto. Rio São Mateus basin: MNRJ 50973 , 16 (2 for DNA, 2 CS), 39.2-48.0 mm SL (4, 39.2-48.0 mm SL), MBML 13549, 5, 42.0- 48.8 mm SL, collected with holotype. Rio Doce basin: MNRJ 41832 , 33 (10 for DNA), 40.7-55.9 mm SL (9, 40.7-55.9 mm SL), Santa Teresa municipality, rio Santa Maria do Rio Doce at Cachoeira do Rúdio waterfall, 19°50’5”S 40°41’12”W, 159 m above sea level, 5 Oct 2013, J. R. Gomes, E. Neuhaus & C.C.D. Quijada .

Non-types. Locality data has been abbreviated to include only the municipality, state abbreviation, and name of main river. All from Brazil, grouped according to hydrographic basins from north to south. Rio Jucuruçu basin: MBML 7501 (3 for DNA), 12, 22.2-31.2 mm SL, Palmópolis, MG, rio do Prado; MBML 7508 (1 for DNA), 2, 39.3- 39.7 mm SL, Palmópolis, MG, córrego Paulino Laranjeira; MBML 12879, 1, 30.3 mm SL, Itamaraju, BA, tributary of rio Jucuruçu. Rio Itanhém-Alcobaça basin: MBML 3846, 5, 33.7-36.7 mm SL, Medeiros Neto, BA, córrego do Medeiros Neto. Rio Mucuri basin: MNRJ 38334, 8, 29.2-50.2 mm SL, Teófilo Otoni, MG, rio Todos os Santos; MNRJ 38335, 32, 23.3-39.3 mm SL, Teófilo Otoni, MG, rio Todos os Santos; MNRJ 38336, 11, 33.4-40.5 mm SL, Teófilo Otoni, MG, rio Todos os Santos; MZUSP 54818, 25, 18.4-34.4 mm SL, Teófilo Otoni, MG, riacho São Pedro; MZUSP 54821, 8, 25.0- 38.6 mm SL, Teófilo Otoni, MG, rio Todos os Santos. Rio Itaúnas basin: CZNC 874 (2 for DNA), 41, 38.8-44.0 mm SL, Montanha, ES, rio Itaúnas; CZNC 1376, 57, 28.4-40.7 mm SL, Pedro Canário, ES, rio Itaúnas; MBML 4782, 1, 31.4 mm SL, Conceição da Barra, ES, córrego Grande; MBML 4807, 4, 22.7-25.2 mm SL, Conceição da Barra, ES, córrego Grande; MBML 4808, 6 (1 for DNA), 22.2-23.1 mm SL, Pedro Canário, ES, córrego Douradão; MBML 4830, 2, 29.6 mm SL, Conceição da Barra, ES, córrego Grande; MBML 7217, 4, 34.7 mm SL, Pinheiros, ES, córrego Santo Antônio; MBML 7223, 19, 22.8-31.9 mm SL, Pinheiros, ES, tributary of córrego Santo Antônio; MBML 7227, 1 (1 for DNA), 25.8 mm SL, Pinheiros, ES, córrego Santo Antônio; MBML 7235, 2, 31.0- 32.1 mm SL, Pinheiros, ES, córrego do Veado; MBML 7248, 2, 32.8-38.5 mm SL, Pinheiros, ES, córrego São Roque; MBML 7261, 1, 22.1 mm SL, Pinheiros, ES, córrego do Veado; MNRJ 37635, 41 (4 for DNA), 21.3- 40.8 mm SL, Barra de São Francisco, ES, rio Itaúnas; MZUSP 54817, 80, 18.5-39.6 mm SL, Pinheiros, ES, rio do Sul. Rio São Mateus basin: CZNC 167, 6, 32.4-34.4 mm SL, São Mateus, ES, rio Cricaré; CZNC 1047, 6, 39.1- 41.2 mm SL, São Mateus, ES, rio Cricaré; CZNC 1389, 8, 39.6-39.8 mm SL, Ecoporanga, ES, rio Dois de Setembro; CZNC 1443, 45 (1 for DNA), 37.4-40.0 mm SL, Vila Pavão, ES, córrego da Rapadura; CZNC 1467, 9, 45.4 mm SL, Vila Pavão, ES, córrego Alecrim; CZNC 1477 (2 for DNA), 17, 36.4-48.7 mm SL, Barra de São Francisco, ES, rio Cricaré; CZNC 1506 (1 for DNA), 20, 36.8-47.0 mm SL, Água Doce do Norte, ES, rio Preto; CZNC 1531, 11, 34.3 mm SL, Ecoporanga, ES, rio Cotaxé; CZNC 1535, 6, 34.3- 36.8 mm SL, Ataléia, ES, rio do Norte; CZNC 1545, 10, 26.4-47.7 mm SL, Ecoporanga, ES, rio Dois de Setembro; CZNC 1556, 50, 32.1-34.1 mm SL, Ataléia, ES, rio Paraju; CZNC 1576, 1, 37.4 mm SL, Ecoporanga, ES, ribeirão Santa Rita; CZNC 1593 (1 for DNA), 37, 33.6-34.5 mm SL, Ecoporanga, ES, córrego Lajeado; MBML 3480, 1, 40.9 mm SL, Água Doce do Norte, ES, córrego Água Doce; MBML 3484, 15, 28.0- 43.2 mm SL, Água Doce do Norte, ES, rio Preto; MNRJ 50845, 1, 34.8 mm SL, Boa Esperança, ES, rio Cotaxé; MZUSP 54820, 5, 36.0- 37.2 mm SL, Boa Esperança, ES, rio Braço Norte. Rio Doce basin: MBML 4369, 6, 31.4-43.7 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MBML 4371, 15, 29.1-40.1 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MBML 4381, 2, 32.4-46.6 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MBML 4388, 6, 38.7-53.0 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MBML 4403, 1, 34.2 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MBML 4422, 3, 33.5- 33.9 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MBML 4456, 1, 44.5 mm SL, Santa Teresa, ES, tributary of rio Santa Maria do Rio Doce; MBML 7636, 10, 44.6-52.7 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MNRJ 22420, 11, 38.7-55.6 mm SL, Santa Teresa, ES, rio Santa Maria; MNRJ 36190, 10, 23.9-42.6 mm SL, Laranja da Terra, ES, ribeirão Lagoa; MNRJ 37625, 5, 30.0- 39.2 mm SL, Baixo Guandu, ES, rio Mutum; MNRJ 37730, 8, 27.7-34.3 mm SL, Águia Branca, ES, rio Braço Sul; MNRJ 37737, 6, 32.6-50.5 mm SL, Águia Branca, ES, córrego Água Calma; MNRJ 41879, 7 (5 for DNA), 34.9-49.2 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce; MNRJ 50842, 55, 36.1-46.7 mm SL, Santa Teresa, ES, rio Santa Maria do Rio Doce at cachoeira do Rúdio.

Diagnosis. The new species is included in a group of species of Characidium with an unscaled area in the isthmus (vs. isthmus scaled in C. bahiense , C. bimaculatum Fowler, 1941 , C. boehlkei Géry, 1972 , C. borellii (Boulenger, 1895) , C. caucanum Eigenmann, 1912 , C. chupa Schultz, 1944 , C. clistenesi , C. deludens , C. etheostoma Cope, 1872 , C. etzeli Zarske, Géry, 2001 , C. geryi (Zarske, 1997) , C. heinianum Zarske, Géry, 2001 , C. heirmostigmata Graça, Pavanelli, 2008 , C. interruptum Pellegrin, 1909 , C. lagosantense Travassos, 1947 , C. lanei Travassos, 1967 , C. laterale (Boulenger, 1895) , C. litorale Leitão, Buckup, 2014 , C. longum Taphorn, Montaña, Buckup, 2006 , C. marshi Breder, 1925 , C. mirim Netto-Ferreira, Birindelli, Buckup, 2013 , C. nana Mendonça, Netto-Ferreira, 2015 , C. nupelia Graça, Pavanelli, Buckup, 2008 , C. occidentale Buckup, Reis, 1997 , C. orientale Buckup, Reis, 1997 , C. papachibe Peixoto, Wosiacki, 2013 , C. pellucidum Eigenmann, 1909 , C. phoxocephalum Eigenmann, 1912 , C. pteroides Eigenmann, 1909 , C. rachovii Regan, 1913 , C. roesseli Géry, 1965 , C. samurai , C. sanctjohanni Dahl, 1960 , C. satoi Melo, Oyakawa, 2015 , C. schindleri Zarske, Géry, 2001 , C. serrano Buckup, Reis, 1997 , C. steindachneri Cope, 1878 , C. stigmosum Melo, Buckup, 2002 , C. tapuia Zanata, Ramos, Oliveira-Silva, 2018 , C. tenue (Cope, 1894) , C. vestigipinne Buckup, Hahn, 2000 , C. xanthopterum Silveira, Langeani, da Graça, Pavanelli, Buckup, 2008 , C. xavante Graça, Pavanelli, Buckup, 2008 , and C. zebra Eigenmann, 1909 ). Within this group of species, Characidium cricarense is distinguished from other species, except C. alipioi Travassos, 1955 , C. fasciatum , C. hasemani Steindachner, 1915 , and C. kamakan , by the presence of 5-7 scales and distance between the anus and the anal-fin greater than 10% SL (vs. 2-4 scales and distance shorter than 10% SL in C. boavistae Steindachner, 1915 , C. grajahuense Travassos, 1944 , C. helmeri , C. japuhybense Travassos, 1949 , C. lauroi Travassos, 1949 , C. oiticicai Travassos, 1967 , C. pterostictum Gomes, 1947 , C. schubarti Travassos, 1955 , C. summum Zanata, Ohara, 2015 , C. timbuiense , C. travassosi Melo, Buckup, Oyakawa, 2016 , and C. vidali Travassos, 1967 ), and 14 series of scales around the caudal peduncle (vs. 12 or less series of scales around caudal peduncle in C. amaila Lujan, Agudelo-Zamora, Taphorn, Booth, López-Fernández, 2013 , C. brevirostre Pellegrin, 1909 , C. crandellii Steindachner, 1915 , C. declivirostre Steindachner, 1915 , C. gomesi Travassos, 1956 , C. macrolepidotum (Peters, 1868) , C. purpuratum Steindachner, 1882 and C. sterbai (Zarske, 1997)) . Characidium cricarense is distinguished from C. alipioi by having 4 series of scales above the lateral line (vs. 5 series) and a greater distance between the anus and the anal fin (11.4-15.5 vs. 10.1-11.1% SL); from C. fasciatum and C. kamakan by the smaller body depth at the dorsalfin origin (16.3-20.0 vs. 20.1-23.1 and 20.6-21.8% SL, respectively), at the anal fin origin (13.0-15.5 vs. 16.1-17.2 and 16.6-17.8% SL, respectively), and at the caudal peduncle (8.8-10.5 vs. 11.6-12.5 and 12.0-12.9% SL, respectively); from C. hasemani by the short distances between the tip of the snout and the pelvic fin (46.1-50.2 vs. 53.2-55.6% SL), the tip of the snout and the anal fin (72.1-76.4 vs. 76.4-79.9% SL), and the tip of the snout and the tip of anal fin (87.5-92.2 vs. 94.4-100.6% SL).

Description. In Tab. 2 , morphometric data is summarized for holotype and paratypes. Characidium cricarense has elongate, fusiform body ( Fig. 1 ). Greatest body depth at dorsal-fin origin. Dorsal profile in lateral view convex from snout to origin of dorsal fin; convex, posteriorly slanted at base of dorsal fin, almost straight between base of dorsal fin and adipose fin; posteriorly slanted at base of adipose fin, almost straight between adipose fin and caudal fin. Ventral profile in lateral view slightly concave between lower lip and base of pectoral fin, slightly straight from pectoral base to anal-fin origin, straight and slightly slanted at base of anal fin, straight below caudal peduncle. Pelvic-fin origin at vertical line about two scales posterior to dorsal-fin origin. Adipose fin at vertical line about two and a half scales posterior to anal-fin origin.

Snout directed downwards, dorsally convex in lateral view ( Fig. 1a ). Mouth subterminal, aligned with or lower to ventral margin of orbit. Posterior tip of maxilla not reaching level of anterior margin of orbit. Orbit oval, slightly elongate along axis from tip of snout to parietal bone; margin of eye free. Nares widely separated; distance between nares larger than distance between posterior naris and eye. Margin of anterior naris expanded, forming circular rim; posterior naris with expanded margin on its anterior-most portion. Parietal branch of supraorbital laterosensory canal present, reaching parietal bone in larger specimens; frontal branch of supraorbital laterosensory canal present. Fontanel reduced, limited anteriorly by frontals, laterally by parietals and posteriorly by supraoccipital bones.

Premaxillary teeth 6(3) in single series; teeth decreasing gradually in size posteriorly, unicuspid or tricuspid. An anomalous specimen with premaxillae fused, with 13 teeth in total. Dentary teeth in two rows; outer series of dentary teeth 8(2) or 9(2) unicuspid or triscuspid with small medial and lateral cuspids; teeth decreasing gradually in size posteriorly; inner series of dentary teeth with several tiny conical teeth on edge of replacement tooth trench. Maxillary teeth absent. Ectopterygoid teeth present, in single row, conical, unicuspid. Mesopterygoid teeth absent. Branchiostegal rays 4(1), 5(2) or 6 including one as vestigial (1); 3(1), 4(2) or 4 including one as vestigial (1) attached to anterior ceratohyal. Gill rakers on first arch 11(3) or 12(1).

Scales cycloid; parallel radii present on posterior field. Lateral series of scales 35(4), 36(25*), or 37(3), all perforated by lateral-line canal. Scales above lateral line 4(32*). Scales below lateral line 4(32*). Scales around caudal peduncle 14(32*). Pre-dorsal scale series arranged in single, regular row; scales in pre-dorsal series 8(1), 9(7), 10(17), 11(6*), or 12(1). Scales between anus and origin of anal fin 5(5), 6(23*), or 7(4). Isthmus without scales; chest scaled ( Fig. 1d ).

Dorsal-fin rays i,9(1), ii,8(3), ii,9(27*), or ii,10(1). Adipose fin present (32*). Pectoral-fin rays iii,8,i(3), iii,9(4), iii,9,i(15*), iii,10(9), or iii,10,i(1). Pelvic-fin rays i,7,i(23*), or i,8(9). Anal-fin rays i,6(2), i,7(3), ii,6(5), or ii,7(21*); adnate rays on last pterygiophore of anal fin 1(21*), or 2(8). Principal caudal-fin rays i,9,7,i(1) or i,9,8,i(12*).

Postcleithrum I absent; and postcleithrum II and postcleithrum III present in pectoral girdle; cleithrum with posteriorly directed process at region immediately below ventral tip of supracleithrum. Precaudal vertebrae 19(2) or 20(6); total vertebrae 33(1), 34(3), 36(3) or 37(1). Supraneurals 4(3), 5(3) or 6(1). Upper procurrent rays 7(1), 8(6) or 9(2); lower procurrent rays 6(7), 7(1) or 8(1). Epurals 3(3).

Coloration in alcohol. Background color yellowish ( Fig. 1 ). Black longitudinal stripe about half scale thick extending from tip of snout to base of caudal fin. Body with 8 to 13 triangular bars, wider dorsally, thinner ventrally; bars from opposite sides united across dorsal midline by a corresponding series of dorsal blotches; bars located posteriorly to pelvic fin extending to ventral surface of body. Humeral blotch partially merged into first bar. Dark spot present at base of middle caudal fin rays. All fins mostly hyaline, with scattered chromatophores along rays forming bands of variable intensity. Dorsal fin with three bands; main (intermediate) band dark, not reaching anterior-most margin of fin. In smaller specimens, caudal-fin pigmentation restricted to some scattered chromatophores at outer lobes, ontogenetically developing into conspicuous incomplete B-shaped pattern in larger specimens, formed by a posterior extension of the lateral stripe and diagonal bands across the dorsal and ventral portions of fin. Pectoral, pelvic and anal fins with dark, diffuse distal bands.

Color in life. Slightly translucent body, with olive-greenish or light-brownish background color on head and dorsum. Silvery background below lateral stripe. Dark chromatophore patterns as described for specimens preserved in ethanol.

Sexual dimorphism. No hooks on fins or other sexually dimorphic features were observed externally on the examined specimens.

Molecular delimitation. DNA barcodes based on the amplified 656bp (base pairs) partial sequence of COI mitochondrial gene were generated for a total of 36 specimens of Characidium cricarense ( Tab. 1 ), encompassing specimens from the Jucuruçu basin (4 individuals), Itaúnas basin (6 individuals), São Mateus basin (11 individual), and Doce basin (15 individuals). All samples exhibit less than 2% of genetic divergence, which is the usually accepted upper limit of conspecific similarity ( Pereira et al., 2013). Characidium cricarence received the Barcode Index Number (BIN) BOLD:ACI3743 through the BoldSystems automated algorithm ( Ratnasingham, Hebert, 2013).

The genetic distance of Characidium cricarense from other congeneric species analyzed were 6% divergence from C. alipioi specimens from Macaé (MNLM 3004, MNLM 3007) and São João (MNLM 5462) basins; 11% divergence from C. timbuiense specimen from Reis Magos basin (MNLM 5403); 12% divergence from C. vidali specimen from Guanabara basin (MNLM 2043) ( Fig. 2 ). The specimen used as outgroup, C. litorale from São João basin (MNLM 7660), is 23% divergent from the remaining species in the analysis.

Geographical distribution. Characidium cricarense is known from the southern portion of Bahia State, northern portion of Espírito Santo State, and eastern portion of Minas Gerais State, where it is known from the rio Jucuruçu, rio Itanhém-Alcobaça, rio Mucuri, rio Itaúnas, and rio São Mateus drainages, extending to the rio Doce drainage in the central portion of the Espírito Santo State, Brazil ( Fig. 3 ). The rivers from the northern portion of the distribution are within the coastal tablelands of the Barreiras sedimentary formation. These coastal tablelands correspond to a region of mild relief, with elevations usually not surpassing 150 m above sea level, formed by Cenozoic sediments, well-seen along the border between the States of Espírito Santo and Bahia ( Sarmento-Soares, Martins-Pinheiro, 2013). On the right bank of the rio Doce, C. cricarense reaches the lower slopes of the Serra de Santa Teresa. According to a global biogeographic regionalization of freshwater systems, this geographic distribution belongs to the Northeastern Mata Atlântica ecoregion ( Abell et al., 2008).

Ecological notes. Characidium cricarense inhabits lowland stretches of rivers, usually below 200 m above sea level, except in the Mucuri drainage, where it has been found in the upper rio Todos os Santos, above the city of Teófilo Otoni at about 500 m above sea level (G. M. Mattox, pers. comm.), and in the Jucuruçu drainage, in tributaries around the city of Palmópolis at about 300 and 600 m above sea level. Based on our field observations, individuals of this species usually occupy the edge of turbulent, moderately fast flowing rivers and streams ( Fig. 4 ). Specimens aggregate along stretches less than 1.5 m deep, with transparent waters, occupying the waterfalls in rocky areas, and unconsolidated bottom over sandy, pebbles or gravel areas at the edge of the waterfall.

Etymology. The name refers to the rio Cricaré, where type locality of this fish is found, and is used as a Latinized geographic adjective.

Conservation status. Characidium cricarense occurs in coastal basins from Northeastern Mata Atlântica ecoregion (sensu Abell et al., 2008) along three Brazilian states: Bahia, Espírito Santo and Minas Gerais. The species inhabits turbulent waters in rapids, occupying the rocky areas as well as the unconsolidated bottom near the rapids. Even though a mining disaster occurred in Doce basin, where Characidium cricarense might occur, no specific threats to the species is detected in the whole area of distribution. Consequently, the conservation status of Characidium cricarense may be classified as Least Concern (LC), according to the International Union for Conservation of Nature (IUCN) categories and criteria (IUCN, 2017).

Figures

Fig. 1. Characidium cricarense, new species. MNRJ 50962, 45.8 mm SL, paratype in a. lateral, b. dorsal, and c. ventral views; rio Itaúnas, rio Itaúnas basin. In details: d. the unscaled isthmus, the scaled pectoral base, and e. the area between the anus and the anal-fin.

Fig. 2. Phylogenetic tree based on molecular data, generated with the maximum likelihood method and HKY+G model. MNLM codes and associated data are listed in Tab. 1. River basins are indicated next to species names. Divergence values are indicated along the branches, except for those below 0.5% which were suppressed. Characidium alipioi species group represented in yellow, Characidium grajahuense species group in green, and C. litorale (outgroup) in blue.

Fig. 3. Map of eastern South America showing Characidium cricarense distribution: the red star indicates the type locality, yellow stars indicate localities collection of paratypes, and black dots represent localities collection of non-type specimens. The National Hydrographic Division (CNRH 2003) is represented in thick black lines on highlighted map; red-shaded area represents the main map, at the edge of East Atlantic and Southeast Atlantic basins.

Fig. 4. Fast flowing stream, type locality of Characidium cricarense, at Cachoeira do Inferno rapids in rio Cricaré, São Mateus, ES, Brazil. GPS coordinates: 18°42’24.5”S 40°16’7.2”W.

Tables

Tab. 2. Morphometric data for Characidium cricarense n. sp. based on holotype (MNRJ 51282) and paratypes (MNRJ 41832, n=9; MNRJ 50962, n=13; MNRJ 50973, n=4; MBML 13549, n=5). Distances are based on landmarks described in Melo, Oyakawa (2015); ‘n’ is the number of specimens examined; SD is the standard deviation.

Distances	Character	Holotype	n	Range	Mean	SD
1–2v	Total length (mm)	58.02	32	47.29 – 64.80	53.8	-
1–3	Standard length (mm)	49.00	32	39.23 – 55.92	45.16	-
Percentage of standard length
1–4	Head length	20.41	32	19.01 – 24.05	21.63	1.24
1–5	Prepectoral distance	19.23	32	19.10 – 22.40	20.40	0.90
5–6	Pectoral-fin height	18.17	32	16.60 – 20.49	18.17	0.91
1–7	Predorsal distance	41.48	32	41.48 – 44.66	43.29	0.82
7–8	Dorsal-fin height	17.23	32	15.13 – 20.33	17.39	1.36
7–9	Dorsal-fin base	15.21	32	11.85 – 15.84	13.89	0.96
1–10	Prepelvic distance	47.44	32	46.14 – 50.16	48.02	1.12
10–11	Pelvic-fin height	11.37	32	10.28 – 13.62	11.50	0.78
1–13	Preanal distance	73.22	32	72.06 – 76.37	74.15	1.12
1–15	Anal-apex distance	89.30	32	87.48 – 92.21	89.60	1.09
12–13	Anus to anal-fin distance	12.78	32	11.36 – 15.55	13.20	1.09
13–14	Anal-fin height	13.62	32	10.73 – 13.79	12.85	0.69
13–16	Anal-fin base	7.41	32	5.77 – 8.37	7.03	0.65
17–18	Adipose-fin height	5.96	32	4.09 – 6.65	5.57	0.61
16–3	Peduncle length	19.84	32	17.18 – 21.94	19.80	1.13
7–7’	Body depth at dorsal-fin origin	17.62	32	16.33 – 20.03	18.26	0.97
13–13’	Body depth at anal-fin origin	14.86	32	13.03 – 15.46	13.89	0.64
19–19’	Body depth at caudal peduncle	10.02	32	8.78 – 10.54	9.83	0.40
5 (left-right)	Body width	10.43	32	8.72 – 11.99	10.25	0.72
	Percentage of head length
1–20	Snout length	28.80	32	24.47 – 30.93	28.25	1.42
1–23	Snout-maxillary tip	24.70	32	22.00 – 27.27	25.04	1.44
24–20	Anterior naris-orbit	14.00	32	11.63 – 15.56	13.61	0.87
25–20	Posterior naris-orbit	7.20	32	5.73 – 8.28	6.72	0.64
22–26	Cheek	14.10	32	11.38 – 15.24	13.04	0.85
20–21	Orbital diameter	25.50	32	25.11 – 29.00	26.84	0.98
27 (left-right)	Interorbital diameter	27.90	32	23.24 – 31.12	27.57	1.9