Pseudophallus elcapitanensis (Meek & Hildebrand 1914)

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 : 91-94

Pseudophallus elcapitanensis (Meek & Hildebrand 1914)

( Figs. 2–3 , 5 , 7 ; Table 2)

Siphostoma auliscus (non Swain): Gilbert & Starks, 1904: 57 (Rio Grande, Miraflores, near Panama; comparison with specimens from Magdalena Bay).

Syngnathus auliscus (non Swain): Regan, 1908: 53 (brief description).

Siphostoma elcapitanense Meek & Hildebrand in Meek, 1914: 119 (original description; type locality: “El Capitan, Panama ”); Grey, 1947: 187 (type specimens, listed); Dawson, 1982: 46, 53 (as a synonymy of P. elcapitanensis ); Ibarra & Stewart, 1987:80 (listed as paratypes at FMNH, see remarks).

Syngnathus elcapitanense: Meek & Hildebrand, 1923: 260 , pl. 19 (new combination; description; distribution in Panama and Costa Rica, USNM 81735 indicated as type); Breder, 1927: 136, 144, 148, 159, 165 (in key, euryhaline, behavior, listed in Río Chico, Panama); Jordan et al., 1930: 242 (listed; West Coast of Panama, synonymic list); Fowler, 1944 a: 496 (listed; distribution in Central America); Bussing, 1967: 236 (listed, Costa Rica); Whatley, 1969: 448 (cited as euryhaline fish, misspelled); Dawson, 1982: 46, 53 (synonymic list of P. elcapitanensis ).

Syngnathus elcapitanensis: Herald, 1940: 61 , 63 (identification key; Panama); Gunter, 1956: 349 (listed; Central America); Miller, 1966: 796 (listed; Pacific slopes of Costa Rica and Panama).

Syngnathus (Syngnathus) elcapitanensis: Dawson, 1982: 53 (synonymic list of P. elcapitanensis ).

Pseudophallus elcapitanensis: Herald, 1959: 473 (new combination; brief mention to brood-pouch morphology; occurrence in the Pacific coast); Follett, 1960: 224 (affinities with congeners); Dawson, 1974: 409, 410 (meristic characters in table and identification key); Lüling, 1974:201 (reference); Dawson, 1982: 5, 23, 45, 47–55, figs. 19, 20 (redescription, distribution, comparisons with congeners, identification key); Dawson, 1985: 161, 209, figs. 239, 241 (characters, distribution, comparisons with congeners, identification key); Lyons & Schneider, 1990: 4, 6, 7, 8 (listed; Costa Rica, biology, ecology); Bussing, 1998: 252–254, figures and map (identification key, species account, distribution in Costa Rica); Ferraris Jr, 2003: 591 (listed, distribution; USNM 81735 indicated as lectotype of species); Vega et al., 2006: 92 (listed; Panama); Kuiter, 2009: 241 (distribution, figure from Dawson, 1982); Angulo et al., 2013: 998 (listed, distribution; Costa Rica); Angulo et al., 2015: 7, 9, 10; fig. 5J (listed, distribution in Costa Rica, identification key); Angulo et al., 2020: 9, 18, 19, 25; fig. 8M (listed in Costa Rica, identification key).

Diagnosis. Pseudophallus elcapitanensis differs from species of Pseudophallus from the Atlantic coastal drainages, P. brasiliensis , P. mindii and P. galadrielae in having all of the specimens with 14 trunk rings (vs. most usually with 13 trunk rings).

Pseudophallus elcapitanensis differs from P. starksii in the following combination of characters: 30–35 dorsalfin rays (87.2% of specimens with 31–33), 47-51 body rings (89.7% of specimens with 48–49), 17–19 brood-pouch rings (90.9% of specimens with 17–18) and 6.75–7.5 subdorsal tail rings vs. 36–44 dorsal-fin rays (89.5% of specimens with 38–41), 49–53 body rings (91.9% of specimens with 50–51), 19–21 brood-pouch rings (64.9% of adult male specimens with 21) and 8.5–10.5 subdorsal tail rings (86.7% of specimens with 9.25–10.0).

Description. Meristic and morphometric characters detailed in Table 2. General appearance in Figures 2B and 3B . 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 14. Tail rings 33–37 (89.7% of specimens with 34–35). Body rings 47–51 (89.7% of specimens with 48–49). Subdorsal rings 6.75–7.5 (97.4% of specimens with 6.75–7.25). Subdorsal trunk 0–0.25 rings. Subdorsal tail rings 6.75–7.5 (97.4% of specimens with 6.75–7.25). Dorsal-fin origin between 0.25 of posteriormost trunk ring and anteriormost caudal ring. Posterior tip of superior trunk ridge at vertical through superior tail ridge, between 6.0 and 7.5 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 depth of body of females and juveniles at middle of trunk; in males at 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 on snout region, more inclined on orbital region and straight on 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 30–35 (82.1% of specimens with 31–33), 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, distal margin of pectoral-fin rays rarely extending beyond third trunk ring. Pectoral-fin rays 13–16 (66.7% of specimens with 14). 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 17–19 rings (90.9% of specimens with 17–18 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. Head and body ranging from a uniform light to dark brown, with small white punctuations. Dorsal-fin rays with scattered brown chromatophores, pectoral fin hyaline. Caudal fin with the same coloration of the body, sometimes varying in tone ( Figs. 2B and 3B ).

Distribution. Freshwater coastal drainages of the Pacific, from Río Mamoni, Province of El Capitain, Panama, to Río Turrubales, Province of Orotina, Costa Rica ( Fig. 5 ).

Remarks. Meek & Hildebrand (in Meek, 1914) provided a brief description of Siphostoma elcapitanense (sic) stating that the species would be further detailed in an upcoming account of fishes of Panama, a study that came out years later ( Meek & Hildebrand, 1923). In the original description, the type locality was stated as El Capitan, Panama. No catalogue number of lots was mentioned, but two specimens from Costa Rica were listed at the end of the description: one from Jesus Maria and another from Río Turrubales.

The species was described in more detail under a different genus, as Syngnathus elcapitanensis by Meek & Hildebrand (1923: 260), who also listed and illustrated a male specimen, from Río Mamoni , El Capitan, Panama, catalogued as USNM 81735 , as the type specimen. Those authors also mentioned that they had a total of 11 specimens, two males and nine females, from five different localities in Panama and that all, except one, originated in freshwater .

Ten of those specimens are currently deposited in the fish collections of the National Museum of Natural History, Smithsonian Institution (5 specimens, one in each lot of USNM 79687, 79689–90; 81375 and 108645) and the Field Museum of Natural History (5 specimens, one in each lot of FMNH 8290–8294). The last specimen of the 11 mentioned by Meek & Hildebrand (1923) originated in Chame Point, Panama, which according to those authors was not a locality from freshwater. According to Dawson (1982: 54) that is a specimen of Pseudophallus starksii that was misidentified by Meek & Hildebrand (1923).

Associated with each lot from the Field Museum of Natural History are handwritten labels with numbers that correspond to the catalogue numbers of the lots from the National Museum of Natural History as follows: FMNH 8290 (number in label: 79688), FMNH 8291–8292 (number in label: 79690, also marked as cotypes in a label), FMNH 8293 (number in label: 79689) and FMNH 8294 (number in label: 79687).

All the 10 specimens listed above were collected by S. E. Meek and S. F. Hildebrand in March 1911 or February 1912, as part of the expedition Smithsonian Biological Survey of the Panama Canal Zone. Of those, only five specimens: USNM 79690, USNM 81735, USNM 108645, FMNH 8291 and FMNH 8292 were collected at Río Mamoni, El Capitan, the type locality of P. elcapitanensis . In the original description of the species, Meek & Hildebrand (in Meek, 1914) did not expressly indicate a holotype. Later, in addition to citing USNM 81735 as the type of the spe-cies, another specimen was indicated as a paratype ( Meek & Hildebrand, 1923: plate XIX b), which is currently catalogued as USNM 108645 and, according to the institution catalogue records, was removed from the lot USNM 81735. The specimens FMNH 8291 and 8292 seem to have originated from USNM 79690 and were also available to the authors when they established the nominal species (based on information on the labels inside the jars, mentioned above). Therefore, those are the five specimens that constitute the type series of the species (ICZN, 1999: Art. 72.4.1.1 and 73.2). Meek & Hildebrand´s (1923) citation of USNM 81735 as type, would therefore constitute a lectotype designation, an interpretation in line with that of Ferraris Jr. (2003: 591). The additional five specimens from Panama, (USNM 79687, 79689 and FMNH 8290, FMNH 8293 and FMNH 8294) and the two specimens from Costa Rica, listed at the end of the description of P. elcapitanensis by Meek & Hildebrand (in Meek, 1914) (FMNH 7901 and FMNH 7916) have been sometimes erroneously considered as paratypes (e.g. Grey, 1947; Ibarra & Stewart, 1987: 80). The two specimens from Costa Rica were identified by Dawson (1982) as Pseudophallus starksii , an identification confirmed in the present study.

Type material: lectotype. USNM 81735 , 1 , El Capitain, Río Mamoni , Panamá, S. E. Meek & S. F. Hildebrand 20 Mar 1912 (not examined) . Paralectotypes. FMNH 8291/8292, 2, 108.5– 130.8 mm SL, same data as lectotype; USNM 79690 , 1 , same data as lectotype (not examined); USNM 108645 , 1 , same data as lectotype (not examined) .

Non-types. Panama: Darien: AMNH 11376, 1, 125.0 mm SL, Río Chico , Marsh Expedition, 24 Feb 1924 ; FMNH 8290, 1, 122.9 mm SL, Río Tuyra, Boca de Cupe , S. E. Meek & S. F. Hildebrand 28 Feb 1912 ; FMNH 8294, 1, 73.9 mm SL, Río Calobra , S. E. Meek & S. F. Hildebrand, 18 Mar 1911; MCZ 88623 , 1, 110.6 mm SL, Río Pucuro just above confluence with Río Tuira , 08º0’N 77º32’W, K. E. Hartel, 16 Feb 1985 ; Panama: CAS 106919 , 2 , 86.6–119.3 mm SL, Río Grande, Miraflores, C. H. Hilbert et al., 1900; FMNH 8293, 1, 85.3 mm SL, Río Mamoni , Chepo , S. E. Meek & S. F. Hildebrand 21 Mar 1911 ; Chiriqui: ANSP 99933, 1, 115.9 mm SL, creek 5 mi W. of Boca del Monte on old road to David, H. G. Loftin & E. L. Tyson, 18 Nov 1961 ; ANSP 104124, 4, 64.4–130.0 mm SL, Creek at bridge 5 mi. W. of San Felix bridge on Interamerican Highway new section, H. G. Loftin & E. L. Tyson, 18 Nov 1961 ; ANSP 104212, 2, 105.2– 106.3 mm SL, Río Tinta, creek at bridge 3 mi. W. of Río Tabasara on Sona-Remedios road, H. G. Loftin, E. L. Tyson & M. Canipe, 11 Nov 1961 ; ANSP 104342, 1, 80.6 mm SL, Creek Chacarrera on Sona-Remedios Rd. 14 mi. E. of Remedios, H. G. Loftin, E. L. Tyson & M. Canipe, 12 Nov 1961 ; USNM 208369 , 8 , 72.3–100.1 mm SL, Creek 14 m. W. of San Juan on I. Am. Highway, H. G. Loftin & E. L. Tyson, 18 Nov 1961 ; USNM 208371 , 12 , 87.6–135.2 mm SL, Río Estero Salado near David, H. G. Loftin, 1 Dec 1961 .

Costa Rica: FMNH 7916, 1, 104.8 mm SL, Río Turrubales , Orotina, S. E. Meek, 21 Apr 1912 ; TNHC 14736, 2, 129.7– 148.2 mm SL, Río Pavo , P. N. Corcovado 08º51’N 83º58’W, K. Winemiller, Aug 1986 .

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 5. Distribution map of species of Pseudophallus occurring in drainages along the Pacific coast based on specimens examined in this study.

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).