Taxonomic review of the pipefish genus Pseudophallus Herald, with the description of a new species (Syngnathiformes: Syngnathidae)
Author
Dallevo-Gomes, Caio I. A.
0000-0001-7788-5861
Departamento de Zoologia, Instituto de Biociências da Universidade de São Paulo, Rua do Matão, travessa 14, no. 101, 05508 - 090, São Paulo, SP Brazil. & caioisola @ yahoo. com. br; https: // orcid. org / 0000 - 0001 - 7788 - 5861
caioisola@yahoo.com.br
Author
Mattox, George M. T.
Departamento de Biologia, Universidade Federal de São Carlos, Campus Sorocaba, São Paulo, Brazil.
Author
Toledo-Piza, Mônica
0000-0001-7596-1410
Departamento de Zoologia, Instituto de Biociências da Universidade de São Paulo, Rua do Matão, travessa 14, no. 101, 05508 - 090, São Paulo, SP Brazil. & mtpiza @ usp. br; https: // orcid. org / 0000 - 0001 - 7596 - 1410
mtpiza@usp.br
text
Zootaxa
2020
2020-10-05
4859
1
81
112
journal article
8217
10.11646/zootaxa.4859.1.3
ff9ddcdc-1340-4533-ae1e-da83b895158c
1175-5326
4412809
C213B95B-6707-4F80-A1EA-58245141F24B
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
).
FIGURE 5.
Distribution map of species of
Pseudophallus
occurring in drainages along the Pacific coast based on specimens examined in this study.
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
.