A taxonomic revision of the catshark genus Haploblepharus Garman 1913 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae)
Author
Human, Brett A.
text
Zootaxa
2007
1451
1
40
journal article
10.5281/zenodo.176248
a273c73d-75c7-456d-a815-a47a8768428b
1175-5326
176248
Haploblepharus fuscus
Smith, 1950
(
Fig. 7
,
Table 3
)
Haploblepharus fuscus
Smith, 1950
: 883
.
Haploblepharus edwardsii
:
Günther, 1870
: 401
(in part);
Smith, 1949
: 54
(in part).
Type
Series and Locality.
Holotype
,
RUSI
21, adolescent male
530 mm
TL
(measured as
495 mm
TL
in the current study), collected off East London, Eastern Cape,
South Africa
(by J. L. B. Smith?), approx.
33°00'S
27°55'E
, from the shore “in shallow water among rocks” using hook and line, in good condition, although there is a large gash present on the dorsal midline of the head above gill slits 3 to 5, and there are a number of patches where the skin has been damaged, particularly on the right trunk, that may be due to poor preservation (
Fig. 7
).
FIGURE 7.
Photographs of the holotype of
Haploblepharus fuscus
, RUSI 21, 530 mm TL adolescent male.
Paratype
, missing presumed lost, possibly never catalogued,
680 mm
TL
male, collected from Knysna, Western Cape,
South Africa
(by J. L. B. Smith?), approx.
34°03'S
23°02.5'E
, from the shore using hook and line.
Diagnosis.
The largest member of the genus,
H. fuscus
has a relatively stocky body at all stages of maturity, particularly in mature individuals; abdomen width 11.0% TL for the
holotype
(mean 11.4% TL); snout rounded in the
holotype
, not coming to a point, becoming more broadly rounded in larger individuals, head width at the posterior margin of the orbit 12.0% TL in the
holotype
(mean 12.2% TL); head strongly depressed, head height at the posterior margin of the orbit 6.8% TL for the
holotype
(mean 6.1% TL); trunk depressed, trunk height 8.9% TL and trunk width 12.1% TL in the
holotype
(mean 8.7% TL and 12.2% TL, respectively); claspers of mature males stout, inner length 6.9 times the base in the
holotype
(mean inner length 4.3 times base). The
holotype
of
H. fuscus
has 84 rows of teeth in the upper jaw (mean 71.2) and 83 rows of teeth in the lower jaw (mean 74.7). The
holotype
of
H. fuscus
has a total of 134 (mean 134.8) vertebral centra.
Haploblepharus fuscus
always has a chocolate brown or dull brown background colouration, occasionally with indistinct saddles, and occasionally with white spots present, rarely with dark spots present.
FIGURE 8.
Photographs of
Haploblepharus fuscus
specimens: A) RUSI 25925; B) SAM 32614, to show the presence of spots or saddles, respectively, in this species.
Description.
The morphometric and meristic data for
H. fuscus
are given in
Table 3
.
Holotype
, adolescent male
495 mm
TL (mean of all specimens examined in
Table 3
, including the
holotype
, see Study material):
H. fuscus
is a stocky bodied
Haploblepharus
shark, although juveniles are slender, with a relatively broad head, head width at the pectoral origin 4.06 (3.91) times the preoral length; head length 1.24 (1.28) times distance from snout tip to first gill slit; height of first gill slit 1.86 (1.82) times the height of the fifth gill slit; eye length 5.67 (4.14) times longer than spiracle length; basimandibular cartilage found at the symphysis of the Meckels cartilage in the lower jaw; mouth length 1.33 (1.47) times the prenarial length; mouth width 8.5 (6.27) times the upper labial furrow length; labial cartilages present; nasal lobes fused into a nasal flap that covers the excurrent apertures and extends to the mouth; interorbital width 1.19 (1.15) times the nasal flap width; head strongly depressed, head width at the posterior margin of the orbit 1.76 (2.0) times its height; head width 1.44 (1.60) times its height; trunk strongly depressed, trunk width 1.36 (1.40) times its height; abdomen depressed, abdomen width 1.29 (1.18) times its height; tail not depressed, tail width equal to its height; caudal peduncle strongly compressed, caudal peduncle width 0.70 (0.57) times its height; precaudal length 1.86 (1.79) times the distance from snout to first dorsal fin; dorsal fins rounded; height of first dorsal fin 0.96 (1.08) times that of the second dorsal fin; first dorsal fin length equal to the length of its anterior margin; second dorsal fin length 1.08 (1.04) times the length of its anterior margin; pectoral fin to pelvic fin space 1.14 (1.28) times the interdorsal space; pectoral and pelvic fins rounded; pectoral fin height 2.18 (1.98) times the height of the pelvic fin; pectoral fin length 0.91 (0.90) times the length of its anterior margin; pelvic fin length 1.35 (1.36) times the length of its anterior margin; claspers of mature males stout, clasper inner length 6.89 (4.33) times the base; anal fin to caudal fin space 1.48 (1.42) times the head height at the origin of the pectoral fin; length of anal fin base 1.19 (1.29) the length of the second dorsal fin base; anal fin length 1.37 (1.43) times the length of its anterior margin; distance from pectoral fin insertion to the midpoint of the first dorsal fin length 1.52 (1.53) times the caudal dorsal margin length. Vertebral counts: total 134 (128–141), 34 (34–40) monospondylous, 60 (52–62) precaudal diplospondylous and 40 (35–48) caudal diplospondylous vertebrae. Dental formula: upper jaw (left) 40 (23–43), (right) 44 (23–49); lower jaw (left) 43 (23–46), (right) 40 (25–46). Spiral valve turns: NA (9).
Size and sexual maturity.
Haploblepharus fuscus
is the largest
Haploblepharus
shark, both in terms of overall size and mass. In this study, males were found to be juvenile at
438 mm
TL to
460 mm
TL, adolescent at
495 mm
TL to
543 mm
TL, and mature at
550 mm
TL to
649 mm
TL. Females were found to be adolescent at
496 mm
TL to
568 mm
TL, and mature at
609 mm
TL to
631 mm
TL, no female juveniles were available for examination. There appears to be no sexual dimorphism in
H. fuscus
.
Colouration.
Haploblepharus fuscus
is the least patterned of the
Haploblepharus
sharks (
Figs. 7
&
8
). The
holotype
is a uniform chocolate brown, slightly paler ventrally, without saddles, spots, or any other markings (
Fig. 7
). In other
H. fuscus
individuals, the background dorsal colouration is most often uniform chocolate brown or dull grey brown with no markings, although sometimes individuals have small inconspicuous black spots (rarely) or inconspicuous white spots (
Fig. 8
A), never both; saddles variably present and inconspicuous when present (
Fig. 8
B), more conspicuous in smaller specimens, centre of saddle more orange than background with darker margins anteriorly and posteriorly, colour in the centre of the saddle dull, number of saddles and position on body highly variable, usually between 2 to 4, although never on the head; one individual had white spots and indistinct saddles. No markings on any fins. Ventral colouration uniform white, off white, cream or paler than the background dorsal colouration, pectoral and pelvic fin webs darker, anal fin with dorsal background colour.
Comparison with other species.
Haploblepharus fuscus
is the least patterned of the
Haploblepharus
sharks. The head is bluntly rounded as in
H. pictus
, and the body is depressed more so than in
H. edwardsii
and
H. kistnasamyi
. Claspers of mature males are equivalent in size to
H. kistnasamyi
and
H. pictus
, although longer and stouter than in
H. edwardsii
.
Haploblepharus fuscus
has the highest total tooth count of the
Haploblepharus
sharks.
Haploblepharus fuscus
is most similar to
H. pictus
in overall morphology, and distinguished from that species by having indistinct saddles and spots, when present, with spots and saddles rarely occurring together, more vertebrae and fewer teeth rows. The colouration of
H. fuscus
is distinct from
H. edwardsii
and
H. kistnasamyi
(except possibly for the juveniles of the latter species, which can be distinguished from
H. fuscus
in possessing obvious saddles; see
Fig. 13
), however,
H. fuscus
has also been confused with
H. pictus
.
Remarks.
Smith (1950)
made his original description of
H. fuscus
from two males, one
530 mm
TL from East London, Eastern Cape (
RUSI
21) and one
680 mm
TL from Knysna, Western Cape. The only reference to the designation of a
type
specimen made by Smith was on the illustration of the
530 mm
individual, to which he labelled “
type
”. This, in accordance to Articles 73.1.1 and 73.1.4 of the International Commission for Zoological Nomenclature (ICZN, 1999), establishes that individual (
RUSI
21) as the
holotype
. In accordance with Recommendation 73D (ICZN, 1999), the remaining specimen is a
paratype
of
H. fuscus
, however, from examination of the SAIAB collection catalogue, this specimen apparently no longer exists, and there are no specimens of
H. fuscus
in that collection with the locality of Knysna, suggesting that the
paratype
was never catalogued.
Bass
et al
. (1975)
and
Springer (1979)
refer to the two specimens described by Smith, but make no mention of the status of the
paratype
.
The only material examined by
Springer (1979)
for
H. fuscus
was SAM 24545, which he illustrated and gave morphometric data for. In the present study, this specimen was determined to be
H. pictus
, therefore the morphometric proportions given by Springer for
H. fuscus
are actually referable to
H. pictus
.
The dentition and denticles of
H. fuscus
are described and illustrated by
Smith (1950)
,
Bass
et al
. (1975)
, and
Compagno (1984b)
.
Bass
et al
. (1975)
found no evidence of sexual heterodonty in
H. fuscus
, and found the teeth to be quite different to other
Haploblepharus
in being longer and narrower.
The juveniles of this species are scarce (
Bass
et al
., 1975
; M. J. Smale, pers. comm.; current study), and there appears to be an unknown habitat that is used by
H. fuscus
for egg laying, and where juveniles spend that stage of their life history. Only two juvenile specimens could be found in the present study.
RUSI
6079 is a juvenile male
460 mm
TL, collected from Port Alfred, Eastern Cape (
33°36’S
26°54’E
), and SAM 32614 (
Fig. 8
B) is also a juvenile male
438 mm
TL, collected from Langebaan Lagoon, Western Cape (Klein Oostervaal farm). Both of these juveniles are much larger than the juveniles of other members of the genus.
Haploblepharus fuscus
is regularly caught by shore anglers, who often regard them as pests (
Compagno, 1984b; pers. obs.
). This species is also occasionally used in aquaria, however, there is no directed fishery for this species for the aquarium trade at present (pers. obs.).
The biology of this species is virtually unknown, which is somewhat disturbing given that this is an inshore shark and caught with relatively high frequency by shore anglers in the Eastern Cape (pers. obs.). The gathering of biological data for this species should be considered a priority given that it is an endemic with a restricted range, with a habitat preference that is in a zone that experiences significant fishing pressure.
Distribution.
Haploblepharus fuscus
is verified as occurring coastally, south of latitude 33°S in
South Africa
(
Fig. 9
).
Haploblepharus fuscus
is most commonly found in the Eastern Cape, from Storms River mouth to East London, however it is a rare visitor to the Western Cape and has been verified from a specimen collected in Langebaan Lagoon (SAM 32614), assuming that the locality given for this specimen is accurate.
Haploblepharus fuscus
almost certainly ranges north of East London, however, its northern limit on the east coast of
South Africa
is unknown. It apparently does not range as far north as Durban.
Bass
et al
. (1975)
record
H. fuscus
from southern kwaZulu-Natal, without specifics.
Bass
et al
. (1975)
and
Bass (1986)
also record a specimen from Bredasdorp, Western Cape (true locality probably Arniston, Stuisbaai or Cape Agulhas because Bredasdorp is more than
20 km
inland), which they did not examine, however this specimen was examined by the author and was determined to be
H. pictus
(SAM 24545).
FIGURE 9.
Distribution of
Haploblepharus fuscus
as identified by the current study. Red dots are locality records for the species, the light blue line represents the 200 m isobath and the dark blue line represents the 1000 m isobath.
TABLE 3.
Morphometric and meristic data for the holotype (HT) of
Haploblepharus fuscus
, plus sample size, mean and range for
H. fuscus
specimens (including the holotype). TL and WT are actual measurements in millimetres and grams, respectively. All other morphometric measurements are expressed as percentage of TL. Abbreviations as in Table 1.
HT N Mean Min Max HT N Mean Min Max to be continued.
| WT |
468.7 |
23 |
679.9 |
100.0 |
1210.0 |
P1B |
5.8 |
21 |
5.8 |
4.5 |
6.6 |
| TL |
495 |
23 |
554.5 |
340.0 |
649.0 |
P1H |
8.5 |
21 |
9.9 |
7.1 |
12.3 |
| PCL |
84.2 |
23 |
81.7 |
78.1 |
84.8 |
P1I |
6.9 |
21 |
6.8 |
5.6 |
8.4 |
| PD2 |
68.3 |
21 |
66.1 |
63.3 |
68.8 |
P1P |
10.1 |
21 |
10.4 |
8.9 |
12.0 |
| PD1 |
45.3 |
21 |
45.6 |
43.4 |
48.8 |
P1R |
6.6 |
21 |
7.7 |
6.3 |
10.2 |
| BDL |
65.9 |
21 |
63.4 |
59.1 |
66.7 |
D1L |
10.0 |
21 |
10.6 |
8.6 |
12.0 |
| IDS |
14.9 |
21 |
13.4 |
11.3 |
16.8 |
D1A |
10.1 |
21 |
10.5 |
8.8 |
12.2 |
| D2C |
29.3 |
21 |
27.1 |
25.8 |
29.3 |
D1B |
6.6 |
21 |
7.4 |
5.5 |
8.4 |
| DCS |
9.5 |
20 |
8.6 |
6.9 |
10.3 |
D1H |
4.4 |
21 |
5.2 |
4.3 |
6.3 |
| PAL |
54.5 |
21 |
57.4 |
54.5 |
59.9 |
D1I |
3.4 |
21 |
3.2 |
2.8 |
4.4 |
| PP2 |
36.0 |
21 |
38.7 |
35.7 |
42.0 |
D1P |
4.8 |
21 |
5.2 |
3.7 |
6.0 |
| PP1 |
14.5 |
21 |
17.9 |
14.5 |
20.5 |
P2L |
11.2 |
21 |
11.7 |
10.2 |
12.6 |
| SVL |
39.2 |
21 |
40.5 |
36.5 |
43.0 |
P2A |
8.3 |
21 |
8.6 |
6.3 |
10.4 |
| TRL |
21.4 |
21 |
22.3 |
19.4 |
25.1 |
P2B |
5.2 |
21 |
6.6 |
5.2 |
8.1 |
| PPS |
17.0 |
21 |
17.2 |
14.1 |
19.2 |
P2H |
3.9 |
21 |
5.0 |
3.4 |
6.8 |
| PAS |
14.5 |
20 |
13.3 |
11.6 |
14.6 |
P2I |
5.1 |
21 |
5.6 |
3.9 |
7.4 |
| VCL |
64.0 |
21 |
60.4 |
56.5 |
64.1 |
P2P |
7.8 |
21 |
7.4 |
6.2 |
9.0 |
| PCA |
35.5 |
21 |
34.3 |
29.7 |
36.8 |
CLO |
2.1 |
12 |
6.6 |
2.0 |
9.0 |
| ACS |
13.3 |
21 |
11.8 |
10.0 |
14.0 |
CLI |
6.2 |
14 |
10.4 |
4.4 |
13.2 |
| HDH2 |
6.8 |
21 |
6.1 |
5.2 |
7.5 |
CLB |
0.9 |
12 |
2.4 |
0.7 |
7.6 |
| HDW2 |
12.0 |
21 |
12.2 |
9.8 |
13.1 |
D2L |
11.1 |
21 |
11.1 |
9.7 |
12.4 |
| INO |
6.4 |
21 |
6.1 |
5.5 |
6.6 |
D2A |
10.3 |
21 |
10.7 |
8.7 |
11.9 |
| HDH |
9.0 |
21 |
8.3 |
7.2 |
10.2 |
D2B |
8.3 |
21 |
8.0 |
6.7 |
9.2 |
| HDW |
13.0 |
21 |
13.3 |
8.9 |
14.7 |
D2H |
4.6 |
21 |
4.8 |
3.4 |
6.2 |
| TRH |
8.9 |
21 |
8.7 |
6.2 |
10.8 |
D2I |
2.6 |
21 |
3.1 |
2.6 |
3.7 |
| TRW |
12.1 |
21 |
12.2 |
9.8 |
16.6 |
D2P |
4.8 |
21 |
5.2 |
4.2 |
6.2 |
| GIR |
34.9 |
21 |
35.0 |
30.4 |
42.9 |
ANL |
11.8 |
21 |
12.9 |
11.7 |
14.3 |
| ABH |
8.5 |
21 |
9.7 |
6.8 |
11.4 |
ANA |
8.7 |
21 |
9.0 |
7.4 |
10.3 |
| ABW |
11.0 |
21 |
11.4 |
7.9 |
15.1 |
ANB |
9.9 |
21 |
10.3 |
8.8 |
11.6 |
| TAH |
7.1 |
21 |
6.4 |
5.8 |
7.4 |
ANH |
4.0 |
21 |
4.4 |
3.5 |
5.0 |
| TAW |
7.1 |
21 |
6.4 |
5.5 |
7.6 |
ANI |
2.5 |
21 |
2.6 |
2.1 |
3.3 |
| CPH |
3.7 |
21 |
3.7 |
3.3 |
4.1 |
ANP |
5.6 |
21 |
6.5 |
5.1 |
7.9 |
| CPW |
2.6 |
21 |
2.1 |
1.7 |
2.6 |
CDM |
19.2 |
21 |
18.7 |
17.3 |
20.1 |
| HDL |
18.0 |
21 |
18.6 |
16.9 |
20.2 |
CPV |
9.3 |
21 |
10.2 |
8.9 |
12.0 |
| PG1 |
14.5 |
21 |
14.5 |
12.3 |
15.4 |
CPU |
9.7 |
21 |
9.0 |
7.3 |
10.7 |
| PSP |
9.1 |
21 |
9.0 |
8.1 |
9.4 |
CST |
5.1 |
21 |
4.4 |
3.4 |
5.5 |
| POB |
5.8 |
21 |
5.9 |
5.1 |
6.3 |
CSW |
3.1 |
21 |
2.9 |
2.1 |
3.2 |
| EYL |
3.4 |
21 |
2.9 |
2.5 |
3.4 |
CTR |
4.4 |
21 |
4.6 |
3.5 |
5.6 |
| EYH |
1.3 |
21 |
1.5 |
1.1 |
1.7 |
CTL |
5.5 |
21 |
5.4 |
5.0 |
5.8 |
|
TABLE 3
(continued).
|
| HT N |
Mean |
Min |
Max |
HT |
N |
Mean |
Min |
Max |
| SOD 0.2 17 |
0.3 |
0.1 |
0.5 |
DPI |
29.1 |
21 |
28.6 |
24.5 |
30.8 |
| SPL 0.6 21 |
0.7 |
0.4 |
0.9 |
DPO |
11.7 |
21 |
11.2 |
1.1 |
14.3 |
| ESL 0.3 21 |
0.4 |
0.2 |
0.6 |
PDI |
7.7 |
21 |
7.4 |
5.8 |
8.8 |
| EMA 1.5 21 |
1.4 |
0.4 |
2.5 |
PDO |
22.8 |
21 |
20.6 |
17.8 |
24.9 |
| ING 3.7 21 |
4.4 |
3.6 |
5.4 |
DAO |
7.8 |
21 |
7.5 |
5.5 |
10.9 |
| GS1 2.6 21 |
3.1 |
2.4 |
3.6 |
DAI |
6.1 |
21 |
5.1 |
4.0 |
6.2 |
| GS2 2.3 21 |
2.8 |
2.2 |
3.4 |
ANFL |
2.1 |
21 |
2.2 |
1.8 |
2.4 |
| GS3 2.1 21 |
2.6 |
2.1 |
3.0 |
ANFM |
1.3 |
21 |
1.6 |
1.1 |
2.2 |
| GS4 1.8 21 |
2.3 |
1.8 |
2.7 |
ANFW |
5.4 |
21 |
5.3 |
3.4 |
5.8 |
| GS5 1.4 21 |
1.7 |
1.3 |
2.2 |
P2 fused |
0 |
11 |
0.9 |
0.0 |
1.7 |
| POR 3.2 21 |
3.4 |
3.1 |
3.8 |
P2 free |
4.5 |
11 |
5.3 |
4.5 |
6.0 |
| PRN 1.8 21 |
1.9 |
1.6 |
2.3 |
VERT |
134 |
17 |
134.8 |
128 |
141 |
| MOL 2.4 21 |
2.8 |
2.0 |
3.5 |
mono |
34 |
17 |
35.6 |
34 |
40 |
| MOW 8.5 21 |
9.4 |
6.6 |
11.0 |
predip |
60 |
17 |
57.2 |
52 |
62 |
| ULA 1.0 21 |
1.5 |
1.0 |
2.0 |
caudaldip |
40 |
17 |
41.9 |
35 |
48 |
| ULH 0.9 21 |
1.2 |
0.7 |
2.7 |
UL |
40 |
6 |
37.7 |
23 |
43 |
| LLA 1.6 21 |
2.6 |
0.9 |
3.2 |
UR |
44 |
5 |
40.2 |
23 |
49 |
| NOW 2.1 21 |
2.3 |
2.0 |
2.6 |
U tot |
84 |
6 |
71.2 |
40 |
90 |
| INW 2.1 21 |
1.8 |
1.5 |
2.3 |
LL |
43 |
6 |
37.2 |
23 |
46 |
| IOW 6.5 21 |
6.7 |
5.5 |
7.4 |
LR |
40 |
6 |
37.5 |
25 |
46 |
| P1L 11.7 21 |
12.1 |
10.5 |
13.0 |
L tot |
83 |
6 |
74.7 |
48 |
92 |
| P1A 12.8 21 |
13.4 |
11.9 |
15.0 |
Spiral valve |
- |
1 |
9.0 |
9 |
9 |
Etymology.
Although
Smith (1950)
did not give the etymology in his original description, there is little doubt that the specific name comes from the Latin,
fuscus
, which means dusky, dark, or swarthy, and is in reference to the general drab brown colour of this species.
Common name.
In an attempt to introduce species specific common names for this genus, this shark was given the name plain happy in
Compagno &
Human
(2003)
. It is also known as the brown shyshark.
Study material.
BAH
20020304.05, male
340 mm
TL
, Hamburg, Eastern Cape,
South Africa
,
33°17.2'S
27°28.9'E
;
BAH
20020304.06, male
580 mm
TL
, Hamburg, Eastern Cape,
South Africa
;
LJVC
820916, gravid female
631 mm
TL
, Port Elizabeth, Eastern Cape,
South Africa
,
33°53'S
25°39'E
; MJS 941004, mature female
625 mm
TL
, Cape Recife, Eastern Cape,
South Africa
,
34°01.7'S
25°42.1'E
; MJS 990217, mature female
609 mm
TL
, Cape Recife, Eastern Cape,
South Africa
;
RUSI
21
holotype
of
Haploblepharus fuscus
, see under
Type
Series and Locality for details;
RUSI
3701, mature male
605 mm
TL
, Cape Padrone, Eastern Cape,
South Africa
,
33°46'S
26°28'E
;
RUSI
6079, previously
ORI
2470, juvenile male
460 mm
TL
, Port Alfred, Eastern Cape,
South Africa
,
33°36'S
26°54'E
;
RUSI
6081, previously
ORI
2787, mature female
610 mm
TL
, Port Alfred, Eastern Cape,
South Africa
;
RUSI
6082, previously
ORI
2471, adolescent male
543 mm
TL
, Port Alfred, Eastern Cape,
South Africa
;
RUSI
7617, adolescent female
568 mm
TL
, Cape Padrone, Eastern Cape,
South Africa
;
RUSI
10289, mature male
551 mm
TL
, Algoa Bay, Eastern Cape,
South Africa
,
34°02'S
25°42'E
;
RUSI
12826, mature male
602 mm
TL
, locality not recorded;
RUSI
13144,
2 specimens
, mature male
649 mm
TL
, Algoa Bay, Eastern Cape,
South Africa
;
RUSI
14005,
2 specimens
, one of which is referable to
H. kistnasamyi
,
H. fuscus
specimen is a mature male
637 mm
TL
, Cape Recife, Eastern Cape,
South Africa
;
RUSI
19993,
3 specimens
, 2 adolescent females
496 mm
TL
and
540 mm
TL
, mature male
550 mm
TL
, locality not recorded;
RUSI
25182, mature male
577 mm
TL
, Paradise Beach, Eastern Cape,
South Africa
,
34°07'S
24°52.5'E
;
RUSI
25925, gravid female
571 mm
TL
, Cape Recife, Eastern Cape,
South Africa
;
RUSI
41963, mature male
559 mm
TL
, Fish River Lighthouse, Eastern Cape,
South Africa
,
33°29'S
27°08'E
;
SAM
32523,
4 specimens
, Storms River Mouth, Eastern Cape,
South Africa
,
34°01.3'S
23°54.7'E
;
SAM
32614, immature male
438 mm
TL
, Langebaan Lagoon (Klein Ostervaal farm), Western Cape,
South Africa
,
33°04.5'S
18°02.5'E
.