Pseudophallus brasiliensis Dawson, 1974

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 : 86-90

Pseudophallus brasiliensis Dawson, 1974

( Figs. 2–4 , 6–7 ; Tables 1 , 3–6 )

Pseudophallus brasiliensis Dawson, 1974: 405 , figs. 1–3 (original description, type locality: “ Brazil, Pará, Rio Tocantins, Igarapé Inó, Faro (sic) de Panaquera; 01 o 52’ S, 49 o 10’ W ” (=Furo de Panaquera); holotype: MZUSP 10278); Dawson, 1982: 50–52 (synonym of P. mindii ); Ferraris Jr., 2003: 591 (as junior synonym of P. mindii , listed); Menezes et al., 2003: 71 (listed as valid; Brazil); Kuiter, 2009: 242 (distribution, as a freshwater species, listed as valid, figures from Dawson, 1982), Hamilton et al., 2017: 393–395, figs. 1–3 (phylogeny, molecular data).

Pseudophallus aff. brasiliensis: Mol et al., 2012: 278 (checklist, Corantijn and Suriname River, Suriname).

Pseudophallus cf. brasiliensis: Mol, 2012: 45–46 , 48–52, figs. 3, 4 (Corantijn River, Suriname: distribution, conservation, ecology).

Pseudophallus mindii (non Meek & Hildebrand, 1923): Dawson, 1982: 23, 45, 47–53 (in part; as a junior synonym of P. mindii , description, distribution, comparisons with congeners, identification key); Dawson, 1985: 203 (in part; as a junior synonym of P. mindii , distribution, biogeography); Menezes et al., 2003: 71, 153 (in part, listed as valid; Brazil); Carvalho-Filho, 1999: 90 (in part, listed, distribution; Brazil); Szpilman, 2000:83 (in part, checklist, Brazil); Fritzsche & Vincent, 2003: 1222 (in part; as a junior synonym of P. mindii , listed, distribution); Sazima & Carvalho-Filho, 2003: 177 (recorded at same habitat as Lupinoblennius paivai ); Lasso et al., 2004: 144 (listed, Orinoco Delta, Venezuela); Corrêa & Uieda, 2007: 489 (diet, listed, Parque Estadual da Serra do Mar, São Paulo State, Brazil); Bonecker et al., 2009: 243–244 (listed, Mucuri Estuary, Espírito Santo State, Brazil); Kuiter, 2009: 241 (in part; as a junior synonym of P. mindii , distribution, freshwater; figures from Dawson, 1982); Mattox & Iglesias, 2010: 110 (listed, Rio Jurubatuba, Santos, São Paulo State, Brazil); Sarmento-Soares et al., 2012: 8, 16, 21, 23 (listed, Piúma, Espírito Santo State, Brazil); Di Dario et al., 2013: 261, 263, 270, 278 (listed, Restinga de Jurubatiba National Park, Rio de Janeiro State, Brazil, identification key); Vieira et al., 2014: 19, 85, 96 (listed, Rio Benevente, Espírito Santo State, Brazil, distribution, regional conservation status and ecological aspects, photograph, identification key); Lima et al., 2015: 217 (larvae; listed, Goiana Estuary, Pernambuco State, Brazil); Araújo et al., 2016: 112 (Sepetiba Bay, Rio de Janeiro State, Brazil, ecology).

Pseudophallus mindi (sic) (non Meek & Hildebrand, 1923): Figueiredo & Menezes, 1980: 11, fig. 19 (Southeastern Brazil;, characters, distribution, identification key); Godefroid et al., 2003:50,53 (listed, Paranaguá bay, Paraná State, Brazil); Neves et al., 2011:121 (listed, rio Mambucaba, Rio de Janeiro State, Brazil).

Pseudophallus starksi mindii: Lüling, 1974: 196 , fig. 7 (new combination based on identification by E. S. Herald; listed for Rio Magé-Roncador, Rio de Janeiro, Brazil); Dawson, 1982: 50 (synonymy list of P. mindii ).

Diagnosis. Pseudophallus brasiliensis differs from species of Pseudophallus from the Pacific coastal drainages, P. starksii and P. elcapitanensis in having 90.5% of specimens with 13 trunk rings (vs. 14 trunk rings).

Pseudophallus brasiliensis differs from P. mindii in the following combination of characters: 12–15 pectoralfin rays (82.4% of specimens with 13), 28–41 dorsal-fin rays (98.5% of specimens with 28–37) and 13–17 broodpouch rings in adult males (87.7% of the adult males with 14–16) vs. 13–16 pectoral-fin rays (88.2% of specimens with 14–15), 32–42 dorsal-fin rays (76.7% of specimens with 35–38) and 15–23 brood-pouch rings in adult males (93.8% of adult males specimens with 15–21) in P. mindii .

Pseudophallus brasiliensis differs from P. galadrielae in having 28–41 dorsal-fin rays (98.5% of specimens with 28–37), 6.5–10.5 subdorsal rings (86.5% of specimens with 7.0–9.5), 12–15 pectoral-fin rays (82.4% of specimens with 13) and 40–50 total body rings (95.3% of specimens with 43–49) vs. 27–34 dorsal-fin rays (91.6% of specimens with 27–31), 6.0–8.25 subdorsal rings (92.9% with 6.0–7.25), 12–15 pectoral-fin rays (60.6% of specimens with 14–15) 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 2A and 3A. Body without scales, covered with bony plates forming rings along its axis, bony plates horizontally oval, with elevation in midline forming bony ridges along body. Trunk rings 12–14 (90.5% of specimens with 13). Tail rings 27–37 (83.9% of specimens with 31–35). Body rings 40–50 (95.3% of specimens with 43–49). Subdorsal rings 6.5–10.5 (86.5% of specimens with 7.0–9.5). Subdorsal trunk rings 0–0.5. Subdorsal tail rings 6.5–11.0 (72.3% of specimens with 8.0–9.75). Dorsal-fin origin at vertical between posterior half 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 body depth of females and juveniles at middle of trunk; in males, at anterior region of brood pouch. Body depth decreasing posteriorly along superior trunk ridge to minimum depth at caudal peduncle. Dorsal profile of head straight or slightly inclined at snout region, more inclined at orbital region, 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 28–41 (98.5% of specimens with 28–37), 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, with distal margin rarely extending beyond third trunk ring. Pectoral-fin rays 12–15 (82.4% of specimens with 13). 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 13–17 rings (86.5% of specimens with 14–16 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 life. Coloration based on six specimens captured in Rio Jurubatuba, Santos, Brazil (ZUEC 17250). Coloration varies according to surroundings. Specimens near leaves on the substrate with ground color dark brown with darker brown spots on some rings along the body; spots of variable size, but never larger than width of one bony ring. Specimens on sandy substrate or plants light beige with small brown spots at every two to four rings; spots usually smaller than width of one bony ring. Live image also available in Mol (2012).

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 strip. Dorsal-fin rays with scattered brown chromatophores. Pectoral fin hyaline. Caudal fin with same general coloration of body, sometimes in slightly different tone ( Figs. 2A and 3A ).

Distribution. Coastal freshwater drainages from Santa Catarina State, Brazil, to the Río Orinoco Delta, Delta Amacuro, Venezuela ( Fig. 4 ).

Remarks. The southern limit of distribution of P. brasiliensis based on specimens examined in the present study is Rio Ribeira de Iguape (24º70´S 47º56´W), Registro, São Paulo State (USNM 144050). Godefroid et al. (2003) recorded the presence of Pseudophallus south from that region, at Baía de Paranaguá (25º32’57”S 48º22’15”W), however, they did not mention voucher specimens. One specimen of P. brasiliensis from Rio Guaraqueçaba, state of Paraná (MZUEL 8928) was recently discovered but it was not included in the statistical analysis. A second specimen (MZUSP 860) from Rio Itajaí, Blumenau, state of Santa Catarina is damaged, rendering a detailed analysis of its features impossible. Both specimens further extend the southern limit of the geographic distribution of P. brasiliensis .

Type material: holotype. MZUSP 10278 : 1, 84.7 mm SL, male, Brazil, Pará, Rio Tocantins, Igarapé Inó , Furo do Panaquera , 01º52’S, 49º10’W, Expedição Permanente da Amazônia , 1 Sept 1970 . Paratypes. All from Brazil, Pará: GCRL 12755, 2, 79.1–112.2 mm SL, MZUSP 10279 , 2 , 56.8–62.2 mm SL, Igarapé Mapará , Samaúma , Rio Tocantins, 02º05’S, 49º20’W, Expedição Permanente da Amazônia , 5 Sept 1970; MZUSP 10280 : 1, 48.7 mm SL, Igarapé Acicurá , Rio Tocantins, Cametá , 02º15’S, 49º30’W, Expedição Permanente da Amazônia , 7 Sept 1970; MZUSP 10627 , 1 , 41.5 mm SL, USNM 212058 , 1 , 69.1 mm SL, taken with holotype.

Non - types. Brazil: São Paulo: MZUSP 58747 , 1, 109.3 mm SL, Rio Tetequeira, Peruíbe , C.P. Silva & N. Silva, Mar 1997 ; MZUSP 3214 , 2 , 94.4–100.8 mm SL, Santos, A. Marques, 27 Feb 1941 ; MZUSP 37374 , 2 , 69.8–82.4 mm SL, Rio Cabuçu, Santos , 23º54’36’’S, 46º18’36’’W, H. A. Britski, S. Kullander, H. Ortega & M. Villacorta, 21 Aug 1980 ; MZUSP 93649 , 4 , 43.7–73.1 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51” W, G. M. T. Mattox & J. M. P. Iglesias, 21 Mar 2007 ; MZUSP 93655 , 10 , 39.65 –86.0 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 22 Mar 2007 ; MZUSP 103632 , 1 , 83.5 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 02 Oct 2007 ; MZUSP 103649 , 2 , 57.0– 102.1 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 04 Oct 2007 ; ZUEC 17247 , 12 , 58.5 –104.0 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 03 Oct 2007 ; ZUEC 17248 , 5 , 52.2–70.4 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 04 Dec 2007 ; ZUEC 17249 , 5 , 49.2–110.5 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 06 Dec 2007 ; ZUEC 17250 , 6 , 74.0–83.0 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, C. I. A. Dallevo-Gomes & G. M. T. Mattox, 24 Aug 2016 ; ZUEC 17251 , 6 , 42.5–83.5 mm SL, Rio Jurubatuba, Santos , 23°51’50”S, 46°17’51”W, G. M. T. Mattox & J. M. P. Iglesias, 20 Mar 2007 ; MZUSP 9992 , 1, 106.7 mm SL, Mercado de São Paulo, R. P. Lambalot, Dec 1970 ; MZUSP 77995 , 3 , 97.6–108.4 mm SL, Rio Ubatubamirim on Rio-Santos Hwy, Ubatuba , 23º43’S, 45º07’W, Excursão MZUSP, 17 Jan 2000 ; MZUSP 93930 , 1 , 50.5 mm SL, Rio da Fazenda , Ubatuba, V. Uieda & Corrêa, 2004 ; USNM 144050 , 1, 129.9 mm SL, Rio Ribeira de Iguape , Registro, J.P. Carvalho, 23 Jun 1941 ; ZUEC 4963 , 3 , 33.8–55.6 mm SL, Rio da Fazenda, Ubatuba , 23º21’59’’S, 45º50’15’’W I. Sazima, 15 Jun 2000 ; ZUEC 6111 , 1 , 76.7 mm SL, Rio Escuro, Ubatuba , 23º28’59’’S, 44º09’00’’W, I. Sazima, 30 Nov 1980 ; ZUEC 8640 , 1 , 87.3 mm SL, Rio Escuro (200m from beach), Ubatuba , 23º29’27’’S, 44º09’00’’W A. Carvalho-Filho, 22 Apr 2000 ; Rio de Janeiro: CAS 14378, 1, 101.9 mm SL, Rio Roncador , Rio de Janeiro, K. H. Lüling, 14 Jan 1972 ; CAS 136439 , 1, 107.6 mm SL, Rio Grande, Arroio Fundo, Santa Cruz , Rio de Janeiro, 22º56’S, 43º12’W, Thayer Expedition, Apr 1865 ; CAS 162398 , 1 , 64.2 mm SL, Canal, Guandu, above Itaguaí , Baixada Fluminense, Rio de Janeiro, W. C. Freihofer, Sep 1962 ; MCZ 11732, 3 , 100.3 – 120.2 mm SL, Rio Grande, Arroio Fundo, Santa Cruz , Rio de Janeiro, 22º56’S, 43º12’W, Thayer Expedition, Apr 1865 ; MNRJ 19178 , 1 , 86.2 mm SL, Rio Saco, Angra dos Reis, W. D. Bandeira & D. F. Moraes, 16 Jul 1987 ; Espírito Santo: CIUFES 131446, 3, 84.2–86.1 mm SL, Estuary of Rio Santa Maria , Vitória, J. L. Helmer, 04 Oct 1989 ; CIUFES 131055 , 1 , 83.2 mm SL, 500 from Monte Agha, Piúma, J. L. Helmer, 30 Mar 1981 ; MNRJ 27160 , 14 , 48.4 –149.0 mm SL, Rio Piraquê Açu, ca. 1 km downstream from where crossed by road ES–124, Aracruz , 17º22’12’’S, 47º31’12’’W, M. Britto & R. Campos-da-Paz, 27 Jun 2004 ; MNRJ 27135 , 1 , 82.3 mm SL, Rio Piraquê Açu near concrete bridge on road ES–124, Aracruz , 19º51’00’’S, 40º16’48’’W, M. Britto & R. Camposda-Paz, 24 Jun 2004 ; MNRJ 27198 , 1 , 80.3 mm SL, Ribeirão Sahuanhá or Timbuí near bridge at road ES–351, 20º03’S, 40º34’W, M. Britto & R. Campos-da-Paz, 26 Jun 2004 ; MZUSP 865 , 1 , 67.1 mm SL, Rio Doce , Linhares, E. Garbe, 1906 ; Bahia: FMNH 70588, 2, 63.9–73.6 mm SL, Rio Paraguassu , Cachoeira, J. D. Haseman, 14 Apr 1908 ; FMNH 88221 , 1 , 63.1 mm SL, Rio Paraguassu , J. D. Haseman, 14 Apr 1908 ; MBML 1308 , 7 , 56.2–107.2 mm SL; Rio Mucuri, Mucuri , 18º08’S, 39º55’W, J. L. Gasparini & C. Zamprogno, 01 Jan 1991 ; MZUSP 64514 , 1 , 61.6 mm SL, creek near Cachoeira, Rio Paraguaçu , N. A. Menezes & P. Whitehead 11 Dec 1974 ; Piauí: MCZ 46862, 1, 36.7 mm SL, mangrove near Parnaiba , 02º52’12’’S, 41º42’36’’W, T. R. Roberts, 01 Aug 1968 ; Pará: MZUSP 64503 , 7 , 40.2–83.8 mm SL, Igarapé Inó, Furo de Panaquera , Expedição Permanente à Amazônia, 01 Sep 1970 ; MZUSP 64504 , 1 , 40.2 mm SL, Igarapé Bahia near vila Maiauatá , 01º51’0’’S, 49º00’36’’W, Expedição Permanente à Amazônia, 28 Aug 1970 ; MZUSP 64505 , 4 , 60.7–76.9 mm SL, Igarapé Apicurá, Cametá, Expedição Permanente à Amazônia, 07 Sep 1970 .

Suriname: Sipaliwini: CAS 230532 , 1 , not measured, damaged specimen, Potoka Bank, Corantijn River , 5º3’21’’N, 57º18’57’’W, J. H. Mol, 11 Sep 2007 ; CAS 230533 , 1 , not measured, damaged, Pidgeon Island, Corantijn River , 04º59’17’’S, 57º 25’26’’W, J. H. Mol, 14 Sep 2007 ; CAS 230535 , 1 , not measured, damaged specimen, Suriname River near Carolina Bridge Km 118, K. Y. W. Tong & A. de Kom, 11 Jun 2011 ; FMNH 119605, 2, 36.6– 51.2 mm SL, Corantjin River at sand landing in Apoera, J. H. Mol, 22 Mar 2007 ; ROM 100796, 3, 51.5–57.9 mm SL, Kabalebo River , 05º03’24’’S, 57º18’58’’W, K. Gajapersad, M. Kolmann, S. Steele & K. Wan, 11 Mar 2015 .

Venezuela: Delta Amacuro: GCRL 18831, 2, 78.3–82.5 mm SL, Río Orinoco near Curiapo , 08º36’24’’S, 61º00’00’’W, J. Baskin, 19 Nov 1979 ; LACM 43348–9, 5, 68.8–86.4 mm SL, Río Orinoco near Barrancas , 08º43’S, 62º18’W, J. G. Lundberg, 12 Nov 1979 ; USNM 233547 , 3 , 58.9 –99.0 mm SL, Small caño on south shore of isla Guacujana , 50–51 NM from sea buoy, 08º36’51’’N, 60º51’34’’W, J. Baskin, 22 Jun 1982 . Monaguas: USNM 219325 , 1 , 58.8 mm SL, no information (same collector and date as USNM 219326 ) ; USNM 219326 , 1 , 83.3 mm SL, Río Orinoco , small cano 3/ 4 mile upstream from mouth of Caño Guarguapo near Barrancas, J. N. Baskin & J. G. Lundberg, 18 Feb 1978 .

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).