Pseudophallus mindii ( Meek & Hildebrand 1923 )

Dallevo-Gomes, Caio I. A., Mattox, George M. T. & Toledo-Piza, Mônica, 2020, Taxonomic review of the pipefish genus Pseudophallus Herald, with the description of a new species (Syngnathiformes: Syngnathidae), Zootaxa 4859 (1), pp. 81-112 : 96-98

Pseudophallus mindii ( Meek & Hildebrand 1923)

( Figs. 2–4 , 6–7 ; Tables 1 , 3–6 )

Syngnathus mindii Meek & Hildebrand, 1923: 261 , pl. 18, fig. 2 (original description, “creek near Mindi, Canal Zone, Panama ”, holotype USNM 81770); Jordan et al., 1930: 242 (listed in Central America, synonym list of P. mindii ); Fowler, 1944: 461, 496 (listed in Central America); Dawson, 1982: 49 (synonym list of P. mindii ); Ferraris Jr, 2003: 591 (synonym list of P. mindii ).

Pseudophallus mindii: Herald, 1942: 132 (new combination, identification key, distribution); Schultz, 1949: 100, 101 (identification key, Río Sanchón and Río Cumboto, Venezuela); Caldwell et al., 1959: 18 (listed; Tortuguero, Costa Rica); Boeseman, 1960: 123 (presumed occurrence in Trinidad); Follett, 1960: 224 (affinities with congeners); Herald, 1965: 364 (listed in the Atlantic coast, freshwater pipefish); Miller, 1966: 797 (distribution in Central America); Mago-Leccia, 1970: 92 (listed, Venezuela); Gilbert & Kelso, 1971:18, 29 (listed in Tortuguero, Costa Rica); Dahl, 1971: 173 (listed: Ríos Sinú and Pechilin, Colombia); Dawson, 1974: 409, 410 (meristic characters; table and identification key); Shino, 1976: 109 (listed, common names of fishes); Dawson, 1982: 5, 23, 45, 47–53, figs. 18, 20 (in part, senior synonym of P. brasiliensis , description, distribution in Central America, comparisons with congeners, identification key); Dawson, 1985: 203 (distribution, biogeography); Bussing, 1987: 164 (listed, Costa Rica); Winemiller & Leslie, 1992: 49 (listed, Tortuguero National Park, Costa Rica); Greenfield & Thomerson, 1997: 136 (listed, Belize); Bussing, 1998: 253, 255–257, figures and map (identification key, species account, distribution in Costa Rica); Fritzsche & Vincent, 2003: 1222 (in part, senior synonym of P. brasiliensis , listed, distribution); Ferraris Jr, 2003: 591 (listed, distribution); Smith et al., 2008: 998, 1000 (listed, Puerto Rico): Kuiter, 2009: 241 (in part, senior synonym of P. brasiliensis , , distribution, freshwater, figure from Dawson 1982); Matamoros et al., 2009: 16, 35 (listed, Honduras); Mol, 2012: 45, 46, 51 (mention of putative taxonomic problem); Page et al., 2013: 114 (listed, Central America); Phillip et al., 2013: 10, 21, 26 (listed, Trinidad, identification key); Van Oosterhout et al., 2015: 697 (listed, Costa Rica).

Pseudophallus mindi (sic): Herald, 1959:473 (distribution in the Atlantic coast); Lülling, 1974:200 (geographic race of P. starksii ); Angulo et al., 2013:998 (listed in Costa Rica, distribution).

Pseudophallus starksi mindi: Lüling, 1974: 200 (as a geographic race of P. starksii ).

Diagnosis. Pseudophallus mindii differs from species of Pseudophallus from the Pacific coastal drainages, P. starksii and P. elcapitanensis , in having 78.9% of specimens with 13 trunk rings (vs. 14 trunk rings).

Pseudophallus mindii can be distinguished from P. brasiliensis by the following combination of characters: 13– 16 pectoral-fin rays (88.2% of specimens with 14–15), and 15–23 brood-pouch rings (93.8% adult male specimens with 15–21) vs. 12–15 pectoral-fin rays (82.4% of specimens with 13), 28–41 dorsal-fin rays (98.5% of specimens with 28–37) and 13–17 brood-pouch rings (87.7% of the adult male specimens with 14–16) in P. brasiliensis .

Pseudophallus mindii differs from P. galadrielae in having 32–42 dorsal-fin rays (96.4% of specimens with 34–42), all specimens with 7.5–10.5 subdorsal rings, 45–51 total body rings (94.4% of specimens with 46–51) vs. 27–34 dorsal-fin rays (91.6% of specimens with 27–34), 6.0–8.25 subdorsal rings (92.9% of specimens with 6.0–7.25) and 41–45 total body rings (97.1% of specimens with 41–44) in P. galadrielae .

Description. Meristic and morphometric characters detailed in table 1. General appearance in Figures 2C and 3C . Body without scales, covered with bony plates forming rings along its axis, bony plates horizontally oval, with an elevation in midline forming bony ridges along body. Trunk rings 12–14 (80.3% of specimens with 13). Tail rings 32–37 (90.7% of specimens with 34–35). Body rings 45–51 (85.1% of specimens with 46–48). Subdorsal rings 7.5–10.25 (93.0% of specimens with 7.75–9.75). Subdorsal trunk rings 0–0.25. Subdorsal tail rings, from 7.5–10.25 (91.0% of specimens with 7.75–9.75). Dorsal-fin origin between posterior half of posteriormost trunk ring and anteriormost caudal ring. Posterior tip of superior trunk ridge at vertical through superior tail ridge, between 7.0 and 10.0 anteriormost tail rings. Lateral trunk ridge continuous with superior tail ridge with origin on first caudal ring. Inferior trunk ridge continuous with inferior tail ridge.

Body slender and cylindrical, transversal section of trunk heptagonal, with transition to quadrangular in tail at end of superior trunk ridge. Maximum body depth of females and juveniles at middle of trunk; in males at anterior region of the brood pouch. Body depth decreasing posteriorly along superior trunk ridge to minimum depth on caudal peduncle. Dorsal profile of head straight or slightly inclined at snout region, more inclined at orbital region and straight at posterior region, with two nuchal plates between occipital region and first trunk ring. Ventral profile of head straight.

Dorsal fin not elevated, curved on first rays, straight along median portion with pointed end. Dorsal-fin rays of similar length, except the first two rays. Dorsal-fin rays 32–42 (76.7% of specimens with 35–38), posteriormost two rays closer to each other, difficult to distinguish in some specimens. Pectoral fin small, middle rays longer than outer and inner rays. Pectoral-fin origin on anteriormost trunk ring, and distal margin rarely extending beyond third trunk ring. Pectoral fin rays 13–16 (88.2% of specimens with 14 –15). Caudal fin small and fan like, with 10 rays (5 rays in each hypural plate). Pelvic and anal fins absent.

Mouth terminal, slightly inclined dorsally, jaws edentulous. Lower jaw slightly prognathous. Snout short, tubular, without dorsal or lateral ridges. Dorsal profile of snout with slight inclination following dorsal margin of eyes. Eyes located longitudinally at middle of head. Opercle occupying most of lateral surface of head posterior to eye, opercular opening reduced to pore on posterodorsal margin of opercle. Opercular ridge generally indistinct, except for some specimens with small protuberance on anterior region of opercle, following superior margin of bone. Lateral line and lateral line canals absent.

Sexual dimorphism. Males with brood pouch. Origin of brood pouch on posteriormost trunk rings, brood pouch rings extending along 15–23 body rings (93.3% of specimens with 15–21 brood pouch rings). Bony plates of the brood pouch well developed and T-shaped encasing entire lateral portion of pouch, decreasing in size posteriorly and parallel with the inferior tail ridge. Ventral region of pouch with contralateral dermal folds extending towards the midline, dividing brood pouch longitudinally. Adult females with anal papilla on anal ring with small, barely visible spicules.

Color in alcohol. Ground color of head and body usually uniform, from light beige to brown, sometimes with mottled patterns formed by different tones of brown spots. Thicker bony regions at body ridges with less or no chromatophores, forming nearly white longitudinal lines. Some specimens with series of dark arch-shaped blotches in lateral view, at every two or four rings. Bony rings with one or two white spots underneath bony plates; spots circular in shape when single spot present and oval shaped when two spots present, due to proximity of spots. Dorsolateral part of snout with layer of skin and mesethmoid forming light stripe. Dorsal-fin rays with scattered brown chromatophores. Pectoral fin hyaline. Caudal fin with same general coloration of body, sometimes in slightly different tone. ( Figs. 2C and 3C ).

Distribution. Freshwater Atlantic coastal drainages, from Río Cumboto, Ocumare, Venezuela, to Río Hondo, San Antonio, Belize, including freshwater drainages in the Caribbean islands ( British Virgin Islands, Puerto Rico, Jamaica and Cuba) ( Fig. 4 ).

Remarks. The species Syngnathus mindii Meek & Hildebrand was described based on a specimen captured in a creek near Mindi, Canal Zone, Panama (USNM 81770). A tag inside the jar of the specimen has the name Siphostoma mindii , however, to our knowledge, such combination has never been used in the literature.

Type material: Holotype. USNM 81770 , 1 , 90.3 mm SL, male, Creek Mindi, Canal Zone, Panama, S. E. Meek & S. F. Hildebrand, 11 Mar 1911.

Non-types. Venezuela: Aragua: UMMZ 146437, 1, 89.9 mm SL, Río Cumboto , near Ocumare, F. F. Bond, 05 Jan 1938; UMMZ 146438 , 1, 112.3 mm SL, Boca del Río Cumboto , 2km NW of near Ocumare, F. F. Bond, 05 May 1939; UMMZ 146441 , 1 , 95.1 mm SL, Río Sanchon , 5 km W of Tavorda & 10 km W of Puerto Cabello, F. F. Bond, 26 Jan 1938; USNM 123667 , 1, 110.8 mm SL, Río Cumboto near mouth, 2 km northwest of Ocumare , F. F. Bond, 5 May 1939 .

Panama: Kuna Yala: USNM 208359 , 3 , 118.2 – 134.1 mm SL, Río Narka 5 miles upstream, Battelle Nw Lab, 2 Mar 1967 ; USNM 293481 , 2 , 97.4–102.5 mm SL, Río Carti Grande at Carti rd. 3km from Carti, W. C. Starnes et al., 1 Mar 1985 ; Colón: ANSP 104451, 2, 95.4–119.7 mm SL, Río Piedras about 4 mi. above mouth, where small road fords river, H. G. Loftin & C. Lewis, 23 Feb 1962 ; GCRL 4734, 1, 47.9 mm SL, Portobello Road, mouth of creek entering Caribbean , ca 20 km E of Maria Chiquita, C. E. Dawson, 27 Jun 1970 ; GCRL 12775, 2, 79.1–112.2 mm SL, Río Guanche , ca 1-1.5 km upstream from mouth, L. G. Abele et al., 17 Apr 1974 ; LSUMZ 15741, 1, 62.1 mm SL, upstream off road to Errebache Embera from Portobello , 09º26’17’’N 79º43’ 50’’W, C. McMahan, W. Matamoros & M. Davis, 05 Apr 2011 .

Costa Rica: Limón: TNHC 14060, 1, 66.4 mm SL, Caño Agua Fria Viejo, tributary of Río Tortuguero ca 0.6 km from village, P. N. Tortuguero , 10º30’N 83º30’W, K. Winemiller, 12 Apr 1985 ; TNHC 14134, 1, 79.0 mm SL, Caño Agua Fria Viejo, tributary of Río Tortuguero ca 0.6 km from village, P. N. Tortuguero , 10º50’N 83º50’W, K. Winemiller, 23 Jun 1985 ; TNHC 14170, 4, 101.8– 110.5 mm SL, Caño Agua Fria Viejo, tributary of Río Tortuguero ca 0.6 km from village, P. N. Tortuguero , 10º30’N 83º30’W, K. Winemiller, 21 Aug 1985 ; TNHC 14252, 1, 100.3 mm SL Caño Agua Fria Viejo, tributary of Río Tortuguero ca. 0.6 km from village, P. N. Tortuguero , 10º30’N 83º30’W, K. Winemiller, 18 Nov 1985; TU 189786 , 2, 68.5–96.6 mm SL, Río Blanco , West of Puerto Limon, 1 Apr 1974 ; UF 7179, 3, 59.5–83.9 mm SL, small creek 1/ 2 mile from Tortuguero Inlet, 10º58’N 83º52’W, L. Ogren, 19 Aug 1958 ; UF 11167, 1, 84.9 mm SL, Río Palacio , ca. 2 mi. above mouth ca. 5 mi. NNW of Tortuguero, C. Gilbert & D. P. Kelso, 23 Aug 1963 ; UF 11242, 1, 92.9 mm SL, Río Agua Fria , ca. 4 mi. upstream from Tortuguero Village, 10º52’N 83º56’W, C. Gilbert et al., , 26 Aug 1963 ; UF 16356, 1, 72.9 mm SL, Tortuguero lagoon, westside, sidewater area, about 100 yd below mouth, C. R. Gilbert et al., 29 Aug 1963 ; Alajuela: ANSP 163716, 1, 87.7 mm SL, stream by road 5.0 km SE of Upala , km 85, ruta 4, D. Fromm & P. Fromm, 12 Mar 1988 .

Nicaragua: North Atlantic: GCRL 15645, 2, 61.8–65.8 mm SL, Río Prinzapolka , near junction of Ríos UIi and Wani, ca 150 from Atlantic Caribbean, Zelaya, A. Cajina, 6 Apr 1977 ; LSUMZ 15346, 1, damaged, La Mosquitia at Río Prinzapolka , 500m S of mouth from canal, 13º24’14’’N 83º36’40’’W, C. McMahan, W. Matamoros, P. House & M. Palacios, 05 Aug 2011; UF 228493 , 1, 63.6 mm SL, Chihuahua creek, near Río Mico , L. R. Riva, 30 Jun 1980 .

Honduras: Atlantida: LACM 32462.11, 1, 113.8 mm SL, Río Cangrejal 4.8 km southeast of Barrio La Isla , La Ceiba, M. Martin, 16 Mar 1970 ; LSUMZ 17598, 1, 94.3 mm SL, Laguna Bacalar, J. C. Carrasco, 26 Nov 2010; UMMZ 173353, 1, 57.4 mm SL, 48.5 km West of Ceiba , near San Juan Benque, Gordon, Chable & George, 11 Apr 1951 .

Guatemala: Izabal: AMNH 35246, 1, 152.2 mm SL, Río Montagua at Finca Hopi , D. E. Rosen, J. M. Bailey, R. Vari & L. C. Stuart, 17 Apr 1974; UMMZ 146068 , 1 , 63.4 mm SL, Río Boca Nueva, trib. to Río Polochic at RR bridge, ca 3.5 km SW of Panzos, Alta Verapaz, L. C. Stuart, 1 Mar 1940; UMMZ 197349 , 2 , 135.9 – 157.1 mm SL, Motagua River at Finca Hopi , 17 km by map above mouth, Izabal, D. E. Rosen, J. M. Bailey, R. Vari & L. C. Stuart, 17 Apr 1974; USNM 114345 , 1 , 41.1 mm SL, Jicotea creek tributary of Río Sarstoon , 1/ 4 mile above mouth, R. R. Miller, A. D. Holloway & J. Midence, 30 Apr 1947 .

Belize: Belize: FMNH 103802, 7, 98.5–120.5 mm SL, Hector creek , Hattieville, N. Aspinwall, Mar 1970 ; UMMZ 202904, 1, 71.9 mm SL, Hector Creek , Sibun River drainage, M. Gordon & Fairweather, 23 Mar 1949 ;

Orange Walk: FMNH 82329, 1, 111.1 mm SL, Rio Hondo, 7 mi. W of Orange Walk, San Antonio, Greenfield , Carter , Anderson , 20 Dec 1975 .

Puerto Rico: East: GCRL 16070, 1, 95.0 mm SL, Espirito Santo river estuary, east of San Juan , UPR, 7 Sep 1977 ; West: ANSP 144683, 1, 111.5 mm SL, Mayaguez, no collector specified, 1939.

British Virgin Islands: USNM 108413 , 1 , 88.3 mm SL, largest specimen damaged, Leeward Isles , 18º39’30’’N 64º5130’’W, Johnson-Smithsonian Deep-Sea Expedition, 4 Mar 1933 .

Jamaica: Saint James: USNM 78138 , 1 , 88.3 mm SL, Montego River , C. B. Wilson, 1910; Saint Mary: ANSP 102022, 1, 100.8 mm SL, Fort George, Pencar river, 2 miles S. of Annotto Bay , C. W. Hart & G. Thomas, 4 Aug 1959 .

Cuba: Guantánamo: UF 10376, 1, 81.4 mm CP, Yateras River , Station # 2, 2.25 mi. upstream from mouth, Oriente, W. McLane et al., 19 Sep 1952 .

Figures

FIGURE 2: Species of the genus Pseudophallus: A—Pseudophallus brasiliensis (paratype) USNM 212058, 69.1 mm SL, Igarapé Inó, Furo de Panaquera, Rio Tocantins, Pará, Brazil. B—Pseudophallus elcapitanensis MCZ 88623, 110.6 mm SL, Río Picuro, Darien, Panama. C—Pseudophallus mindii (holotype) USNM 81770, 90.3 mm SL, Mindi Creek, Canal Zone, Panama. D—Pseudophallus starksii USNM 208365, 171.1 mm SL, upper tributary of Río Jaque, Província de Darien, Panama. E—Pseudophallus galadrielae. (holotype) FMNH 126156, 63.1mm SL, Lago Izabal, Guatemala. Red arrow indicates the anal papilla.

FIGURE 3: Detail of anterior portion of body of species of the genus Pseudophallus A—Pseudophallus brasiliensis (paratype) USNM 212058, 69.1 mm SL, Igarapé Inó Furo de Panaquera, Rio Tocantins, Pará, Brazil. B—Pseudophallus elcapitanensis MCZ 88623, 110.6 mm SL, Río Picuro, Darien, Panama. C—Pseudophallus mindii (holotype) USNM 81770, 90.3 mm SL, Mindi Creek, Canal Zone, Panama. D—Pseudophallus starksii USNM 208365, 171.1 mm SL, upper tributary of Río Jaque, Província de Darien, Panama. E—Pseudophallus galadrielae. (holotype), FMNH 126156, 63.1 mm SL, Lago Izabal, Guatemala.

FIGURE 4: Distribution map of species of Pseudophallus occurring in drainages along the Atlantic coast, based on specimens examined in this study.

FIGURE 6. Scatterplot of scores of linear discriminant analysis for morphometric and meristic characters of the three species of Pseudophallus occurring in Atlantic drainages. LD1: Linear discriminant 1, LD2: Linear discriminant 2, A: Standard length, B: Head length, C: Snout Length, D: Snout depth, E: Pectoral-fin length, F: Pectoral-fin base length, G: Tail length, H: Dorsal-fin base length, I: Anal ring depth, J: Body depth, K: Head depth, L: Body width, M: Trunk depth, N: Orbital diameter, O: Snout width, P: Dorsal-fin rays, Q: Pectoral-fin rays, R: Trunk rings, S: Caudal rings, T: Subdorsal trunk rings, U: Subdorsal caudal rings, V: Origin of dorsal fin.

FIGURE 7. Maps of distribution of specimens of Pseudophallus available for study. Left side: distribution of specimens examined for the morphological analysis in this study. Right side map: distribution of voucher specimens with genetic sequences on GenBank (as of April 2020).