Aphanotorulus emarginatus ( Valenciennes 1840 )

Ray, C. Keith & Armbruster, Jonathan W., 2016, The genera Isorineloricaria and Aphanotorulus (Siluriformes: Loricariidae) with description of a new species, Zootaxa 4072 (5), pp. 501-539 : 512-516

Aphanotorulus emarginatus ( Valenciennes 1840)

( Fig. 7 )

Hypostomus emarginatus Valenciennes , in Cuvier & Valenciennes, 1840: 500 (369 in Strasbourg deluxe edition). Type locality: Probablement originaire du Brésil. Holotype: MNHN a-9447 (dry). Plecostomus annae Steindachner, 1881: 112 , pl. 3 (figs. 2–2a). Type locality: Pará [= Belem, Brazil]. Holotype: NMW 44073. Listed as Chaetostomus annae in figure caption (p. 146). Hypostoma squalinum Jardine , in Schomburgk, 1841: 142, pl. 2. Type locality: rios Rios Branco, Negro, and Essequibo. No types known.

Material examined. Brazil: ANSP 187270 (3) (ex MZUSP 88601), Rio Culuene (Xingu basin), Canarana/Gaucha do Norte, Gaucha do Norte, Mato Grosso, 13°30'53''S 53°5'34''W, O.Oyakawa, J.L. Birindelli, C. Moreira et al, 19 October 2004; MCZ 7805 (1) Pará [Belem and environs], Brazil, col. by L. Agassiz and D. Bourget; MCZ 7807 (2), Ilha de Marajo, Furo Tajapuru at Tajapuru, Pará, Brazil, 20–21 August 1865, col. by L. Agassiz; MCZ 7824 (1), Santarem, Pará, Brazil, August 1865, col. by D. Bourget; MCZ 7840 (2), Rio Ica; Rio Putomajo [Rio Ica (tributary of Rio Solimoes) near the Brazilian-Colombian border], Amazonas, Brazil, September 1865, col. by W. James and S. Talisman; MCZ 7860 (1), Obidos; Obydos [Rio Amazonas at Obidos], Pará, Brazil, December 1865, col. by W. James and W. Hunnewell; MCZ 7992 (1), GURUPA [Rio Amazonas delta at Gurupa], Pará, Brazil, 22 August 1865, col. by L. Agassiz et al.; MNRJ 12710 (1), rio Passa Três, afluente esquerdo rio Maranhão, alto Tocantins, Uruaçu, Goias, 14°30’S, 49º9’W, G.W. Nunan, D.F.. Moraes Jr., 26 October 1996; MNRJ 12781 (1), Rio Bagagem, à montante da foz do córrego Barriguda, afluente direito Rio Maranhão, alto Tocantins, Niquelândia, Goiás, 14º5’S 48º14’W, col. by G.W. Nunan & D.F. Moraes Jr., 15 October 1985; MNRJ 12932 (1), Rio Tocantins, em Porto do Garimpo, mun. Minaçu/Cavalcante, Minaçu, Goiás, 13º34’5”S 48º6’4”W, col. by D.F. Moraes Jr., 13 January 1988; MNRJ 12944 (1), Afluente do córrego Lageado (afluente margem esquerda Rio Tocantins) na estrada para Porto Rubião, Minaçu, Goiás, 13º38’85”S 48º19’12”W, col. by D.F. Moraes Jr., 16 January 1988; MNRJ 18360 (1), poças formadas por riacho da margem esquerda do Rio Tocantins, a montante de Ponte Rubão durante enchimento, Colinas do Sul, Goiás, col. by D.F. Moraes, D.A. Habolth, O.T. Oyakawa et al., 28 October 1996; MNRJ 19338 (1), Rio Tocantins, próximo da cidade de Tucuruí, Tucuruí, Pará, col. by L.C. Alvarenga; MNRJ 29536 (1), Riacho (drenagem do Juruena) na estrada MT-208, 1 km a oeste da balsa Ariel, Fazenda São Nicolau, município de Cotriguaçu. 9º52’42”S 58º14’40”W, col. by P.A. Buckup, L.F.S. Ingenito, I.L. Assumpção, F.B. Freitas, 6 May 2006; MNRJ 31177 thru 31196, Rio Tocantins, a jusante da UHE Serra da Mesa, poções formados durante o enchimento do reservatório, Município de Minaçu/Colinas do Sul, 13º49’S, 47º17’W, col. by D.F. Moraes, D.A. Halboth, O.T. Oyakawa, et al., 28 October–4 November 1996; MZUSP 24064 (1), Rio Tocantins, entre Mocajuba e Baião, Baião, Pará, 2º40’S 49º42’W, col. by Expedição Permanente à Amazônia, 9 September 1970; MZUSP 24086 (1), Lago às margens do igarapé Espírito Santo, entre Baiao e Tucuruí, Rio Tocantins, Baião/Tucuruí, Pará, 3º46’S 49º41’W, col. by Expedição Permanente à Amazônia, 11 September 1970; MZUSP 27637 (1), Rio Negro, Pedra do Gavião, Moura, Amazonas, 1º28’S 61º38’W, col. by L.P.S. Portugal, 13 November 1982; MZUSP 34550 (1), Igarapé do Cujobim, Rio Branco, em frente da ilha de Maracá, Roraima, 3º25’N 60º20’W, col. by M. Goulding, 13 January 1984; MZUSP 40478 (1), Rio Paranã (cachoeira), fazenda Olho d'Água, Flores de Goiás, Goiás, 14º26’S 47º3’W, col. by J.C. Oliveira & W.J.M. Costa, 12 September 1988; MZUSP 52308 (1), Rio Araguaia, Rio Tocantins, Mato Grosso; MZUSP 52327 (1), Rio Araguaia, Bandeirantes, Rio Tocantins, Mato Grosso, 13º40’S 50º48’W; MZUSP 52342 (1), Rio Araguaia, Araguaiana, Rio Tocantins, Mato Grosso, 15º43’S 51º51’W; MZUSP 54208 (1), Rio Araguaia, Ilha do Bananal, GO, Mato Grosso; MZUSP 54539 (1), Rio Araguaia, s/ dados de localidade, Rio Tocantins; MZUSP 86893 (2) Rio 7 de Setembro, afl. rio Xingu, ca. 20 km W Canarana pela estr. MT-020 (Canaran-Garapu), Brasil, Mato Grosso, Canarana, 13°30'19.0"S, 52°24'57.0"W, col. by Moreira, Landim, Nolasco and Datovo, 17 October 2004; MZUSP 87081 (3), Rio Batovi (Tamitatoala) afl. Xingu Salto da Alegria, faz. Dois Netos (prop. André Parente), Mato Grosso, Gaúcha do Norte, 13°14'46.0"S, 54°1'30.0"W, col. by Equipe AXE, 21 October 2004; MZUSP 87082 (4), Rio Batovi (Tamitatoala) afl. Xingu Salto da Alegria, faz. Dois Netos (prop. André Parente), Mato Grosso, Gaúcha do Norte, 13°14'46.0"S, 54°1'30.0"W, col. by Equipe AXE, 21 October 2004; MZUSP 89734 (1), Rio Culuene, trecho entre cachoeira até ribeirão Corgão, Mato Grosso, Paranatinga, 13°49'0.0"S, 53°15'0.0"W, col. by A. Akama & J.L. Birindelli, 15 January 2006; MZUSP 92720 (1), Rio Tapajós, na corredeira do Pajaú e arredores, Pará, Pimental, 4°35'4.0"S, 56°15'32.0"W, col. by L.M. Sousa and J.L. Birindelli, 11 November 2006; MZUSP 94085 (1), Rio Culuene, fazenda do Sr. Zezé (c. 2 km acima da ponte), Mato Grosso, Gaúcha do Norte, 13°30'53.0"S, 53°5'40.0"W, col. by F.C.T. Lima, F.A. Machado, C.A. Figueiredo and J.L. Birindelli, 21–26 May 2007; MZUSP 94415 (1), Lago da Miriam, margem direita do rio Culuene (23–24.v.2007), Mato Grosso, Canarana, 13° 25' 48.0" S, 53° 2' 24.0" W, col. by F.C.T. Lima, F.A. Machado, C.A. Figueiredo and J.L. Birindelli, 23 May 2007; MZUSP 95627 (5), Rio Teles Pires próximo da balsa da rodovia MT-416, Mato Grosso, Paranaíta, 9° 27' 7.0" S, 56° 30' 46.0" W, col. by L.M. Sousa and A.L. Netto-Ferreira, 2007; MZUSP 96132 (1), corredeiras no rio Teles Pires, abaixo das "Sete Quedas", Pará, Jacareacanga, 9°19'34.0"S, 56°46'52.0"W, col. by L.M. Sousa and A.L. Netto-Ferreira, 2007; MZUSP 96598 (1), Rio Peixoto de Azevedo, afl. Rio Teles Pires, próximo a cidade de Peixoto de Azevedo, Mato Grosso, Peixoto de Azevedo, 10°13'14.0"S, 54°58'2.0"W, col. by J.L. Birindelli, L.M. Sousa, A.L. Netto-Ferreira, M. Sabaj-Perez, N.K. Lujan, 16 October 2007; MZUSP 97217 (2), Rio Curuá, bacia do Iriri, na vila de Castelo dos Sonhos, Pará, Altamira, 8° 19' 7.0" S, 55° 5' 23.0" W, col. by J.L. Birindelli, L.M. Sousa, A.L. Netto-Ferreira, M. Sabaj-Perez, N.K. Lujan, 22 October 2007; NMW 44073 (1) (Holotype: Plecostomus annae Steindachner 1881 ), Pará [=Belem, Brazil]; USNM 191582 (1), Rio Araguaia, Near Aruana, -14.967 -51.400, col. by H.R. Axelrod, 1960. Columbia: ANSP 146867 (1), Rio Negrito at bridge on road joining Puerto Lopez and Villavicencio; 200– 400 yd downstream of bridge, Meta, J.E.Bohlke et al., 15 March 1973; AUM 35432 (1), Rìo Manacìas, Sandbar on right (E) bank 31 airkm SSW Puerto Gaitan, Meta, J.S. Ramsey, R. J. Scully, M.C. Blanco, et al., 10 October 1978. Guyana: ANSP 175912 (1), Essequibo River: 180 yd. upstream from Essequibo campsite (Maipuri), Siparuni VIII- 2, D. Allicock, 27 January 1997; ANSP 175913 (1), Essequibo River: sandbar ca. 800 m downstream from Essequibo campsite (Maipuri), Siparuni VIII-2, W.G. Saul, G.G. Watkins, N.R. Liley, C. Watson, 29 January 1997; AUM 35514 (1), Rupununi River 3.7 km SSE Massara, col. by J.W. Armbruster, M.H. Sabaj, D.C. Werneke, C.L. Allison, M.R. Thomas, C.J. Chin, D. Arjoon, S. Mario, S.M. James, 27 October 2002; AUM 35535 (2), Rupununi River 4.6 km NW Massara, col. by J.W. Armbruster, M.H. Sabaj, D.C. Werneke, C.L. Allison, M.R. Thomas, C.J. Chin, D. Arjoon, S.M. James, S. Mario, 26 October 2002; AUM 35551 (3), Rupununi River at Kwatamang, 4 km SE Annai, col. by J.W. Armbruster, M.H. Sabaj, D.C. Werneke, C.L. Allison, M.R. Thomas, C.J. Chin, D. Arjoon, S.M. James, 25 October 2002; AUM 35666 (2), Rupununi River at Karanambo, col. by J.W. Armbruster, M.H. Sabaj, D.C. Werneke, C.L. Allison, M.R. Thomas, C.J. Chin, D. Arjoon; AUM 36611 (1), Rìo Macaruma, 134 km SE. of Cuidad Guiana, 5 km SE. of Guasipati, at old bridge just W. of the main road, Bolivar, col. by J.W. Armbruster, D.C. Werneke, T.P. Pera, N.K. Lujan, and O. Leon, 11 June 2003; AUM 38885 (1), Takutu River 3.77 km SSW Lethem, Reg. 9, col. by J.W. Armbruster, M.H. Sabaj, M. Hardman, D. Arjoon, N.K. Lujan, L.S. de Souza, 1 November 2003; AUM 44344 (2), Essequibo River, shoreline and sandbeach downstream of Kurukapari Falls and upstream from Iwokrama, Region 8, col. by L.S. deSouza, N.K. Lujan, D.C. Taphorn, J.A. Hartsell, E. Liverpool, and S. Lord, 23 November 2005; AUM 45033 (1), Pond at Yukupari, Region 9, col. by N.K. Lujan, D.C. Taphorn, and E. Liverpool, 29 November 2005. Venezuela: ANSP 182988 (3), Rio Siapa (Casiquiare Dr.), raudale Gallineta, 142 km E of San Carlos de Rio Negro, M.H.Sabaj, D.C. Werneke, N.K.Lujan, M.Arce, 17 March 2005; AUM 39309 (1), Rio Manapiare, 14.5 km NW of San Juan de Manapiare, Amazonas, col. by N.K. Lujan, M.H. Sabaj, L.S. deSouza, and D.C. Werneke, 12 April 2004; AUM 39507 (1), Rio Ventuari, at Raudales Tencua, 56 km ESE of San Juan de Manapiare, Amazonas, col. by D.C. Werneke, N.K. Lujan, O.Leon, A. Luna, and R. Pajua; AUM 39836 (1), Rio Manapiare, 10 km NW of San Juan de Manapiare, Amazonas, col. by N.K. Lujan, L.S. deSouza, D.C. Werneke, and M.H. Sabaj, 14 April 2004; AUM 39844 (1), Rio Ventuari, beach below Raudales Tencua, 56 km ESE of San Juan de Manapiare, Amazonas, col. by N.K. Lujan, O. Leon, and R. Pajua; AUM 40579 (1), Rio Orinoco, at Macuruco landing, 75 km E. of San Fernando de Atabapo, Amazonas, col. by M.H. Sabaj, L.S. deSouza, D.C. Werneke, and N.K. Lujan, 4 April 2004; AUM 40751 (1), Rìo Ventuari, beach across the river from Picua Village, 34 km ENE of Macuruco, 104 km E of San Fernando de Atabapo, Amazonas, col. by M.H. Sabaj, N.K. Lujan, D.C. Werneke, L.S. deSouza, and O. Leon, 5 April 2004; AUM 41558 (2), Rio Manapiare, at Laja Pelada landing, 27 km SSW of San Juan de Manapiare, Amazonas, col. by O. Leon, D.C. Werneke, and N.K. Lujan, 18 April 2004; AUM 42092 (2), Rìo Orinoco, beach, 16.1 km W of La Esmeralda, Amazonas, col. by N.K. Lujan, M. Arce, E.L. Richmond, M.B. Grant, J. Valadez, D. Brooks, and T.E. Wesley, 25 March 2005; AUM 42122 (2), Rìo Orinoco, 33.9 km W of La Esmeralda, Punto Piaroa, Amazonas, col. by N.K. Lujan, M. Arce, T.E. Wesley, M.B. G., E.L. R., J. B., D. B., 29 March 2005; AUM 42128 (1), Rìo Casiquiare, bedrock in stream, 73 km NE of San Carlos de Rìo Negro, Amazonas, col. by N.K. Lujan, D.C. Werneke, M.H. Sabaj, M. Arce, R. Betancur, and T.E. Wesley, 9 March 2005; AUM 42165 (3), Rìo Orinoco, bedrock outcrop, 52.9 km SE of San Antonio, 102 km W of La Esmeralda, Amazonas, col. by N.K. Lujan, D.C. Werneke, M.H. Sabaj, O. Leon, M. Arce, R. Betancur, and T.E. Wesley, 4 March 2005; AUM 42182 (6), Rìo Siapa, Raudales Gallineta, 142 km E of San Carlos de Rìo Negro, Amazonas, col. by N.K. Lujan, D.C. Werneke, M.H. Sabaj, O. Leon, M. Arce, and T.E. Wesley, 17 March 2005; AUM 42196 (1), Rìo Casiquiare, bedrock outcrop, 59.5 km SW of La Esmeralda, Amazonas, col. by N.K. Lujan, D.C. Werneke, M.H. Sabaj, M. Arce, R. Betancur, T.E. Wesley, and O. Santa Ella, 8 March 2005; AUM 42199 (1), Rìo Casiquiare, 153 km NE of San Carlos de Rìo Negro, Amazonas, col. by N.K. Lujan, O. Leon, M. Arce, E.L. Richmond, M.B. Grant, J. Valadez, D. Brooks, and T.E. Wesley, 24 March 2005; AUM 42208 (1), Rìo Ventuari, near ornamental fish market in the river, Amazonas, col. by N.K. Lujan, M. Arce, E.L. R., M.B. G., and T.E. Wesley, 3 March 2005; AUM 42222 (3), Rìo Orinoco, near Puerto Ayacuho on a beach called Playa Bagre, Amazonas, col. by N.K. Lujan, M. Arce, and T.E. Wesley, 13 March 2005; AUM 42994 (2), Rìo Orinoco, at Puerto Venado, 4.3 km S of Samariapo, 56.4 km SSW of Puerto Ayacucho, Amazonas, col. by N.K. Lujan, D.C. Werneke, M.H. Sabaj, M. Arce, R. Betancur, and T.E. Wesley, 26 February 2005; AUM 43345 (3), Rìo Casiquiare, at mouth of Caño Caripo, 37 km WSW of La Esmeralda, Amazonas, col. by N.K. Lujan, D.C. Werneke, M.H. Sabaj, O. Leon, M. Arce, R. Betancur, and T.E. Wesley, 5 March 2005; AUM 44971 (15), Essequibo River, at Kwaimatta, beach in main channel and mouth of side channel, Reg. 9, col. by L.S. deSouza, N.K. Lujan, D.C. Taphorn, J.A. Hartsell, E. Liverpool, and S. Lord, 1 December 2005; INHS 29085 (1), Rio Mavaca, Amazonas, Venezuela, 1991; INHS 34779 (1), Rio Masparro, Barinas, Venezula, 1995; MCNG 20045 (1) Rio Capanaparo 200m. del Paso de la MAcanilla, en una playa, Dto. Pedro Camejo, 7°04’58”N 67°71’11”W, 5 April 1989; MCNG 21575 (1), Río Caura, río arriba de caño la Prisión, Bolivar, Dtto. Sucre, 7°00’00”N 64°66’67”W, col. by D.C.Taphorn, E. Suttón, E. Sanoja,A. Fernandez, 2 February 1989; MCNG 23800 (1), Rio Matacuni, al frente del N°4, Amazonas, Atabapo, col. by Basil Stergios, 25 January 1990; MCNG 28843 (1), Caño entrando a una laguna del Río Mavaca, Amazonas, Dpto. Río Negro, col. by L. Nico, G. Aragua, 29 January 1991; MCNG 30360 (1), Rio Orinoco al frente de Macuruco, Amazonas, col. by A.Barbarino, I.Lopez, R.Berrios, 5 August 1994; USNM 265664 (2), Rio Orinoco, Cove, Islote De Fajardo, 182 Naut. mi. Upstream From Sea Buoy, Bolivar, 8.367, - 62.700, col. by J. Baskin, D. Taphorn, 15 February 1978; USNM 269964 (2), Small Cano Connecting With Rio Orinoco Immediately South of El Burro, Bolivar, 6.183 -67.417, co. by R.P. Vari, O. Castillo, C.J. Ferraris, 9 December 1984.

Diagnosis. Aphanotorulus emarginatus differs from A. ammophilus and A. unicolor by the presence of a single, large buccal papilla (vs. multiple buccal papillae); from A. horridus by having circular spots on the dorsal and paired fins which remain unfused across the length of the fin (vs. spots that fuse into bands across the dorsal and paired fins, mostly in adults); and from A. phrixosoma by having fewer elongate hypertrophied odontodes covering the dorsal and ventral surfaces of body except the head (vs. many hypertrophied odontodes present—see A. phrixosoma description); and from A. gomesi only by distribution. Aphanotorulus emarginatus further differs from Isorineloricaria spinosissima , I. tennuicauda , and I. villarsi by having flat compound pterotics (vs. rounded ridge present in I. spinosissima and I. tennuicauda and a sharp ridge present in I. villarsi ); from I. villarsi by lack of sharp ridge on compound pterotics (vs. sharp ridge present); and from I. acuarius by having a smaller orbit diameter to interorbital width ratio (32.4–40.5% vs. 40.6–63.7%), which separates 96% of individuals, and by having smaller, more circular spots on the head (vs. larger, oval spots in I. acuarius ).

Description. Morphometrics given in Table 3 . Head moderately compressed. Supraoccipital crest not elevated, with posterior edge sloping gently into nape. Interorbital surface flat. Nape increasing in depth posteriorly to dorsal fin. Pectoral fin reaches past point of insertion of pelvic fin. Depressed pelvic spine reaches point of insertion of the anal fin. Adipose fin triangular.

Lateral line plates 26–31 (mode= 29; n= 118); dorsal-fin base plates six to nine (mode= seven; n= 118); folded dorsal plates nine to 14 (mode= 12; n= 118); plates between dorsal and adipose fin seven to 12 (mode= nine; n= 118); adipose fin plates two to three (mode= three; n= 117); anal-fin base plates two to three (mode= two; n= 118); plates from anal fin insertion to last plate on caudal peduncle 14–19 (mode= 18; n =118); plates in folded pectoral fin five to eight (mode= six; n =116); number of teeth on dentary 10–45 (mode= 26; n =118); number of teeth on premaxilla 14–45 (mode= 29; n =118).

Sexual Dimorphism. Breeding males with hypertrophied odontodes on pectoral-fin spine, which increase in density and length distally along spine. Paired-fin spines swell and become larger distally. Odontodes present on caudal fin, with largest odontodes on spines and rays and also covering posteromedial edge of most plates on lateral surface of body; most relatively short (less than 2–3mm). Longest plate odontodes occur on mid-dorsal, median, and mid-ventral plate rows. Slightly lengthened odontodes present on cheek.

Color: Light tan to white background. Spotting pattern highly variable across range. Spots small (less than or equal to pupil width) to medium in size; can be very dense (less background color showing) to very sparse (more background color showing) across entire body. Ventral surface with some spots across pectoral girdle or no spots. Spots on fin rays similar to those on body. Spots irregularly placed on paired-fin rays. Dorsal-fin membrane with two distinct rows of spots between each fin ray. Lower caudal fin lobe dark in color, almost black in some specimens.

Distribution. ( Fig. 8 ) A wide-ranging species, A. emarginatus ’s range includes the upper Orinoco River (except the Rio Apure); the Essequibo River and its tributaries; the lower Amazon River including the Rio Negro, the Rio Tapajos, and the Rio Xingu.

Figures

FIGURE 7. Dorsal, lateral, and ventral views of Aphanotorulus emarginatus, MZUSP 97217, 227.0 mm SL.

FIGURE 8. Distribution of Aphanotorulus emarginatus. Type locality is unknown.

Tables

TABLE 3. Selected morphometrics of Aphanotorulus emarginatus and A. horridus.: Dorsal-fin spine lengths are not included for either species because too many specimens had broken spines, and it was too difficult to determine accurate lengths.

	Aphanotorulus emarginatus	Aphanotorulus horridus
Measurement	N Average SD Range	N Average SD	Range
Standard Length (SL, mm)	153 46.5–357.0	73	44.9–513.0
% SL
Predorsal Length	153 36.5 2.3 29.8–41.9	73 36.8 2.2	29.6–41.2
Head Length (HL)	153 29.8 2.5 23.3–36.2	73 30.2 2.3	23.5–38.8
Cleithral Width	153 23.8 1.5 20.0–27.2	72 24.8 1.8	19.2–27.6
Dorsal spine Length	–		–
Folded dorsal-fin Length	153 36.0 1.7 30.0–41.5	72 36.5 1.2	31.1–40.9
Dorsal-fin base Length	153 23.5 1.0 20.1–28.3	73 23.6 1.2	19.6–26.0
Dorsal-adipose Length	153 24.6 2.0 19.6–31.4	73 24.2 1.8	20.3–28.5
Thorax Length	153 23.6 1.4 20.0–27.5	73 24.2 1.6	19.2–28.7
Pectoral spine Length	153 25.4 2.1 18.8–31.4	72 25.9 2.3	18.1–30.3
Abdominal Length	153 20.3 1.1 16.5–24.8	73 20.2 1.2	18.0–24.8
Pelvic spine Length	153 20.1 1.8 14.1–25.6	73 20.8 2.4	14.7–26.0
Postanal Length	153 39.9 2.8 32.9–45.9	73 39.8 4.1	20.6–46.8
Caudal peduncle Depth	153 7.2 0.6 5.7–8.9	73 7.8 0.9	5.6–9.1
Caudal peduncle Width	153 16.7 1.3 13.4–21.0	73 16.7 1.5	10.0–19.2
Adipose-fin spine Length	151 6.7 1.0 4.2–9.8	72 6.8 1.0	4.6–11.0
Anal-fin Length	153 12.0 1.6 6.8–17.5	72 13.2 1.7	10.4–17.9
Anal-fin base Length	152 4.0 0.5 2.3–5.3	72 4.1 0.6	1.9–5.8
Adipose-upper caudal Length	152 20.4 1.2 17.0–23.5	73 21.2 1.6	16.6–24.7
Adipose-lower caudal Length	153 24.8 1.3 20.4–28.2	72 25.7 1.3	22.7–28.9
%HL
Head Depth	153 57.3 3.4 48.7–73.5	73 59.0 3.5	50.5–66.0
Snout Length	153 60.4 2.4 55.7–73.5	73 59.1 3.8	42.0–71.3
Orbit Diameter	153 19.4 3.0 14.4–36.5	73 17.9 2.4	11.7–24.2
Interorbital Width	153 38.5 2.0 34.1–44.3	73 43.5 3.9	30.0–49.5
Snout opercle Distance	153 71.2 2.5 66.7–86.3	73 70.5 3.2	53.0–77.3
Head Width	153 80.3 3.4 72.0–97.3	73 82.6 3.4	67.4–90.4
Head eye Length	153 36.5 2.6 32.5–61.1	73 39.4 1.6	32.8–43.9
Eye nare Length	153 11.4 1.4 6.9–15.1	73 11.9 1.2	9.0–14.3
Internares Width	153 17.4 1.6 12.3–22.5	73 18.0 1.9	11.5–22.2
Snout nares Length	153 41.3 3.1 36.7–68.2	73 38.6 4.5	14.1–44.1
Snout pectoral Length	153 74.1 3.0 68.9–92.0	73 73.0 3.4	54.7–80.1
Pectoral orbit Distance	153 27.1 3.5 19.1–37.0	73 27.2 2.8	20.3–34.2
Mouth Width	153 41.8 3.8 34.8–54.3	73 40.7 3.1	34.8–49.4
Mouth Length	153 43.4 3.7 33.3–59.7	73 41.9 3.5	30.4–47.8
Dentary Length	153 13.1 1.9 7.4–19.9	73 12.5 1.2	8.8–15.0