Profundulidae, Hoedeman & Bronner, 1951

Domínguez-Cisneros, Sara E., Domínguez-Domínguez, Omar, Velázquez-Velázquez, Ernesto & Pérez-Rodríguez, Rodolfo, 2023, Redescription and diagnoses of the genera Profundulus and Tlaloc (Cyprinodontiformes: Profundulidae), Mesoamerican endemic fishes, Neotropical Ichthyology (e 220089) 21 (1) : -

Key to identification of Profundulidae species

1a. Humeral spot absent; dorsal fin origin posterior to a vertical through the anal fin origin; mesethmoid prominent and oval-shaped (c&s specimens), protruding from the posterior margins of the vomer ( Tlaloc ) ........................................................................................................................2

1b. Humeral spot present (except in P. adani ); dorsal fin origin at or anterior to a vertical through the anal fin origin; mesethmoid small and crescent-shaped (c&s specimens), not protruding from the body of the vomer ( Profundulus ) ...................................................................................................5

2a. Lower jaw broad, heavy, and protruding, so that the upper jaw is included ................. 3

2b. Jaws equal, or the lower jaw is included in the upper jaw ...................................... 4

3a. Thirty-six or 37 vertebrae in total; dark spot at the base of the caudal fin; 36 scales in lateral series (range: 33 to 37); origin of the anal fin to the base of the caudal fin relatively long (36.3–41.9% in the SL). Interior basin of the Chiapas highlands, Mexico (endemic) ................................................................. T. hildebrandi

3b. Thirty-three or 34 vertebrae in total; 33 scales in lateral series (range: 29 to 34); origin of the anal fin to the base of the caudal fin relatively short (29.0– 36.7% in SL). Interior basin of the highlands of Honduras (endemic) T. portillorum

4a. Jaws equal; 33 to 36 vertebrae in total; dorsal fin in males with two or three series of black points; middle rays of the anal fin in females, similar in size to that of males. Interior basin, Comitán River and adjacent basins of Mexico and spring at Candelaria, Guatemala............................................................ T. candalarius

4b. Upper jaw larger than the lower jaw; 37 to 38 vertebrae in total; middle rays of the anal fin in females, much more elongated than that of males. Interior basins of Chiapas, Mexico, and Guatemala................................................. T. labialis

5a. Scales on lateral line usually 30 (29–31); predorsal scales 25 (22–29); origin of the anal fin to the caudal-base, relatively short (25.2–32.6% in SL); pattern on the sides of the body with delineated, faint, vertical marks. Pacific slope, upper Verde River basin, Oaxaca, Mexico (endemic) .......................................... P. oaxacae

5b. More than 30 scales on the lateral line; predorsal scales 19–24 (except in P. adani ); origin of the anal fin to the caudal-base, relatively large (31.2–37.1% in SL) .......................................................................................................................... 6

6a. Less than basal half of caudal fin densely scaled; head width relatively narrow, its mean value is 60.9% in HL (range 44.5–70.8%). Inland basin of Honduras and Guatemala............................................................................................... P. kreiseri

6b. Basal half or more of caudal fin densely scaled; head width relatively large, its mean value varies between 64.3 to 68.7% in HL..................................................... 7

7a. Predorsal scales usually 26 (23–28); pronounced snout, lower jaw strongly forward projected; relatively large upper jaw (21.1–36.6% in HL). Atlantic slope, upper Papaloapan River basin, Oaxaca, Mexico (endemic) ................ P. adani

7b. Less than 25 predorsal scales; snout is not pronounced, lower jaw is not strongly forward projected........................................................................................ 8

8a. Long epiotic processes, surpassing the first epipleural ribs; robust and deep body (26.3–33.8% in SL). Pacific slope, Huatulco River, Oaxaca, Mexico (endemic) ....................................................................................... P. parentiae

8b. Short epiotic processes, not reaching the first epipleural ribs; slender and deep body (21–31.5% in SL (except in P. punctatus )..........................................................9

9a. Body of the adult fish with brown spots on the scales, aligned in such a way that they form a longitudinal band on the sides of the body; humeral spot evident ......................................................................................................................10

9b. Body of the adult fish without brown spots on the scales on the sides of the body; humeral spot inconspicuous ......................................................................... 11

10a. Third neural spine of the caudal complex much thinner than the second; base of the narrow anal fin less than the length of the caudal peduncle. Pacific slope, from the Isthmus of Tehuantepec, Mexico, to Champerico, Guatemala and Atlantic slope, upper Grijalva River basin, Chiapas, Mexico.......................................................................................... P. punctatus

10b. Third neural spine of the caudal complex, as wide as the second; anal-base fin equal to or greater than the length of the caudal peduncle. Atlantic slope, upper basin of the Coatzacoalcos river (Chimalapas region), Oaxaca, Mexico (endemic) ................................................................................ P. chimalapensis

11a. Head typically rectangular and more arched, its dorsal surface generally very rounded; dark band on the sides of the body extending from the vertical of the dorsal fin to the base of the caudal; 4 to 5 preorbital pores. Interior basin of Guatemala and El Salvador, Pacific slope ...................................... P. guatemalensis

11b. Head conical, its dorsal surface concave to nearly flat; without dark band on the sides of the body; four preorbital pores. Interior basins of Guerrero and Oaxaca, Mexico. Pacific Slope ................................................................................12

12a. Relatively long upper jaw (24.3–35.9% in HL); 31 to 34 scales on the lateral midline. Pacific slope, upper basin of the Mixteco river, Oaxaca, Mexico (endemic) ................................................................................. P. mixtlanensis

12b. Relatively short upper jaw (11.1–23.9% in HL); scales on the lateral midline invariably 33. Pacific slope, Papagayo river basin, Guerrero, Mexico (endemic) ........................................................................................ P. balsanus

Material examined. Profundulus adani : Mexico: CNPE-IBUNAM 23796, holotype, 64.7 mm SL, Santa Maria Ixclatlan, Oaxaca, 17°51’03”N 97°11’57.4”W, 19 Jun 2018, E. Velazquez & M. J. Anzueto. CNPE-IBUNAM 23797, paratypes, 3, 39.0– 59.1 mm SL, Santa Maria Ixcatlan, Oaxaca, 17°51’03”N 97°11’57.4”W, 19 Jun 2018, E. Velazquez & M. J. Anzueto. CNPE-IBUNAM 23798, paratypes, 3, 42.7–48.5 mm SL, Arroyo Nodon, San Miguel Huatla, Oaxaca, 17°47’59”N 97°08’11.9”W, 20 Jun 2018, E. Velazquez & M. J. Anzueto. FMNH 145002, paratypes, 3, 43.1–58.9 mm SL, Santa Maria Ixcatlan, Oaxaca, 17°51’03”N 97°11’57.4”W, 19 Jun 2018, E. Velazquez & M. J. Anzueto. FMNH 145003, paratypes, 3, 67.8–75.0 mm SL, Rio Huatla, San Miguel Huatla, Oaxaca, 17°44’28”N 97°20’15.4”W, 20 Jun 2018, E. Velazquez & M. J. Anzueto. MZ-UNICACH 7382, paratypes, 44 (3 c&s), 36.6–68.2 mm SL, Santa Maria Ixcatlan, Oaxaca, 17°47’57.59”N 97°08’10.40”W, 19 Jun 2018, E. Velazquez & M. J. Anzueto. MZ-UNICACH 7342, paratypes, 18 (2 c&s), 38.3–60.8 mm SL, Arroyo Nodon, San Miguel Huatla, Oaxaca, 17°47’59”N 97°08’11.9”W, 20 Jun 2018, E. Velazquez & M. J. Anzueto. MZ-UNICACH 7343, paratypes, 11, 52.3–77.9 mm SL, Rio Huatla, San Miguel Huatla, Oaxaca, 17°44’28”N 97°20’15.4”W, 20 Jun 2018, E. Velazquez & M. J. Anzueto. Profundulus balsanus : Mexico: MZ-UNICACH 6712, topotypes, 22 (7 c&s), 34.2–64.6 mm SL, Rio Malinaltepec, Guerrero, 17°13’38.03”N 98°40’6.31”W, 26 Apr 2014, M. J. Anzueto. Profundulus chimalapensis : Mexico: MZ-UNICACH 7559, 38 (7 c&s), 35.8–80.3 mm SL, Arroyo Piedra del Rio el Corte, Santa Maria Chimalapa, Oaxaca, 20 Jan 2022, E. Velazquez & M. J. Anzueto. MZ-UNICACH 7560, topotype, 1, 35.1 mm SL, Arroyo la Aurora, Santo Domingo Petapa, Oaxaca, 16°50’17.9”N 95°08’59.8”W, 17 Oct 2021, E. Velazquez & S. Dominguez. Profundulus guatemalensis : Guatemala: LSUMZ 17906, 1 c&s, 52.2 mm SL, Rio las Cabezas, Sanarate, 20 Nov 2014, W. Matamoros. MZ-UNICACH 7194, 12 (4 c&s), 38.3–83.8 mm SL, Balneario el cantil, Rio Cantil, Escuintla, 10 Apr 2017, W. Matamoros & A. E. Gonzalez. MZ-UNICACH 7195, 9 (2 c&s), 38.7–50.3 mm SL, Caserío las Cañas, Arroyo tributario del rio Achihuate, Escuintla, 10 Apr 2017, M. Matamoros & A. E. Gonzalez. MZ-UNICACH 7197, 2, 41.6–42.0 mm SL, Puente la Concha, Arroyo la Concha, Jocotillo Villa Canales, 11 Apr 2017, M. Matamoros & A. E. Gonzalez. MZ-UNICACH 7198, 3, 39.4–57.3 mm SL, Las Cabezas, Puente de las Marias, Santa Rosa Oratoria, 11 Apr 2017, M. Matamoros & A. E. Gonzalez. Profundulus kreiseri : El Salvador: LSUMZ 15105, 2 (1 c&s), 41.57 mm SL, Quebrada Casco de la Hacienda, Parque Nacional Monte Cristo, Santa Ana, 7 Jun 2011, W. Matamoros. MZ-UNICACH 7205, 26 (3 c&s), 37.1–72.6 mm SL, Quebrada los Tecomates, tributario del rio Lempa, Chalatenango, 12 Apr 2017, W. Matamoros & A. E. Gonzalez. MZ-UNICACH 7214, 22 (2 c&s), 38.6–63.3 mm SL, Rio Onuapa, tributario del rio Lempa, Chalatenango, 12 Apr 2017, W. Matamoros & A. E. Gonzalez. MZ-UNICACH 7231, 5, 39.1– 4.19 mm SL, Desembocadura del rio Negro, Perquín, 12 Apr 2017, W. Matamoros & A. E. Gonzalez. Profundulus mixtlanensis : Mexico: CIDOAX-300-F1228, 2, 39.0– 41.1 mm SL, Yosondúa, Tlaxiaco, Oaxaca, 14 Jun 2014, E. Martinez. CIDOAX-301-F1229, 2, 45.6–49.2 mm SL, Ojo de Agua, Yutanicani, Oaxaca, 14 Jun 2014, E. Martinez. MZ-UNICACH 6716, 26 (9 c&s), 39.3–60.7 mm SL, Ranchería la Soledad, Rio San Isidro, Putla, Oaxaca, 29 Apr 2015, M. J. Anzueto & E. Velazquez. MZ-UNICACH 6717, 3, 53.6– 65.5 mm SL, Arroyo tributario del Río Atoyac, colonia Nuevo Tenochtitlan, Putla, Oaxaca, 29 Apr 2015, M. J. Anzueto & E. Velazquez. MZ-UNICACH P 6718, 4, 39.5–53.8 mm SL, Puente Tierra Azul, Putla, Oaxaca, 29 Apr 2015, M. J. Anzueto & E. Velazquez. Profundulus oaxacae : Mexico: MZ-UNICACH 6714, 27 (4 c&s), 35.0– 63.1 mm SL, Rio Salado, subcuenca Atoyac, Mitla, Oaxaca, 27 Apr 2015, M.J. Anzueto. MZ-UNICACH 6715, 22 (2 c&s), 39.2–70.5 mm SL, Arroyo Grande tributario del Río la Hormiga, Mitla, Oaxaca, 27 Apr 2015, M. J. Anzueto. Profundulus parentiae : Mexico: CNPE-IBUNAM 28804, holotype, 65.9 mm SL, Rio Huatulco, Santa Maria Huatulco, Oaxaca, 15°50’24.56”N 96°19’48.75”W, 25 Apr 2015, W. Matamoros, M. J. Anzueto, J. A. Jamangape & I. Aguilar. CNPE-IBUNAM 22805, paratypes, 3, 26.9–35.3 mm SL, Rio Huatulco, Santa Maria Huatulco, Oaxaca, 15°50’24.56”N 96°19’48.75”W, 25 Apr 2015, W. Matamoros, M. J. Anzueto, J. A. Jamangape & I. Aguilar. FMNH 131755, paratypes, 2, 26.9–35.3 mm SL, Rio Huatulco, Santa María Huatulco, Oaxaca, 15°50’24.56”N 96°19’48.75”W, 25 Apr 2015, W. Matamoros, M. J. Anzueto, J. A. Jamangape & I. Aguilar. MZ-UNICACH 6574, 6, 27.5–50.2 mm SL, Puente Hondura de Toro, Rio Huatulco, Santa María Huatulco, Oaxaca, 21 Apr 2015, W. Matamoros, M. J. Anzueto, J. A. Jamangape & I. Aguilar. MZ-UNICACH 6575, 10 (5 c&s), 50.3–68.3 mm SL, Toma de Agua, Rio Huatulco, Santa María Huatulco, Oaxaca, 25 Apr 2015, W. Matamoros, M. J.Anzueto, J. A.Jamangape & I. Aguilar. MZ-UNICACH 6576, paratypes, 10 (3 c&s), 40.2–67.7 mm SL, Arroyo de la calle central, colonia San Miguel Figueroa, Pochutla, Oaxaca, 15°47’24.1”N 96°24’03.9”W, 21 Apr 2015, W. Matamoros, M. J. Anzueto, J. A. Jamangape & I. Aguilar. MZ-UNICACH 6713, paratypes, 10 (4 c&s), 42.5–53.2 mm SL, Camino a Comitlan, Rio la Loma, Santa Maria Colotepec, Oaxaca, 16°01’08.1”N 96°50’41.3”W, 22 Apr 2015, W. Matamoros, M. J. Anzueto, J. A. Jamangape & I. Aguilar MZ-UNICACH 6721, 2 c&s, 44.8– 39.3 mm SL, Rio del Rancho Flor de Café, San Gabriel Mixtepec, Oaxaca, 30 Apr 2015, M. J. Anzueto. MZ-UNICACH 7351, 6, 46.3–73.9 mm SL, Puente Mateo Piña, camino a Huatulco, San Mateo Piña, Oaxaca, 22 Sep 2018, E. Velazquez & M. J. Anzueto. Profundulus punctatus : Guatemala: MZ-UNICACH 7169, 32, 37.9–72.6 mm SL, Rio Batza, Colonia Campolla, puente Batza, Quetzaltenango, 9 Apr 2017, A. E. Gomez & E. Velazquez. MZ-UNICACH 7182, 16, 36.3–75.0 mm SL, Rio Nil, puente Nil, El Acintal, Retalhuleo, 9 Apr 2017, A. E. Gomez & E. Velazquez. MZ-UNICACH 7186, 14, 39.4–82.1 mm SL, Rio el Puentecito, Caserio, Retalhuleo, 9 Apr 2017, A. E. Gomez & E. Velazquez. Mexico: MZ-UNICACH 3100, 9, 36.8–57.0 mm SL, Cortina Copainala, Rio Grijalva, Chicoasen , Chiapas, 23 Apr 2009, E. Velazquez. MZ-UNICACH 4332, 6, 38.4-53.6 mm SL, El Chorreadero, Rio Grijalva, Chiapa de Corzo, Chiapas, 23 Oct 2010, E. Velazquez. MZ-UNICACH 4534, 14, 36.7–58.5 mm SL, El Chorreadero, Rio Grijalva, Chiapa de Corzo, Chiapas, 11 May 2011, M. J. Anzueto & E. Velazquez. UNICACH 5067, 3, 36.1–39.3 mm SL, Arroyo Rancho Mujular, Chiapa de Corzo, Chiapas, 12 Apr 2014, M. J. Anzueto & E. Velazquez. MZ-UNICACH 5074, 3, 37.1–46.7 mm SL, Rio Nandalumi, Chiapa de Corzo, Chiapas, 12 Apr 2014, M. J. Anzueto & E. Velazquez. MZ-UNICACH 5143, 4, 36.7–52.7 mm SL, Rio Nandalumi, Chiapa de Corzo, Chiapas, 28 May 2014, M. J. Anzueto & E. Velazquez. MZ-UNICACH 5085, 9, 36.7–52.7 mm SL, Arroyo Rancho Mujular, Chiapa de Corzo, Chiapas, 28 May 2014, M. J. Anzueto & E. Velazquez. MZ-UNICACH 5752, 4, 40.5–43.5 mm SL, Arroyo San Jose, Rio Grijalva, Villaflores, Chiapas, 26 Dec 2014, M. J. Anzueto. MZ-UNICACH 6326, 25, 37.9–72.6 mm SL, Rio Lagartero, Arriaga, Chiapas, 7 Apr 2015, A. E. Gomez & E. Velazquez. MZ-UNICACH 6355, 2 c&s, 67.3– 62.7 mm SL, Rio Zapata, Cacahotan, Chiapas, 9 Apr 2015, A. E. Gomez. MZ-UNICACH 6578, 18 (2 c&s), 45.9–56.3 mm SL, Cascada Velo de Novia, Tonala, Chiapas, 8 Apr 2015, A. E. Gomez & E. Velazquez. MZ-UNICACH 6586, 17, 37.6–58.4 mm SL. Puente Rancho Bonito, Jiquipilas, Chiapas, 7 Apr 2015, A. E. Gomez & E. Velazquez. MZ-UNICACH 6632, 21 (3 c&s), 43.3–72.0 mm SL, Rio Cintalapa, Escuintla, Chiapas, 9 Apr 2015, A. E. Gomez & E. Velazquez. Tlaloc candalarius : Guatemala: MZ-UNICACH 7566, topotypes, 18, 39.9–50.9 mm SL, Arroyo del Cenote, La Candelaria, Nenton, 15°56’59.9”N 91°46’53.6”W, 19 Dec 2019, E. Velazquez & M. J. Anzueto. Mexico: MZ-UNICACH 3898, 4, 39.5–40.8 mm SL, Arroyo Teopisca, Teopisca, Chiapas, 8 Aug 2008, E. Velazquez. MZ-UNICACH 3899, 8 (2 c&s), 39.5–99.2 mm SL, Arroyo Teopisca, Teopisca, Chiapas, 6 Oct 2008, E. Velazquez. MZ-UNICACH 5705, 7 (2 c&s), 41.3–45.6 mm SL, Arroyo Santa Barbara, Teopisca, Chiapas, 14 Dec 2014, A. E. Gomez & E. Velazquez. MZ-UNICACH 5709, 1 c&s, 50.3 mm SL, Rio Grande, Comitan, Chiapas, 14 Dec 2014, A. E. Gomez & E. Velazquez. MZ-UNICACH 6224, 1, 54.93 mm SL, Arroyo El Mangal, Socoltenango, Chiapas, 25 Jan 2015, E. Velazquez.MZ-UNICACH 6813, 7 (4 c&s), 39.6–72.1 mm SL, Laguna Chukumaltik, Comitan, Chiapas, 22 Jul 2015, M. J. Anzueto & E. Velazquez. MZ-UNICACH 7245, 5, 26.5–65.4 mm SL, Lagos de Colon, La Trinitaria, Chiapas, 21 Oug 2015, A. E. Gomez & E. Velazquez. MZ-UNICACH 7565, 52 (2 c&s), 38.8–86.4 mm SL, Arroyo Ojo de Agua, Tzimol, Chiapas, 15 Feb 2021, M. J. Anzueto & E. Velazquez. MZ-UNICACH 7567, 6, 53.1–79.3 mm SL, Arroyo Uninajab, Comitan, Chiapas, 6 Feb 2020, E. Velazquez & M. J. Anzueto. Tlaloc hildebrandi : Mexico: MZ-UNICACH 7567, 10 (2 c&s), 42.2–89.2 mm SL, Rio Amarillo, San Cristobal de las Casas, Chiapas, 6 Jul 2016, M. J. Anzueto & E. Velazquez. MZ-UNICACH 4330, 4 (3 c&s), 67.7–111.4 mm SL, Rio Fogotico, San Cristobal de las Casas, Chiapas, 4 May 2006, E. Velazquez. MZ-UNICACH 5694, 10, 47.9– 63.1 mm SL, Puente Blanco, San Juan Chamula, Chiapas, 11 Dec 2014, A. E. Gomez & E. Velazquez. MZ-UNICACH 7562, 26, 37.9–63.1 mm SL, Rio San Pedro, Chenalho, Chiapas, 1 Sep 2016, M. J. Anzueto. MZ-UNICACH 5711, 9 (2 c&s), 38.1–104.0 mm SL, Rio Osilhucum, Chenalho, Chiapas, 22 Sep 2007, E. Velazquez. MZ-UNICACH 6194, 9 (1 c&s), 49.5–79.7 mm SL, Arroyo las Limas, Chenalho, Chiapas, 7 Mar 2015, E. Velazquez. Tlaloc labialis : Guatemala: MZ-UNICACH 7233, 2 c&s, 35.5–65.6 mm SL. Rio Selegua, Huehuetan, 20 Apr 2017, W. Matamoros. Mexico: MZ-UNICACH 1402, 1, 64.1 mm SL, Rio Negro, Pueblo Viejo, Cintalapa, Chiapas, 11 Oct 2007, E. Velazquez & M. J. Anzueto. MZ-UNICACH 1305, 16, 41.5–79.7 mm SL, Rio Negro Santa Maria, Cintalapa, Chiapas, 12 Jul 2007, M. J. Anzueto & A. E. Gomez. MZ-UNICACH 3467, 16 (2 c&s), 55.5–73.5 mm SL, Rio Zacalapa, Copainala, Chiapas, 6 Aug 2009, M. J. Anzueto & E. Velazquez. MZ-UNICACH 5274, 1, 67.0 mm SL, Rio Frio, Chiapilla, Chiapas, 1 Oct 2014, E. Velazquez. MZ-UNICACH 5353, 1 c&s, 89.4 mm SL, Rio Ojo de Agua, Emiliano Zapata, Chiapas, 2 Oct 2014, M. J. Anzueto. MZ-UNICACH 6607, 7, 38.9–101.9 mm SL, Rio Mazantic, El Bosque, Chiapas, 8, Mar 2015, A. E. Gomez. MZ-UNICACH 6740, 2 c&s, 64.5–75.7 mm SL, Rio La Venta, Ocozocoautla, Chiapas, 23 Apr 2015, M. J. Anzueto & E. Velazquez. MZ-UNICACH 7563, 26, 33.4–53.1 mm SL, Rio Hondo, Ixtapa, Chiapas, 20 Sep 2019, M. J. Anzueto & E. Velazquez. Tlaloc portillorum : Honduras: LSUMZ 31597, topotypes, 14 (3 c&s), 49.8–70.7 mm SL, Rio Calan, Quebrada Potrerillos, Comayagua, 14°32’31”N 87°52’55”W, 5 Dec 2007, W. Matamoros. MZ-UNICACH 7220, 21 (4 c&s), 40.8–62.9 mm SL, Puente de Piedra a Lepaterique, Francisco Morazan, 15 Apr 2017, W. Matamoros & A. E. Gomenz. MZ-UNICACH 7222, 28 (2 c&s), 37.5–71.3 mm SL, Rio Nacaome, Las Tablas, Francisco Morazan, 15 Apr 2017, W. Matamoros & A. E. Gomez.