A review of the flying fish genus Cypselurus (Beloniformes: Exocoetidae). Part 1. Revision of the subgenus Zonocypselurus Parin and Bogorodsky, 2011 with descriptions of one new subgenus, four new species and two new subspecies and reinstatement of one species as valid
Author
Shakhovskoy, Ilia B.
Author
Parin, Nikolay V.
text
Zootaxa
2019
2019-03-23
4589
1
1
71
journal article
22545
10.11646/zootaxa.4589.1.1
375480cc-1f64-4e37-ba9d-a087e7cb34ef
1175-5326
3772209
860E8CF0-A245-46DA-9023-B0BD825730D0
Cheilopogon
(
Ptenichthys
)
katherinae
sp. nov.
urn:lsid:zoobank.org:act:
FA5312DB-2248-460B-A087-04072418261F
(
Figs. 18
,
20–22
; Tables 1–7, 10)
Cypselurus
sp. Ben Tuvia & Steinitz 1952 (cited by:
Fowler & Steinitz 1956
): 5 (Eylath,
Red Sea
).
Cypsilurus socotranus
(non Steindachner).
Fowler & Steinitz 1956: 271–272
(description; Eylath,
Red Sea
).
Cypselurus hexazona
(non Bleeker).
Parin & Bogorodsky 2011: 683–686
[
658–660 in
English translation] (in part;
Red Sea
).
Material examined.
Fourteen specimens
51–212 mm
SL.
Full morphological study
.
ZMMU
P-24081,
holotype
(
150.5 mm
SL),
19°39’N
38°45’E
,
11.06.1977
.
IORAS 03432
(1, 51 mm SL),
17°43’N
39°20’E
,
8.06.1977
.
IORAS 03433
(
1, 174.5 mm
SL),
24°49’N
35°46’E
,
21.10.1988
.
IORAS 03434
,
paratypes
(3, 139, 139 and
142 mm
SL),
19°39’N
38°45’E
,
11.06.1977
.
Partial morphological study
.
IORAS
uncat.* (3, 185–
197 mm
SL),
14°05’N
42°20’E
,
15.10.1960
.
IORAS
uncat.* (
1, 187 mm
SL),
19°40’N
38°41’E
, 13-
14.10.1960
.
IORAS
uncat.* (
1, 212 mm
SL),
12°22’N
43°59’E
,
5.03.1983
.
SMF 213
* (
1, 181 mm
SL),
Rotes Meer
.
USNM 294827
(
1, 112 mm
SL),
21°34’N
38°41’E
,
17.05.1958
.
ANSP
101711
6
(1, ~
192 mm
SL),
Eylath
,
Red Sea
.
Holotype
(
Figs. 20a, b
).
ZMMU
P-24081,
R
/
V
"Vityaz", Cruise 61, Sta. 7630,
19°39’N
38°45’E
, captured with night light on
11.06.1977
. Female (immature),
150.5 mm
SL.
D
12,
A
9,
P
I 14,
Spred
31,
Str
9,
Sp.br
22 (5 + 17),
Vert
44 (30 + 14). Measurements in % SL:
aA
78.5,
aD
71.2,
aV
56.0,
cV
32.0,
pV
41.9,
c
23.9,
po
10.3,
o
7.4,
ao
4.7,
io
8.0,
Hc
17.2,
H
18.5,
h
7.1,
Dc
26.7,
lP
64.2,
lP
1
34.6+,
lV
34.0,
lD
17.4,
lA
10.7,
HD
11.2,
HA
6.7,
p
13.9. Pectoral fins grayish brown with pale “mirror” to 4th ray. Tip of fin reaches 10th dorsal-fin ray. Pelvic fins transparent, their tip reaching end of anal-fin base. Dorsal fin is gray, tip of last ray reaching middle of caudal peduncle. Penultimate dorsal-fin rays not elongate. Anal fin transparent; first anal-fin ray beneath 6th–7th dorsalfin ray. Longest ray of dorsal fin the 2nd; 2nd and 3rd rays of anal fin longest. Caudal fin is grayish brown. Remnants of two pigment bands ventrally on body: in middle of pecto-ventral distance and near pelvic-fin base. Lower jaw longer than upper, teeth small, conical, some with additional cusps. No palatine teeth.
Paratypes
.
IORAS 03434
, three specimens 139, 139 and
142 mm
SL
,
R
/
V
"Vityaz", Cruise 61, Sta. 7630,
19°39’N
38°45’E
,
11.06.1977
.
Description.
The meristic and morphometric characters are given in Tables 1–7 and 10.
D
11–13,
A
8–10,
P
I 12–14 (usually I 14),
Spred
30–34,
Str
8–9½,
Sp.br
20–24 (5–7 + 15–18),
Vert
44–46 (29–30 + 14–17). Palatine teeth absent (only 1 out of
8 specimens
with row of 3–4 teeth on anterior of palatines). Snout not oblong, upper jaw is slightly pointed at end. Lower jaw longer than upper or, rarely, equal. Jaw teeth very small, usually unicuspid, conical. Occasional specimens with some teeth with additional cusps.
Body of fish
139–212 mm
SL elongate, greatest body depth 5.2–6.0 in SL. Body width 1.10–1.34 and depth of caudal peduncle
2.23–2.74 in
greatest body depth. Greatest head depth and head length 5.8–6.25 and 4.0–
4.25 in
SL, respectively. Head length
1.01–1.16 in
dorso-caudal distance. Eyes large, their diameter 12.2–14.3 times in SL,
2.55–3.45 in
c
and 1.0–
1.15 in
io
.
Pectoral fins relatively long, their length increasing somewhat with growth: in fish
139–150.5 mm
SL tip of fin reaches (or almost reaches) end of dorsal-fin base and
lP
1.54–1.6 in
SL; in larger fish (
174.5–212 mm
SL) tip of pectoral fin reaches from end of dorsal-fin base to middle of caudal peduncle, and
lP
1.44–1.51 in
SL. First pectoral-fin ray unbranched, its length almost not changing with growth,
2.65–2.95 in
SL and
1.75–1.92 in
lP
. Pelvic-fin base much closer to posterior edge of head than to beginning of lower lobe of caudal fin (
cV/pV
= 0.74– 0.79). The length of pelvic fins
2.8–3.3 in
SL and
1.85–2.3 in
lP
. Tip of pelvic fins reaches (almost reaches) end of anal-fin base or protrudes slightly beyond it.
Anal-fin origin far behind dorsal-fin origin, 1st anal-fin ray beneath 5th–7th dorsal-fin ray (usually 5th or 6th). Dorsal fin with 2–4 rays more than anal fin. Heights of dorsal and anal fins 8.3–10.1 and
13.3–16.4 in
SL, respectively. Longest ray in dorsal and anal fins 2nd or 3rd. Tip of last dorsal-fin ray reaches middle of caudal peduncle. Central and posterior rays of dorsal fin not elongate.
Pigmentation.
Fish
139–212 mm
SL have the typical pelagic coloration: dark from above and pale from below. There are no body bands. Only some fish
139–150 mm
SL show remnants of two bands: near pelvic-fin base and in middle of pecto-ventral distance. Gill covers usually with few small specks or large dots; absent below eye.
6. This specimen was studied by the first author basing only on photographs and X-ray photographs (by the courtesy of K.E. Bemis (VIMS)).
FIGURE 19.
Flying fish
Cheilopogon
(
Ptenichthys
)
sp.2. (A)—31 mm SL, AMS I.26494-013, 14°39’S 145°26’E; (B)—38.5 mm SL, AMS I.26529-008, 14°40’S 145°27’E; (C)—48 mm SL, AMS I.19461-060, 14°40’S 145°28’E; (D)—67 mm SL, AMS I.23902-001, 32°28’S 152°05’E; (E)—the same specimen, close up of head.
FIGURE 20.
Flying fish
Cheilopogon katherinae
. (A)—150.5 mm SL, ZMMU P-24081 (holotype), 19°39’N 38°45’E, lateral view; (B)—the same specimen, pectoral fin; (C)—142 mm SL, IORAS 03434 (paratype), 19°39’N 38°45’E, pelvic fin.
Pectoral fins (
Fig. 20b
) grayish-brown or gray with large pale “mirror” reaching to 3rd–4th (sometimes to 6th) ray, and usually with narrow pale posterior edge. In the smallest specimen of the
type
series (
139 mm
SL) the “mirror” reaching upper edge of fin. Pelvic fins transparent, and only some fish retaining remnants of pigmentation distally along the 3rd–4th rays (
Fig. 20c
). Dorsal fin gray; in fish
139–142 mm
SL its upper margin blackish. Anal fin transparent. Caudal fin pale brown, and in fish
139–150 mm
SL the posterior margin pale.
Coloration in life.
No data.
Maximum size.
The maximum length (SL) of
Ch. katherinae
in our materials was
212 mm
(IORAS uncat.,
12°22’N
43°59’E
). The only male available at our disposal was
174.5 mm
SL. The length of the largest female studied for sex and maturity was
187 mm
SL.
Juveniles.
Two juveniles 51 and
112 mm
SL (the latter specimen in poor condition) were preliminarily identified as
Ch. katherinae
(
Fig. 21
). A description of the smaller juvenile with some data for the larger specimen is given below.
The
51 mm
SL juvenile is similar to large fish in measurements of body and fins (Table 10), worth noting only a higher value of body width (15.8% SL), more anterior position of pelvic fins (
cV/pV
= 0.68), much longer pelvic fins (44.9% SL) extending beyond origin of caudal fin lower lobe, and higher dorsal fin (17.2% SL), the penultimate ray of which is longer than the last one and reaches the beginning of the upper lobe of the caudal fin.
The jaw teeth are small and conical; palatine teeth absent. There is a pair of short, pointed barbels on the chin (
Fig. 21b
), nearly reaching the middle of the eye. The barbels consist of a thick stem with a narrow fringe along the outer (posterior) margin. The barbels are uniformly covered with dots and only near the base they are pale. In the
112 mm
SL juvenile barbels are absent.
There are four distinct dark bands on the body (
Fig. 21a
): near the bases of pectoral and pelvic fins, between them and above the anus. The underside of the head is pale with only isolated dots below the eye; the gill covers are densely pigmented. The snout end and the space around the nostrils are covered with dots.
FIGURE 21.
Juvenile of flying fish
Cheilopogon katherinae
(51 mm SL, IORAS 03432, 17°43’N 39°20’E). (A)—lateral view; (B)—close-up of barbel; (C)—pectoral fin; (D)—pelvic fin.
The pectoral fins are mottled (
Fig. 21c
), their proximal part and the lower edge of the distal part dark, with two rows of darker spots and with a paler band near the fin base. Distally the fins are generally pale with several dark streaks along the rays. In the
112 mm
SL juvenile the pectoral fins are gray proximally and brown distally, with a pale “mirror” to the 5th–6th rays.
The pelvic fins (
Fig. 21d
) are brown with faded (paler) areas along the rays. The
112 mm
SL juvenile has a similar pattern of pigmentation, but its intensity is noticeably less—the fin grayish.
The dorsal fin is covered with dots forming three oblique stripes: near the bases of the anterior rays, in central and postero-superior parts of the fin. The last ray is pale. The anal fin is brownish anteriorly between the 1st–3rd rays and there is a cluster of dots (forming a faint spot) on the rear of the fin, between the 4th–8th rays. In the
112 mm
SL juvenile the dorsal fin is pale with some feint pigmentation along its upper margin; the anal fin is transparent.
The caudal fin is pigmented near its base and along the lower lobe rays, where there are two bands. In the
112 mm
SL juvenile the caudal fin is pale brown.
Intraspecific variation.
Specimens of the
type
series of
Ch. katherinae
139–150.5 mm
SL differ slightly from larger fish (
174.5–212 mm
SL) in shorter pectoral fins (62.4–64.8 vs 66.1–69.2% SL). We consider this difference a consequence of allometric growth, although it is also possible that the large specimens belong to another, undescribed, species. Furthermore, one specimen (IORAS 03433,
174.5 mm
SL,
24°49’N
35°46’E
) has a row of 3– 4 teeth on the palatines, while the other seven specimens examined had no palatine teeth. We regard this as intraspecific variation, although it is possible that this specimen may also belong to another species. To resolve these issues it is necessary to study a more representative collection of
Ch. katherinae
.
Etymology.
The species is named in honor of Katherine Elliott Bemis (VIMS) to acknowledge her kind help with our work on this review.
Common names.
The name “Katherine’s
Red Sea
flying fish” (Russian: “кpacнoмopcкий длиннoкpыл Кaтepины”) is proposed here.
Comparative remarks.
Cheilopogon katherinae
differs from
Ch. intermedius
occurring in Indo-West Pacific (for a description of this species see
Parin 1961a
; Parin & Lakshminayana 1993; Lakshminarayana 1993) in the pale pigmentation of pelvic fins, a smaller number of anal-fin rays (usually 9 vs usually 10), larger values of
Spred
,
Str
,
pV
and smaller values of
cv/pV
and
cV
. The
51 mm
SL juvenile differs from the juveniles of
Ch. intermedius
of similar size (see
Parin 1961a
;
Kovalevskaya 1975
) in paler chin barbels with a narrower fringe, in darker pelvic and anal fins and base of pectoral fins, as well as in larger values of
c
,
o
,
lP
and
lD
.
Cheilopogon katherinae
differs from
Ch. furcatus
(see
Parin & Belyanina (1998)
for a description of this species) in pigmentation of pectoral, pelvic, dorsal and anal fins, in a lower number of rays in dorsal, anal and pectoral fins, in smaller values of
lD
and
lA
. It differs from the Atlantic species
Ch. melanurus
(description of this species is given in
Parin & Belyanina (2000))
in fewer rays in pectoral (13–15 vs. usually 16–17) and dorsal fins, higher value of
aV
and lower values of
Dc
,
lD
and
HA
.
Cheilopogon katherinae
is probably most closely related to
Ch. melanurus
or
Ch. intermedius
, because it is most similar to these species in pigmentation and morphometry. However, this issue requires a special study.
Cheilopogon katherinae
can be easily confused with
Ch. marisrubri
, the other species occurring in the
Red Sea
. These two species have very similar pigmentation as adults. However, they can be distinguished by the length of the pelvic fins (
Fig. 18
), the number of vertebrae and transverse scales (
Str
) and by other characters (see Tables 1–7).
FIGURE 22.
Distribution of flying fishes
Cheilopogon katherinae
(Δ),
Cypselurus bosha
(□) and
C. olpar
(○). Filled symbols indicate capture location for fishes ≤100 mm SL, open symbols indicate capture location for fishes>100 mm SL.
TA
BLE
10.
Meristic and morphometric characters of flying fish
Cheilopogon katherinae
№ |
1
|
2
|
3
|
4
|
5
|
6
|
7
|
8
|
9
|
10
|
11
|
12
|
13
|
14
|
SL |
51 |
112 |
139
|
139
|
142
|
150.5
|
174.5 |
181 |
185 |
186 |
187 |
~192 |
197 |
212 |
D
|
12 |
12 |
12
|
11
|
12
|
12
|
13 |
12 |
13 |
13 |
12 |
12 |
12 |
12 |
A
|
8 |
9 |
9
|
9
|
9
|
9
|
10 |
8 |
9 |
9 |
9 |
9 |
9 |
9 |
P
|
15 |
15 |
13
|
15
|
15
|
15
|
15 |
14 |
14 |
14 |
15 |
15 |
14 |
14 |
Spred
|
30 |
- |
32
|
30
|
31
|
31
|
31 |
30 |
34 |
33 |
31 |
31 |
34 |
30 |
Str
|
9 |
- |
9
½
|
8
|
9
½
|
9
|
8 |
8 |
8 |
8½ |
8 |
- |
8 |
- |
Sp.br
|
22 |
24 |
24
|
21
|
23
|
22
|
24 |
22 |
20 |
21 |
21 |
- |
- |
23 |
Vert
|
- |
- |
45
|
46
|
45
|
44
|
- |
44 |
- |
- |
- |
45 |
- |
- |
aA
|
76.6 |
- |
78.2
|
77.6
|
77.4
|
78.5
|
77.4 |
- |
81.3 |
79.8 |
80.2 |
- |
80.1 |
81.6 |
aD
|
69.0 |
- |
71.3
|
71.2
|
70.5
|
71.2
|
71.0 |
- |
73.2 |
72.0 |
72.7 |
- |
72.2 |
73.1 |
aV
|
55.0 |
- |
56.9
|
56.4
|
55.8
|
56.0
|
56.2 |
- |
60.5 |
58.0 |
57.7 |
- |
58.3 |
59.9 |
cV
|
29.4 |
- |
32.6
|
32.3
|
32.9
|
32.0
|
31.9 |
- |
- |
- |
- |
- |
- |
- |
cV
1
|
- |
- |
-
|
-
|
-
|
-
|
-
|
36.1 |
36.7 |
33.2 |
34.6 |
- |
35.4 |
36.3 |
pV
|
43.1 |
- |
41.8
|
42.5
|
41.4
|
41.9
|
43.0 |
40.3 |
40.5 |
41.3 |
42.7 |
- |
40.3 |
40.5 |
c
|
25.2 |
- |
24.4
|
24.1
|
23.5
|
23.9
|
25.1 |
24.4 |
24.8 |
24.7 |
24.6 |
23.7 |
24.3 |
24.0 |
po
|
11.5 |
- |
10.5
|
10.7
|
10.6
|
10.3
|
10.4 |
- |
11.2 |
11.0 |
- |
- |
10.4 |
- |
o
|
9.8 |
- |
8.0
|
7.3
|
7.6
|
7.4
|
7.9 |
8.2 |
7.2 |
7.5 |
7.4 |
- |
7.3 |
7.0 |
ao
|
4.1 |
- |
5.8
|
5.1
|
4.3
|
4.7
|
5.2 |
- |
- |
- |
- |
- |
- |
- |
ao
1
|
- |
- |
-
|
-
|
-
|
-
|
- |
- |
7.3 |
7.6 |
7.0 |
- |
7.3 |
6.6 |
io
|
10.1 |
- |
8.5
|
8.2
|
8.4
|
8.0
|
8.1 |
- |
- |
- |
- |
- |
- |
- |
io
1
|
- |
- |
-
|
-
|
-
|
-
|
- |
8.4 |
8.9 |
8.2 |
7.8 |
- |
8.2 |
7.7 |
Hc
|
17.2 |
- |
16.6
|
16.4
|
16.9
|
17.2
|
16.0 |
- |
- |
- |
- |
- |
- |
- |
H
|
17.4 |
- |
18.0
|
17.9
|
18.6
|
18.5
|
16.7 |
17.1 |
19.1 |
18.4 |
19.2 |
- |
17.5 |
17.6 |
h
|
7.8 |
- |
7.2
|
6.6
|
7.2
|
7.1
|
6.8 |
6.6 |
7.0 |
6.9 |
7.0 |
- |
6.8 |
6.6 |
Dc
|
27.2 |
- |
26.1
|
26.7
|
27.3
|
26.7
|
27.5 |
25.1 |
26.5 |
26.3 |
26.4 |
- |
27.1 |
25.0 |
lP
|
64.9 |
- |
62.4
|
64.8
|
62.8
|
64.2
|
68.7 |
67.4 |
69.2 |
66.1 |
68.9 |
- |
67.0 |
68.3 |
lP
1
|
37.0 |
- |
34.8
|
35.8
|
33.9
|
34.6+
|
- |
36.4 |
37.6 |
35.7 |
35.8 |
- |
36.0 |
- |
lV
|
44.9 |
- |
33.1
|
33.3
|
33.8
|
34.0
|
33.7 |
35.9 |
32.5 |
31.9 |
31.3 |
- |
31.0 |
30.1 |
lD
|
19.8 |
- |
18.0
|
17.2
|
17.4
|
17.4
|
18.9 |
- |
17.8 |
17.7 |
17.6 |
- |
17.4 |
17.4 |
lA
|
11.7 |
- |
11.0
|
11.6
|
10.2
|
10.7
|
12.4 |
- |
10.7 |
10.4 |
10.4 |
- |
10.2 |
10.3 |
HD
|
17.2 |
- |
11.3
|
10.2
|
10.6
|
11.2
|
10.8 |
- |
9.9 |
12.0 |
10.2 |
- |
11.0 |
11.3 |
HA
|
- |
- |
-
|
6.4
|
6.1
|
6.7
|
6.9 |
- |
7.4 |
7.5 |
7.1 |
- |
6.7 |
- |
p
|
15.8 |
- |
14.3
|
14.2
|
14.2
|
13.9
|
14.5 |
13.2 |
14.9 |
14.2 |
14.3 |
- |
14.0 |
14.1 |
lcir
|
10.1 |
- |
-
|
-
|
-
|
-
|
- |
- |
- |
- |
- |
- |
- |
- |
Note to Table 10: No. 1 – IORAS 03432, No. 2 - USNM 294827, Nos. 3-5 – IORAS 03434 (paratypes), No. 6 – ZMMU P-24081 (holotype), No. 7 – IORAS 03433, No. 8 - SMF 213, Nos. 9, 10 and 13 – IORAS uncat., 14°05’N 42°20’E, No. 11 – IORAS uncat., 19°40’N 38°41’E, No. 12 - ANSP 101711, No. 14 – IORAS uncat., 12°22’N 43°59’E. The specimens of the type series are printed in bold.
Biology.
All the specimens of this species studied (
150–187 mm
SL) were immature.
Distribution.
Cheilopogon katherinae
is endemic to the
Red Sea
(
Fig. 22
), where it is widely distributed, penetrating northward to Eylath (Eilat) (ANSP 101711, ~
192 mm
SL,
December 1949
). This species has also been collected in the western part of the
Gulf
of Aden (IORAS uncat.,
212 mm
SL,
12°22’N
43°59’E
,
5.03.1983
).