Redescription and diagnoses of the genera Profundulus and Tlaloc (Cyprinodontiformes: Profundulidae), Mesoamerican endemic fishes
Author
Domínguez-Cisneros, Sara E.
Doctorado en Biodiversidad y Conservación de Ecosistemas Tropicales, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Libramiento Norte, 1150, Col. Lajas Maciel, 29029 Tuxtla Gutiérrez, CHIS, México. (SEDC) sara. dominguez @ unicach. mx. & Colección de Peces, Museo de ZoologÍa, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Libramiento Norte, 1150, Col. Lajas Maciel, 29029 Tuxtla Gutiérrez, CHIS, México. (EVV) ernesto. velazquez @ unicach. mx (corresponding author).
ernesto.velazquez@unicach.mx
Author
Domínguez-Domínguez, Omar
Laboratorio de BiologÍa Acuática, Facultad de BiologÍa, Universidad Michoacana de San Nicolás de Hidalgo, Edificio “ R ” planta baja, Ciudad Universitaria, Francisco J. Mujica S / N, 58030 Morelia, MEX, México. (ODD) goodeido @ yahoo. com. mx, (RPR) rperezr 38 @ gmail. com.
goodeido@yahoo.com.mx
Author
Velázquez-Velázquez, Ernesto
Colección de Peces, Museo de ZoologÍa, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Libramiento Norte, 1150, Col. Lajas Maciel, 29029 Tuxtla Gutiérrez, CHIS, México. (EVV) ernesto. velazquez @ unicach. mx (corresponding author).
ernesto.velazquez@unicach.mx
Author
Pérez-Rodríguez, Rodolfo
Laboratorio de BiologÍa Acuática, Facultad de BiologÍa, Universidad Michoacana de San Nicolás de Hidalgo, Edificio “ R ” planta baja, Ciudad Universitaria, Francisco J. Mujica S / N, 58030 Morelia, MEX, México. (ODD) goodeido @ yahoo. com. mx, (RPR) rperezr 38 @ gmail. com.
goodeido@yahoo.com.mx
text
Neotropical Ichthyology
2023
e 220089
2023-02-20
21
1
1
22
http://dx.doi.org/10.1590/1982-0224-2022-0089
journal article
295747
10.1590/1982-0224-2022-0089
9ec7a6a4-dd99-4d15-a180-0e3dbed303da
1982-0224
11125059
Tlaloc
Álvarez & Carranza, 1951
Tlaloc
Álvarez, Carranza, 1951:40
(
type
species
Fundulus labialis
Günther, 1866
, by monotype).
Diagnosis.
Tlaloc
, one of the two genera of the family
Profundulidae
, is diagnosed here by the following combination of characters: The mesethmoid is prominent and oval in shape, extending beyond the posterior margins of the vomer, encompassing the posterior medial extension and touching the lateral ethmoids (
Fig. 2B
). The anterior portion of the parasphenoid making contact with the mesethmoid and extend beyond the center of the mesethmoid. The autopterotic fossa is reduced (
Fig. 2A
). The dorsal margin of the interoperculum, with a long extension, is exceeding the edge of the bone (
Fig. 3A
). The ventral margin of the lacrimal is straight (
Figs. 4A–B
).
Tlaloc
is further distinguished from
Profundulus
by having less than the basal half of the caudal fin densely scaled (except in
T. portillorum
) (
vs
. more than the basal half or more densely scaled) (
Fig. 5A
); by the absence of a humeral spot (
vs
. humeral spot present); by the origin of the dorsal fin positioned at a vertical line posterior to the origin of the anal fin (
vs
. origin of the dorsal fin positioned at a vertical line slightly anterior to the origin of the anal fin); by long epiotic processes, extending beyond the second vertebra (
vs
. short epiotic processes, not extending beyond the first vertebra).
FIGURE 2 |
Ventral view of the neurocranium, in specimens cleared and stained.
A.
Tlaloc portillorum
(MZ-UNICACH 7222, 70.5 mm SL);
B.
Vomer region of
T. candalarius
(MZ-UNICACH 3899, 61.7 mm SL);
C.
Vomer region of
Profundulus kreiseri
(MZ-UNICACH 7214, 59.6 mm SL); and
D.
Neurocranium of
P. mixtlanensis
(MZ-UNICACH 6716, 55.6 mm SL). Abbreviations: V, vomer (Y-shaped); E, lateral ethmoid; M, mesethmoid; P, parasphenoid; AF, autopterotic fossa.
FIGURE 3 |
Lateral view of left jaws, suspensorium and opercular series in specimens cleared and stained.
A.
Tlaloc portillorum
(MZ-UNICACH 7222, 70.5 mm SL);
B.
Profundulus mixtlanensis
(MZ-UNICACH 6716, 55.6 mm SL). Arrow points to dorsal margin of the interoperculum, with a long or short extension. Abbreviations: de, dentary; pm, premaxilla; ra, retroarticular; mx, maxilla; pl, palatine; qu, quadrate; ar, articular; ms, mesopterygoid; io, interopercle; sy, sympletic; hy, hyomandibula; po, preopercle; op, opercle; so, subopercle.
Description.
Morphometric data appear in
Tab. 1
. Body uniformly slender; head compressed, moderately large (22.4–36.7%). Mouth subterminal to terminal. In
T. hildebrandi
and
T. portillorum
, the lower jaw broad, heavy, and protruding so that the upper jaw is included, whereas in
T. labialis
and
T. candalarius
both equal in forward projection, or the lower jaw is included in the upper jaw. Greatest body depth in the vertical just posterior to pectoral fin (20.0–32.6%). The females are elongated, and the males are more robust than the females. Dorsal and anal fins located posterior to the half of body length. Caudal peduncle relatively large (16.2–26.4%). Median hypural plate divided into subequal parts by an open groove.
TABLE 1 |
Morphometric data of
Tlaloc
and
Profundulus
species. Asterisk mark the number of specimens analyzed, in parenthesis the average values.
|
Tlaloc portillorum
58*
|
T. hildebrandi
63*
|
T. labialis
58*
|
T. candalarius
100*
|
Profundulus punctatus
196*
|
P. guatemalensis
26*
|
| Standard length (mm) |
37.6–71.4 (50.4) |
37.9–111.5 (58.2) |
33.4–101.9 (53.1) |
38.8–99.2 (51.6) |
36.2–82.2 (50.3) |
38.4–83.9 (51.8) |
|
Percents of standard length
|
| Head length |
25.7–32.0 (28.9) |
26.8–32.2 (28.3) |
22.4–31.6 (26.9) |
26.3–36.7 (29.4) |
26.1–32.6 (29.1) |
27.0–31.1 (28.9) |
| Predorsal length |
66.1–73.6 (69.6) |
64.2–68.5 (66.3) |
65.0–70.4 (67.4) |
63.6–73.8 (69.7) |
62.2–74.7(68.0) |
65.5–71.5 (68.4) |
| Prepelvic length |
47.7–55.2 (51.1) |
45.8–51.8 (48.9) |
46.8–54.7 (50.6) |
48.9–61.1 (53.7) |
47.0–57.6 (52.2) |
49.8–53.6 (51.5) |
| Anal origin to caudal base |
29.0–36.7 (33.4) |
36.3–41.9 (39.1) |
32.3–41.8 (36.1) |
27.3–39.7 (33.3) |
19.3–37.4 (33.2) |
31.9–36.7 (34.5) |
| Body, greatest depth |
20.0–27.8 (25.2) |
22.1–29.7 (24.6) |
20.2–29.9 (24.4) |
20.7–32.6 (27.7) |
21.0–33.6 (28.5) |
25.7–31.5 (29.0) |
| Body, greatest width |
11.8–17.0 (14.5) |
12.5–17.6 (14.8) |
10.6–18.1 (14.3) |
11.5–19.5 (15.6) |
14.3–22.8 (18.3) |
10.8–18.2 (14.7) |
| Caudal peduncle, length |
16.3–23.5 (19.8) |
20.3–26.4 (23.3) |
17.8–23.6 (21.5) |
16.2–24.0 (19.5) |
13.7–21.8 (18.3) |
15.5–22.1 (18.8) |
| Caudal peduncle, Least depth |
12.6–15.9 (14.3) |
12.0–15.1 (13.4) |
12.6–15.1 (13.5) |
11.4–16.6 (13.5) |
13.9–18.3 (15.7) |
14.2–16.1 (15.2) |
| Dorsal fin, basal length |
11.1–16.5 (14.1) |
11.7–17.6 (14.1) |
11.6–16.4 (14.0) |
11.1–17.7 (13.9) |
11.1–18.3 (15.0) |
12.5–16.6 (14.7) |
| Anal fin, basal length |
12.1–17.4 (14.3) |
14.0–19.1 (16.0) |
12.7–19.5 (15.5) |
12.3–17.5 (14.5) |
11.6–20.1 (15.7) |
14.3–18.6 (16.0) |
|
Percents of head length
|
| Head depth |
55.6–98.0 (67.3) |
53.3–85.4 (62.5) |
57.5–76.2 (67.0) |
56.6–100.1 (68.5) |
56.5–94.9 (72.7) |
64.8–101.7 (86.0) |
| Head width |
49.3–72.9 (59.6) |
49.2–71.0 (64.6) |
54.8–70.6 (62.1) |
49.2–70.0 (60.7) |
58.8–80.6 (68.7) |
55.8–75.8 (64.3) |
| Interorbital, least bony width |
37.3–48.7 (42.1) |
40.0–48.7 (45.2) |
41.1–51.7 (46.2) |
34.9–48.3 (42.9) |
40.0–53.2 (47.2) |
43.3–54.1 (47.3) |
| Orbit length |
21.1–28.6 (24.5) |
18.0–28.9 (24.8) |
24.9–32.7 (28.6) |
19.5–30.6 (26.5) |
19.0–29.9 (25.4) |
21.6–29.2 (24.0) |
| Snout length |
28.8–35.6 (32.7) |
29.8–39.7 (34.9) |
29.3–39.7 (33.7) |
27.5–38.3 (32.8) |
25.6–37.4 (32.3) |
28.6–36.9 (33.3) |
| Upper jaw length |
23.9–32.9 (28.3) |
26.3–35.3 (31.0) |
22.1–31.4 (27.6) |
21.5–32.3 (27.7) |
16.9–31.7 (25.3) |
20.7–33.2(29.6.3) |
TABLE 1 |
(Continued)
|
P. oaxacae
28*
|
P. balsanus
22*
|
P. kreiseri
52*
|
P. parentiae
40*
|
P. mixtlanensis
59*
|
P. adani
73*
|
p. chimalapensis
38*
|
| Standard length (mm) |
35.0–63.2 (45.5) |
34.2–64.7 (49.7) |
37.2–72.7 (49.4) |
36.8–73.9 (52.7) |
39.0–70.6 (49.7) |
36.7–77.9 (53.7) |
35.9–80.3 (45.6) |
|
Percents of standard length
|
| Head length |
26.3–30.5 (28.7) |
24.1–30.8 (26.8) |
26.2–31.6 (28.8) |
26.9–35.3 (29.4) |
26.8–35.3 (29.1) |
26.1–33.9 (28.6) |
27.4–33.4 (30.8) |
| Predorsal length |
65.5–72.2 (69.0) |
61.9–71.7 (65.5) |
64.1–69.8 (67.0) |
63.7–69.3 (66.9) |
63.9–70.5 (67.1) |
64.7–73.5 (68.2) |
66.3–77.4 (69.4) |
| Prepelvic length |
47.9–57.3 (52.2) |
48.1–55.5 (51.4) |
48.2–54.0 (51.7) |
49.2–54.4 (51.5) |
51.0–57.2 (53.8) |
49.0–57.6 (53.3) |
50.2–56.4 (53.9) |
| Anal origin to caudal base |
25.2–32.6 (29.1) |
29.2–37.1 (33.4) |
29.7–36.8 (34.0) |
29.0–37.4 (33.3) |
30.8–37.8 (34.0) |
29.4–37.9 (33.3) |
27.1–35.6 (31.2) |
| Body, greatest depth |
23.5–30.8 (26.6) |
25.2–31.0 (27.8) |
21.2–30.0 (25.7) |
26.3–33.8 (30.2) |
23.7–31.2 (26.9) |
20.4–31.0 (26.1) |
24.3–30.8 (27.5) |
| Body, greatest width |
14.4–18.6 (16.5) |
11.9–22.2 (17.4) |
10.8–18.7 (14.2) |
13.2–21.4 (17.8) |
10.6–18.6 (15.0) |
11.3–20.8 (15.1) |
15.4–19.4 (17.5) |
| Caudal peduncle, length |
11.1–19.6 (14.8) |
15.5–20.8 (17.2) |
15.5–21.7 (18.5) |
14.3–20.0 (17.9) |
17.3–23.2 (19.7) |
15.5–21.9 (18.0) |
14.2–19.9 (17.0) |
| Caudal peduncle, Least depth |
11.9–16.6 (14.4) |
11.8–15.9 (13.8) |
12.4–16.2 (14.4) |
14.1–16.9 (15.3) |
11.9–15.9 (13.7) |
11.7–15.4 (13.6) |
13.9–17.4 (15.3) |
| Dorsal fin, basal length |
12.8–17.7 (15.2) |
12.5–19.2 (15.2) |
13.3–18.1 (15.5) |
11.6–18.6 (15.0) |
11.9–17.2 (14.2) |
11.7–18.3 (14.7) |
12.5–16.5 (14.7) |
| Anal fin, basal length |
11.7–18.5 (14.7) |
13.2–21.0 (16.1) |
13.7–18.3 (15.8) |
13.2–19.0 (16.1) |
12.7–18.9 (15.3) |
13.5–19.4 (16.2) |
12.6–18.9 (15.2) |
|
Percents of head length
|
| Head depth |
66.4–101.5 (81.0) |
56.1–80.6 (69.3) |
60.1–90.7 (75.0) |
58.3–83.3 (70.5) |
59.7–82.9 (73.2) |
59.0–80.2 (71.9) |
59.9–70.5 (65.8) |
| Head width |
60.1–72.3 (65.0) |
57.9–75.5 (68.1) |
44.5–70.8 (60.9) |
53.8–77.6 (68.5) |
52.9–72.2 (65.7) |
54.8–97.5 (67.6) |
61.0–71.7 (65.6) |
| Interorbital, least bony width |
41.5–48.3 (44.3) |
38.7–55.9 (50.3) |
38.1–48.5 (44.1) |
38.9–58.7 (50.2) |
38.2–52.7 (45.1) |
38.9–50.2 (46.0) |
45.9–52.7 (48.7) |
| Orbit length |
21.2–26.3 (23.8) |
21.4–28.9 (25.0) |
20.4–31.3 (25.2) |
21.5–28.9 (25.4) |
23.1–30.0 (26.3) |
20.5–28.5 (23.6) |
22.0–32.6 (28.0) |
| Snout length |
24.0–32.2 (28.8) |
15.4–34.9 (25.3) |
27.2–38.5 (32.5) |
22.2–34.2 (29.2) |
27.3–37.8 (32.6) |
26.6–38.2 (33.5) |
28.7–37.9 (33.1) |
| Upper jaw length |
21.8–36.1 (26.9) |
11.1–23.9 (16.5) |
24.3–34.4 (28.7) |
11.2–30.0 (18.4) |
24.3–35.9 (29.7) |
21.1–36.6 (33.5) |
23.5–32.0 (26.4) |
FIGURE 4 |
Ventral view of left lacrimal.
A.
Tlaloc portillorum
(MZ-UNICACH 7222, 70.5 mm SL);
B.
T. hildebrandi
(MZ-UNICACH 2266, 75.1 mm SL);
C.
Profundulus mixtlanensis
(MZ-UNICACH 6716, 55.6 mm SL); and
D.
P. punctatus
(MZ-UNICACH 6632, 67.3 mm SL). Arrow point to ventral margin of the lacrimal.
The branching pattern of the cephalic latero-sensory canals in
Tlaloc
is similar to the general pattern of the
Cyprinodontiformes
. The supraorbital series follows the
Type
II designation of Gosline (1949), with canals between pores 1–2a, 2b–4a, 4b–7; preopercular pores 6–7 (mode = 7); preorbital pores 3–5 (mode = 4); mandibular pores 4–5 (mode = 5); in addition, there are 2 to 4 rostral pores usually developed.
Meristic characters are as follows (
Tab. 2
): anal: 9–18; dorsal: 7–14; pelvic: 5–7; pectoral: 13–20; caudal: 16–23; vertebrae totals: 33–38; gill rakers on anterior arm of the first arch: 13–19; branchiostegal rays: 6; scales lateral series: 29–39; scales predorsal: 19–26; scales around body: 25–36; scales around peduncle: 16–28.
Coloration.
The coloration varies strongly among individuals and ontogenetically. The skin of the head and body may show golden reflections, especially in the opercular region and mid flank. The form and coloration of the nuptial adult male and female are shown in
Fig. 6
. Unpaired fins (dorsal and anal) are orange in males, with irregular black markings on the dorsal fin, evident in
Tlaloc candalarius
.
TABLE 2 |
Meristic data of
Tlaloc
and
Profundulus
species.
Asterisk mark the number of specimens analyzed, in parenthesis the modal values.
| Character |
T. labialis
58*
|
T. candalarius
102*
|
T. hildebrandi
63*
|
T. portillorum
58*
|
P. punctatus
202*
|
P. guatemalensis
26*
|
| Dorsal rays |
10–14 |
9–14 (11) |
10–14 (11) |
7–14 (11) |
10–14 (12) |
11–13 (12) |
| Anal rays |
13–18 (16) |
11–16 (14) |
11–16 (15) |
9–14 (13) |
11–16 (14) |
12–15 (14) |
| Pectoral rays |
16–20 (18) |
14–19 (15) |
15–18 (16) |
13–17 (15) |
14–19 (17) |
16–18 (17) |
| Caudal rays |
20–23 (20) |
16–23 (20) |
17–23 (22) |
17–23 (20) |
18–24 (21) |
18–24 (21) |
| Pelvic rays |
5–7 (6) |
5–7 (6) |
5–7 (6) |
5–7 (6) |
5–7 (6) |
6–7 (6) |
| Lateral scales |
33–39 (37) |
30–39 (34) |
33–37 (36) |
29–34 (33) |
29–34 (33) |
30–33 (33) |
| Scales predorsal |
21–26 |
19–26 (23) |
20–24 (22) |
21–26 (23) |
18–24 (20) |
20–23 (22) |
| Scales around body |
28–36 |
25–35 (31) |
27–33 (30) |
25–31 (28) |
24–30 (26) |
26–29 (28) |
| Scales around peduncle |
18–22 (21) |
16–21 (18) |
16–21 (18) |
16–22 (19) |
17–25 (20) |
18–23 (19) |
| Vertebrae |
37–38 (37) |
33–36 (35) |
36–37 (36) |
33–34 (33) |
32–34 |
32–33 (33) |
| Gill rakers |
13–18 (16) |
14–19 (17) |
14–19 (16) |
13–17 (15) |
15–21 (17) |
15–19 (17) |
TABLE 2 |
(continued)
| Character |
P. oaxacae
35*
|
P. balsanus
22*
|
P. kreiseri
50*
|
P. parentiae
41*
|
P. mixtlanensis
60*
|
P. adani
71*
|
P. chimalapensis
38*
|
| Dorsal rays |
11–13 (12) |
11–13 (12) |
10–14 (12) |
10–13 (12) |
10–13 (11) |
10–12 (11) |
11–14 (13) |
| Anal rays |
13–15 (14) |
15–16 (15) |
12–16 (14) |
13–16 (15) |
13–16 (14) |
13–16 (14) |
14–16 (15) |
| Pectoral rays |
15–18 (16) |
15–17 (16) |
14–17 (16) |
15–17 (16) |
15–18 (16) |
15–18 (16) |
15–18 (17) |
| Caudal rays |
17–22 (20) |
18–20 (20) |
19–23 (21) |
18–22 (20) |
18––3 (21) |
18–23 (20) |
18–23 (20) |
| Pelvic rays |
6 (6) |
6 (6) |
5–6 (6) |
6–7 (6) |
6 (6) |
6–7 (6) |
6 (6) |
| Lateral scales |
29–31 (30) |
33 (33) |
32–35 (34) |
31–33 (33) |
31–34 (33) |
32–35 (33) |
29–32 (31) |
| Scales predorsal |
22–29 (25) |
21–24 (24) |
18–23 (21) |
18–22 (22) |
21–25 (22) |
23–28 (26) |
18–21 (19) |
| Scales around body |
23–31 (27) |
26–29 (28) |
23–30 (27) |
25–29 (29) |
26–34 (28) |
25–35 (30) |
25–30 |
| Scales around peduncle |
16–22 (19) |
18–21 (19) |
17–21 (19) |
17–21 (19) |
16–21 (17) |
15–18 (17) |
17–21 (20) |
| Vertebrae |
31–33 |
33–34 (33) |
33–34 (34) |
31–32 (32) |
33–34 (33) |
33–34 (33) |
33 |
| Gill rakers |
15–18 (16) |
14–18 (15) |
14–17 (16) |
14–22 (18) |
16–19 (18) |
16–20 (17) |
16–19 (18) |
FIGURE 5 |
Male general morphology and caudal fin squamation, life colour patterns in:
A.
Tlaloc labialis
; and
B.
Profundulus punctatus
. Solid arrow points to the squamation of the caudal fin, dashed arrow points to the humeral spot.
Sexual dimorphism and contact organs.
There is little sexual dimorphism in species of the genus
Tlaloc
. Males are slightly larger than females; the largest specimen recorded was a male from
Tlaloc hildebrandi
(
111.49 mm
SL), whereas the maximum size recorded in a female was in
Tlaloc labialis
(
101.93 mm
SL). The shape of the anal fin, however, shows a marked sexual difference: the anterior anal rays of the male are not greatly longer than the posterior ones, giving the distal margin of the fin an evenly rounded edge. However, in the female, since the medial rays, from about the sixth to the tenth, are much longer than either the anterior or the posterior rays, the distal margin of the fin is lobate when expanded. Males have contact organs or spinules, articulated with the lateral surfaces of the anal fin rays with some of them conspicuously long; this is the most notable, though not striking difference between males and females of all
Tlaloc
species
(
Fig. 7
).
Geographical distribution.
Restricted to the Atlantic slope of Middle America (
Fig. 8
). From the border between
Oaxaca
and
Chiapas
,
Mexico
, to the center of
Honduras
, with the exception of
Tlaloc portillorum
, this is located on both slopes of
Honduras
, Atlantic and Pacific.
Etymology.
Tlaloc
in reference to
Tlaloc
, a deity of water in Aztec mythology (Álvarez, Carranza, 1951).
FIGURE 6 |
Tlaloc candalarius
.
A.
Male (MZ-UNICACH 7567, 71.4 mm SL);
B.
Female (MZ-UNICACH 7567, 78.8 mm SL); note the orange coloration of the dorsal and anal fins in the male.
FIGURE 7 |
Bony spinules in fin rays of male
Tlaloc labialis
, indicated by arrow (MZ-UNICACH 6740, 75.7 mm SL), articulated with the lateral surfaces of the anal fin rays.
FIGURE 8 |
Geographical distribution of species of the genera
Tlaloc
and
Profundulus
in southern Mexico and Central America.
Nine species are recognized in the genus
Profundulus
(Morcillo
et al.
, 2016; Del Moral-Flores
et al.
, 2020; Domínguez-Cisneros
et al.
, 2021):
P. punctatus
(Günther, 1866)
,
P. guatemalensis
(Günther, 1866)
,
P. oaxacae
(Meek, 1902)
,
P. balsanus
,
P. kreiseri
Matamoros, Schaefer, Hernández & Chakrabarty, 2012
,
P. mixtlanensis
Ornelas-García, Martinez-Ramírez & Doadrio, 2015
,
P. parentiae
Matamoros, Dominguez-Cisneros, Velázquez-Velázquez & McMahan, 2018
,
P. chimalapensis
Del Moral-Flores, López-Segovia
& Hernández- Arellano, 2020, and
P. adani
Dominguez-Cisneros, Velázquez-Velázquez, McMahan & Matamoros, 2021
.