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Redescription of Hydrolagus africanus (Gilchrist, 1922) (Chimaeriformes:

Chimaeridae), with a review of southern African chimaeroids and a key to
their identification

Article in African Journal of Marine Science · July 2015

DOI: 10.2989/1814232X.2015.1033012

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Redescription of Hydrolagus africanus (Gilchrist, 1922)

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(Chimaeriformes: Chimaeridae), with a review of southern
African chimaeroids and a key to their identification
a abc bd e
KA Walovich , DA Ebert , DJ Long & DA Didier
a
Pacific Shark Research Center, Moss Landing Marine Laboratories, Moss Landing, California, USA
b
Department of Ichthyology, Institute for Biodiversity Science and Sustainability, California
Academy of Sciences, San Francisco, California, USA
c
South African Institute for Aquatic Biodiversity, Grahamstown, South Africa
d
Department of Biology, St Mary's College, Moraga, California, USA
e
Department of Biological Sciences, Millersville University, Millersville, Pennsylvania, USA
Published online: 24 Jul 2015.

To cite this article: KA Walovich, DA Ebert, DJ Long & DA Didier (2015) Redescription of Hydrolagus africanus (Gilchrist, 1922)
(Chimaeriformes: Chimaeridae), with a review of southern African chimaeroids and a key to their identification, African Journal of
Marine Science, 37:2, 157-165, DOI: 10.2989/1814232X.2015.1033012

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 African Journal of Marine Science 2015, 37(2): 157–165 Copyright © NISC (Pty) Ltd
Printed in South Africa — All rights reserved AFRICAN JOURNAL OF
MARINE SCIENCE
ISSN 1814-232X EISSN 1814-2338
http://dx.doi.org/10.2989/1814232X.2015.1033012

Redescription of Hydrolagus africanus (Gilchrist, 1922) (Chimaeriformes:

Chimaeridae), with a review of southern African chimaeroids and a key to
their identification§
KA Walovich1*, DA Ebert1,2,3, DJ Long2,4 and DA Didier5

1
Pacific Shark Research Center, Moss Landing Marine Laboratories, Moss Landing, California, USA
2 Department of Ichthyology, Institute for Biodiversity Science and Sustainability, California Academy of Sciences, San
Francisco, California, USA
3 South African Institute for Aquatic Biodiversity, Grahamstown, South Africa

4 Department of Biology, St Mary’s College, Moraga, California, USA

5 Department of Biological Sciences, Millersville University, Millersville, Pennsylvania, USA

* Corresponding author, e-mail: kwalovich@mlml.calstate.edu

Hydrolagus africanus (Gilchrist, 1922) is a little-known chimaeroid species, originally described from the KwaZulu-
Downloaded by [Douglas Long] at 00:05 25 July 2015

Natal coast of South Africa, with a range spanning the western Indian Ocean from Kenya, south to Mozambique and
South Africa, and extending into the south-eastern Atlantic to Namibia. This species is characterised by a lateral
patch of denticles on the male prepelvic tenacula, a second dorsal fin slightly indented in the centre, a long, curved
dorsal spine equal to or sometimes exceeding the height of the first dorsal fin, and a uniform light brown body with
no distinctive markings or patterns and slightly darker brown fins. To clarify the taxonomic status of H. africanus,
we examined and compared specimens from throughout its range in southern Africa. Because the syntypes of
H. africanus are lost, we designate a neotype and redescribe the species based on the neotype and additional
southern African specimens. In addition, we present a review of southern African chimaeroids and provide an
updated key to their identification.

Keywords: African chimaera, conservation, endemic, morphology, neotype, taxonomy

Introduction

Chimaeroid fishes have been a historically understudied
group with poor initial descriptions for many species,
hence making species-specific identification difficult (Didier
1995; Didier et al. 2012). This holds true for the southern
African region despite a prolific ichthyological taxonomic
history that includes the chondrichthyan fauna (Smith and
Heemstra 1986; Gon and Skelton 1997; Ebert and van Hees
2015). Southern Africa, including Namibia, South Africa
and Mozambique, has one of the globally most diverse
chondrichthyan faunas, despite its relatively short coastline
(Compagno 1999; Ebert and van Hees 2015). Chimaeroids,
commonly referred to as ghost sharks, silver sharks or
ratfish, are unlike true shark-like fishes in possessing
elongate bodies, a long filamentous tail, a single gill opening,
smooth and scaleless skin, and paired tooth plates instead
of individual teeth (Didier 1995; Didier et al. 2012). The
family Chimaeridae has two genera, Chimaera Linnaeus,
1758 and Hydrolagus Gill, 1862, both of which occur in the
region; Chimaera is represented by the recently described
species C. notafricana Kemper, Ebert, Compagno &
Didier, 2010, but the composition of Hydrolagus species is
uncertain. The genus Hydrolagus currently is represented
by the poorly known H. africanus (Gilchrist, 1922) and
a species morphologically similar to, if not the same as,
H. trolli Didier & Séret, 2002 (Compagno 1999; Ebert and
van Hees 2015).
Hydrolagus africanus is a small-bodied chimaera
(<500 mm body length, <900 mm total length) that ranges
from northern Namibia to Kenya (Compagno et al. 1989,
1990). It is characterised by a lateral patch of denticles on
the male prepelvic tenacula, a second dorsal fin slightly
indented in the centre, a long, curved dorsal spine equal to,
or sometimes exceeding, the height of the first dorsal fin,
and a uniform light brown body with no distinctive markings
or patterns, and slightly darker brown fins. The species
was originally described from specimens collected in
bottom trawls along the KwaZulu-Natal coast, South Africa
(Gilchrist 1922). Early records from the type locality are
few and brief (Barnard 1925; Norman 1935; Smith 1949,
1965; Compagno et al. 1989). Most recent records are
from the west coast of South Africa northward to Namibia
(Lleonart and Rucabado 1984; Macpherson and Roel 1987;
Compagno et al. 1991).
The original description of H. africanus was based on
several syntypes, all of which have been lost, with no clear
indication of where type specimens, if any, were catalogued

§ This article is based on a paper presented at the ‘Sharks International 2014’ conference, held 2–6 June 2014, Durban, South Africa, and is
part of a special issue ‘Advances in Shark Research’ edited by DA Ebert, C Huveneers and SFJ Dudley

African Journal of Marine Science is co-published by NISC (Pty) Ltd and Taylor & Francis
 158
by Gilchrist or subsequent authors (Eschmeyer 2015).
Reference to the destruction of Gilchrist’s type material
following his death appears in the literature as early as 1926
(Brown 1997; Eschmeyer 2015) and the current authors
could not locate the type specimens. Several illustrations
of H. africanus have been published (Gilchrist 1922; Smith
1965; Compagno et al. 1989), but these lack sufficient detail
to enable reliable species identification, and other reference
material is sparse.
Given the uncertain status of the species and the absence
of original type material, we redescribe H. africanus based
on new information from morphometric individual variation,
ontogeny and sexual dimorphism, and designate a neotype.
We also review the chimaeroid fauna of the southern
African region and provide an updated key to the species
(Appendix).
Material and methods
A total of 66 specimens of H. africanus were examined,
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including 31 females and 35 males. Measurements were
taken on preserved specimens, measured point to point
using digital or dial calipers to the nearest 0.1 mm and
measuring tape to the nearest millimetre. All measurements
and corresponding acronyms follow Didier and Séret (2002).
Body proportions were calculated as proportions of body
length (BDL).
Body measurements included: BDL, dorsal edge of gill
opening to origin of dorsal lobe of caudal fin; total length
(TL); precaudal length (PCL), snout tip to origin of dorsal
lobe of caudal fin; snout-vent length (SVL), snout tip to vent
opening; trunk length (TRL), ventral edge of gill opening
to vent opening; pre-second dorsal length (PD2), snout tip
to origin of second dorsal fin; pre-first dorsal length (PD1),
snout tip to origin of first dorsal fin; pre-orbital length (POB),
snout tip to anterior edge of orbit; prenarial length (PRN),
snout tip to anterior end of nasal apertures; pre-oral length
(POR), snout tip to end of upper labial fold; length of second
dorsal fin base (D2B); maximum height of anterior third of
second dorsal fin (D2AH); maximum height of posterior
third of second dorsal fin (D2PH); maximum height of the
middle of the second dorsal fin (D2MH); length of first
dorsal fin base (D1B); dorsal spine height (DSA); maximum
height of first dorsal fin (D1H); dorsal caudal margin length
(CDM), from origin to insertion of dorsal caudal lobe;
maximum height of dorsal lobe of caudal fin (CDH); total
caudal length (CTL), from origin of dorsal caudal lobe to
end of caudal filament; ventral caudal margin (CVM), from
origin to insertion of ventral caudal lobe; maximum height
of ventral lobe of caudal fin (CVH); caudal peduncle height
(CPH), measured at origin of dorsal lobe of caudal fin; head
length (HDL); pectoral fin anterior margin (P1A); pelvic fin
width (PFW); pelvic fin anterior margin (P2A); interdorsal
space (IDS); dorsal-caudal space (DCS), insertion of
second dorsal fin to origin of dorsal caudal lobe; posterior
base of pectoral fin to anterior base of pelvic fin (PPS);
pelvic-caudal space (PCS), insertion of pelvic fin to origin
of ventral caudal lobe; origin of first dorsal fin to origin of
pectoral fin (D1P1); origin of first dorsal fin to origin of pelvic
fin (D1P2); origin of second dorsal fin to origin of pectoral
fin (D2P1); origin of second dorsal fin to origin of pelvic fin
Walovich, Ebert, Long and Didier
(D2P2); eye length (EYL); eye height (EYH); total clasper
length (CLT), from pelvic fin base to tip; length of medial
clasper branch from fork to tip (CLM); and length of lateral
clasper branch from fork to tip (CLL).
Sexually mature females were identified by dilated and
usually distended oviducal openings. Sexual maturity in
males was determined by the combination of hardened
pelvic claspers, the emergence of frontal and prepelvic
tenacula, and the presence of denticles on the pelvic
claspers (Didier and Séret 2002). All other specimens were
classified as juveniles.
Institutional abbreviations include South African Museum
(SAM), South African Institute for Aquatic Biodiversity
(SAIAB), and California Academy of Sciences (CAS),
following Sabaj Pérez (2013). The neotype specimen is
deposited at the SAM.
Hydrolagus africanus (Gilchrist, 1922) — African
chimaera
Chimaera africana – Gilchrist 1922: 51, Pl. 8; Barnard 1925:
95; Norman 1935: 47; Fowler 1941: 499–500; Smith 1949:
79, Fig. 94; Bigelow and Schroeder 1953: 543; Smith 1961:
573; Smith 1961: 76–77, Fig. 94; Smith 1965: 76, Fig. 94.
Hydrolagus sp. – Compagno et al. 1991: 113; Bianchi et al.
1999a: 101.
Hydrolagus africanus – Smith 1968: 3, Pl. 1a; Karrer 1975:
201; Shcherbachev et al. 1982; Lloris 1986: 148, Fig. 56; Turon
et al. 1986: 71, 137, 178, 236, 300; Compagno 1986: 145;
Macpherson and Roel 1987: 591; Roel 1987: 581; Compagno
et al. 1989: 120; Didier 1995: 15; Compagno 1999; Bianchi et
al. 1999b: 101; Novikov 2002: 279; Compagno and Dagit
2006; Human et al. 2006: 387; Kyne and Simpfendorfer
2007: 19; Kemper et al. 2010: 55; Licht et al. 2012.
Hydrolagus mirabilis? – Lleonart and Rucabado 1984: 43.
Neotype
SAM 34420, mature male, 600+ mm TL, 388 mm BDL,
South-East Atlantic (30°04′59.88″ S, 14°54′6.12″ E), 465 m,
FRS Africana.
Other material
SAM 21923, 1 M, 761 mm TL; SAM 21924, 1 F, 608 mm
TL; SAM 21925, 1 F, 533 mm TL; SAM 21926, 1 F, 897 mm
TL; no tag, 1 M, 513 mm TL; SAM 26324, 1 M, 682 mm TL;
SAM 26325, 1 F, 436 mm TL; SAM 26976, 1 F, 984 mm TL;
SAM 27581, 1 F, 610 mm TL; SAM 27750, 1 M, 28°63′ S,
32°64′ E, 1 000 m; SAM 33194, 1 F, 3 M, 544–871 mm
TL, 227–361 mm BDL, South-East Atlantic (30°14.9′ S,
14°58′ E), 464 m, FRS Africana; SAM 34412, 5 F, 16 M,
418–838 mm TL, 292–413 mm BDL, South-East Atlantic
(27°22.2′ S, 14°16.2′ E), 475 m, FRS Africana, 19 February
1988; SAM 34413, 2 F, 1 M, 770–828 mm TL, 315–337 mm
BDL, South-East Atlantic (35°20′ S, 18°45′ E), 473 m, FRS
Africana; SAM 34414, 2 M, 568–847 mm TL, 389–404 mm
BDL, South-East Atlantic (32°27.7′ S, 16°33′ E), 485 m, FRS
Africana; SAM 34415, 3 F, 446–857 mm TL, 340–461 mm
BDL, South-East Atlantic (27°27.4′ S, 14°25′ E), 425 m,
FRS Africana; SAM 34416, 1 M, 762 mm TL, 336 mm
 African Journal of Marine Science 2015, 37(2): 157–165 159

BDL, South-East Atlantic (30°05.1′ S, 14°50.3′ E), 484 m,
FRS Africana; SAM 34417, 2 F, 1 M, 633–977 mm TL,
388–465 mm BDL, South-East Atlantic (30°35.2′ S,
15°19.5′ E), 490 m, FRS Africana, 20 July 1986; SAM
34419, 4 F, 1 M, 537–844 mm TL, 377–431 mm BDL,
South-East Atlantic (29°03.6′ S, 14°26′ E), 454 m, FRS
Africana; SAM 34421, 1 M, 641 mm TL, 407 mm BDL,
South-East Atlantic (31°47′ S, 16°13′ E), 438 m, FRS
Africana, 21 January 1985; SAM 34422, 1 F, 694 mm TL,
282 mm BDL, South-East Atlantic (30°06′ S, 14°46′ E),
497 m, FRS Africana, 27 January 1985; SAM field no.
A4785048, 1 F, 940 mm TL; A4373046, 1 M, 649 mm TL;
SAM 34498, 6 F, 229–236 mm TL, South-East Atlantic
(30°28′ S, 15°12′ E), 400 m, FRS Africana, 5 February 1996;
SAM 34498, 1 F, 850 mm TL, 361 mm BDL, South-East
Atlantic (30°28′ S, 15°12′ E), 400 m, FRS Africana,
5 February 1996; CAS 229761, 1 M, 230 mm TL; SAIAB
67402, 1 M, 1 F, 527–651 mm TL, South-East Atlantic
(06°58′ S, 11°40′ E), 724 m, 22 March 2002.
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South-East Atlantic (06°38′ S, 11°25′ E), 724 m, 23 March
2002.
Results
Diagnosis
A small species of Hydrolagus distinguished from its
congeners by a lateral patch of 1–3 denticles on the male
prepelvic tenacula, a second dorsal fin slightly indented in
the centre and a long, curved spine, equal to or sometimes
exceeding the height of the first dorsal. Body colour is light
brown, head often darker than the trunk, pale grayish brown
ventral surface and dark brown fins that lighten near the
body margin.
Description
Morphometric measurements, expressed as ratios of
BDL, of all specimens are presented in Table 1. A small-
bodied species with slender head (17.9–31.1% BDL), blunt,
rounded snout that sharply tapers posteriorly to a long,

Hydrolagus mirabilis – SAIAB 67976, 1 F, 625 mm TL, thin caudal filament. Large eyes (length 22.4–41.9% HDL,

Table 1: Morphometric measurements of Hydrolagus africanus expressed as percentage of body length (BDL)

Percentage of BDL
Measurement
Neotype male Range all (n = 65) Range females (n = 31) Range males (n = 34)
BDL 388 (mm) 221–465 (mm) 221–465 (mm) 292–415 (mm)
TL 154.6 117.2–293.8 119.9–293.8 117.2–251.4
PCL 121.4 116.1–130.0 116.1–130.0 116.5–125.3
SVL 53.9 51.9–77.9 54.0–74.7 51.9–77.9
TRL 35.6 30.9–46.8 34.0–46.8 30.9–45.1
PD2 46.4 41.1–57.7 41.1–57.7 41.3–51.7
PD1 25.5 20.3–339 20.3–33.9 20.6–29.8
POB 11.3 8.5–15.0 8.8–15.0 8.5–13.7
D2B 79.6 69.6–86.7 70.5–78.3 69.6–86.7
D2AH 4.9 4.3–7.5 4.5–7.5 4.3–6.9
D2PH 4.1 3.4–6.5 3.4–5.9 3.5–6.5
D2MH – 2.8–5.0 2.8–4.4 2.8–5.0
D1B 9.0 9.0–18.9 13.0–18.9 9.0–17.5
DSA 25.8 18.6–28.3 18.6–25.8 21.2–28.3
D1H 19.3 11.8–20.5 11.8–19.7 13.2–20.5
CDM 21.1 16.0–25.6 16.1–23.7 16.0–25.6
CDH 2.6 2.3–5.0 2.4–5.0 2.3–4.4
CTL 34.3 33.8–163.0 34.4–163.0 33.8–127.7
CVM 33.8 22.8–44.1 25.8–43.0 22.8–44.1
CVH 2.8 2.0–5.0 2.4–5.0 2.0–4.1
CPH 2.6 0.0–3.0 0.0–3.0 0.0–2.6
HDL 21.6 17.9–31.3 17.9–31.3 18.2–26.5
P1A 37.1 29.3–41.4 29.3–41.4 31.52–39.7
P2A 19.3 16.3–23.5 16.3–23.5 16.3–22.1
IDS 12.6 2.1–12.6 3.9–11.9 2.1–12.6
DCS 0.0 0.0–1.3 0–1.0 0.0–1.3
PPS 28.4 25.5–37.6 27.8–37.6 25.5–35.7
D1P1 14.7 14.4–21.9 15.7–21.9 14.4–20.4
D1P2 38.9 25.6–44.0 37.6–44.0 25.6–42.7
D2P1 27.6 23.5–34.1 25.3–32.8 23.5–34.1
D2P2 23.5 18.8–27.7 20.0–27.7 18.8–25.7
EYL 8.2 5.1–9.7 5.8–9.7 5.1–8.3
EYH 4.9 2.9–5.9 2.9–5.7 3.8–5.9
CLT 12.1 3.8–14.2 – 3.8–14.2
CLM 4.9 4.5–7.7 – 4.5–7.7
CLL 4.9 4.3–7.2 – 4.3–7.2
 160
Figure 1: Hydrolagus africanus (Gilchrist, 1922) neotype, SAM 34420, mature male, 600+ mm TL, 388 mm BDL, collected from the west
coast of South Africa. Scale bar = 5 cm
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height 14.5–27.3% HDL) and deciduous skin, which flakes
off in large patches 1–3 cm in diameter (Figure 1).
First dorsal fin triangular with a short base (10.3–18.9%
BDL) and attached to the basal third of the spine. First
dorsal fin spine equal to or slightly longer than height of first
dorsal fin. Spine curved with two rows of serrations along
the distal third of posterior edge and keeled along anterior
edge. Spine length (18.6–28.3% BDL) nearly equal to or
slightly longer than head length (17.9–31.1% BDL). Spine
tip when depressed reaches to origin or slightly past the
origin of second dorsal fin. Interdorsal space usually small
(2.1–11.9% BDL) and connected by a low membrane.
Anterior portion of second dorsal fin (4.3–7.5% BDL) is
slightly higher than the posterior portion (3.4–6.5% BDL).
Second dorsal fin long and slightly depressed in the
centre (2.8–5.0% BDL); distance from second dorsal fin
insertion to caudal lobe origin (0–1.3% BDL) with little to
no separ­ation; posterior portion of second dorsal rounded,
extending to or beyond the insertion of caudal lobe. Caudal
dorsal margin (16.0–25.6% BDL) shorter than ventral
margin (22.8–44.1% BDL); caudal dorsal height (2.3–5.0%
BDL) nearly equal to ventral height (2.0–5.0% BDL) and
extending as a fleshy ridge along the posterior body. Anal
fin absent.
Pectoral fins large, triangular with anterior margin
(29.3–41.4% BDL) extending to or just beyond insertion
of pelvic fins when laid against the body; anterior margin
approximately 1.2–2.2 times larger than pelvic anterior
margin (16.3–23.5% BDL). Pelvic fin pointed distally, broadly
rounded along posterior edge where it joins fin base.
A volmerine and a palatine tooth plate are present on
each side of the upper jaw, with two mandibular plates on
the lower; each volmerine tooth plate with 5–6 tritor ridges.
Palatine tooth plates flat, triangular in shape, and lying
posterior to volmerine plates on upper jaw. Tooth plate colour
varies from yellow to light brown in preserved specimens.
Lateral lines with open grooves and dilations around
the snout. Preopercular and oral canals share a common
branch from infraorbital canal in most specimens; however,
in some individuals the preopercular, oral and infraorbital
canals share a common origin (Figure 2). Trunk lateral line
Walovich, Ebert, Long and Didier
curves downward in sigmoid shape near its origin, before
extending in a straight line to its termination.
Male frontal tenaculum slender with a bulbous tip that
curves slightly upward with several overlapping rows of
fine denticles on ventral surface. Prepelvic tenacula with
five strong denticles along medial edge and 1–3 lateral
denticles next to the medial row (Figure 3). Pelvic claspers
small, trifurcate, pale in colour, and do not extend beyond
distal edge of pelvic fins. Claspers divided along distal
one-third of length, and distal tips with small fleshy bulbs
bearing extremely fine shagreen of denticles (Figure 4).
Females with small anal pads on base of tail posterior to
cloaca.
Body colouration a light brown dorsally, with no distinctive
markings or patterns; head often darker than trunk; ventral
surface pale grey to brown. Fins dark brown to blackish that
lighten in colour near the body margin.
Distribution
Hydrolagus africanus occurs in the western Indian Ocean
from Kenya and Mozambique to the Western Cape province,
South Africa, and in the south-eastern Atlantic along the
west coast of South Africa north to Angola. The occurrence
of this species in Angolan waters is confirmed here for the
first time. Records of this species from south-western Indian
Ocean ridges and seamounts (Novikov 2002; Parin et al.
2008) require confirmation as recent surveys have found
other Hydrolagus species to occur there but no confirmed
specimens of H. africanus have been observed (PJ Clerkin,
Moss Landing Marine Laboratories, pers. comm.). The
species has been recorded from depths of 300–1 030 m, but
occurs most commonly at depths of 300–500 m, on soft or
muddy substrates on the continental slope (Compagno et
al. 1991). The species has been reported from a depth of
1 570 m but the identification of those specimens remains
uncertain (Shcherbachev et al. 1982; Novikov 2002; Parin et
al. 2008).
Biological notes
Very little is known about the biology of H. africanus,
although Macpherson and Roel (1987) speculated that
 African Journal of Marine Science 2015, 37(2): 157–165
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Figure 2: Intraspecific comparison of lateral line canals of (a) Hydrolagus africanus neotype and (b) adult male (SAM 27750); preopercular
(POP), oral (O) and intraorbital (IO). Scale bar = 3 cm
Figure 3: Sexual characters in adult male neotype SAM 34420, (a) lateral view of frontal tenaculum and (b) prepelvic tenaculum with five
medial denticles and one lateral denticle (arrow). Scale bar = 3 cm
it likely feeds on infaunal invertebrates and small fishes.
Reproductive mode is oviparous, but reproductive informa-
tion remains limited. Males and females appear to mature
at body lengths >350 mm. The original species description
details egg cases recovered in the same trawl as adults,
with the eggs described as being 54% of total length and
having poorly developed lateral web-like expansions, a
pointed anterior and narrow posterior, a median keel along
the egg length, and numerous respiratory openings at the
posterior (Gilchrist 1922). However, egg cases were not
taken directly from the female oviduct and hence it cannot
be confirmed that they belonged to H. africanus. Both
Smith (1961) and Compagno (1986) provide illustrations of
egg capsules of H. africanus but neither detail the origin of
the figures, and a search of the literature failed to reveal
additional information on egg capsules for this species.
161
Discussion
The genus Hydrolagus has a rather complex taxonomic
history in southern Africa, with H. africanus having long
been considered to be the only confirmed representa-
tive of the genus. The occurrence of additional southern
Africa species, either named or possibly undescribed
species, within this genus has been reported upon, but not
confirmed (Compagno 1986; Compagno et al. 1989, 1991).
Compagno (1986) reported on a large Hydrolagus species
taken off Durban, South Africa, that appeared to be close
to H. alberti Bigelow & Schroeder, 1951, but not conspe-
cific with it, whereas Compagno et al. (1989) suggested
that H. africanus from off Durban and Mozambique
needed to be compared closely with similar Hydrolagus
species from the west coast of South Africa and Namibia.
 162
Figure 4: Ventral view of neotype SAM 34420 claspers. Scale
bar = 3 cm
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Compagno et al. (1991) referred to the West Coast brown
chimaera as Hydrolagus species, commenting that it could
be H. africanus, the European H. mirabilis (Collett, 1904),
or possibly a different Hydrolagus species. However, this
subject requires further investigation. In a checklist of
southern African chondrichthyans, Compagno (1999) lists
three Hydrolagus species, including two that may possibly
be new species. A more recent checklist includes two
species, H. africanus and H. cf. trolli, from southern Africa
(Ebert and van Hees 2015).
The redescription of H. africanus presented here is crucial
in clarifying its status with the aim of improving the identifi-
cation of this species relative to other Hydrolagus species
that may occur in the southern African region. Our findings
agree in general with the original description of H. africanus
by Gilchrist (1922), but with the additional material now
available we have been able to clarify further the morpho-
logical distinctiveness of this species, including second
dorsal fin shape and clasper morphology.
Examination of additional specimens revealed a lateral
line branching pattern that was not previously reported for
this species. Lateral line morphology can be important in
separating some Hydrolagus species. The preopercular
(POP) and oral (O) canals in the majority of H. africanus
specimens examined share a common branch from the
infraorbital (IO) canal, but the POP, O and IO canals of
several specimens share a common origin (Figure 2).
Intraspecific variation in lateral line morphology has been
observed in other chimaeroid species (Didier and Séret
2002; James et al. 2009; Kemper et al. 2010; Angulo et al.
2014) and hence these branching patterns are most useful
when considered in combination with other morphological
characters (Didier et al. 2012).
Hydrolagus africanus is the only known species to
possess a lateral patch of denticles on the prepelvic
tenacula in addition to five denticles present along the
medial edge. All chimaeroid species exhibit a row of 5–7
denticles on the paired prepelvic tenacula, which articulate
anteriorly with the pelvic girdle to anchor the male to the
female during copulation (Didier et al. 2012). This patch of
denticles was not included in the original description, but
Walovich, Ebert, Long and Didier
can serve as a key diagnostic characteristic in separating it
from other Hydrolagus species. Gilchrist (1922) detailed six
short, recurved spines on the outer margin of the prepelvic
tenacula; however, the lateral patch observed likely accounts
for this discrepancy.
Our current investigation has led to additional observa-
tions on the status of other possible Hydrolagus species from
southern Africa. The identification of additional brown, small-
bodied Hydrolagus species (300–350 mm BDL at maturity)
from this region is difficult due to overlapping morphological
characters, unknown ontogenetic changes and artifacts of
preservation, leading to misidentifications and speculation
about multiple co-occurring species. Hydrolagus mirabilis
is known from throughout the Atlantic and was listed in
a checklist of species from the southern African region by
Krefft (1990), but until now has not been confirmed with
specimens. A small-bodied, brown Hydrolagus species
collected off Angola and found in the fish collection at
SAIAB confirms the presence of both H. africanus and
H. mirabilis from Angola. In fact, we were able to confirm the
occurrence and overlap in distribution in Angolan waters of
both species (SAIAB 67402 = H. africanus, SAIAB 67976 =
H. mirabilis). The distributional overlap of these two species
may include Namibia where H. mirabilis and a Hydrolagus
sp. have been reported, but without specimens, illustrations
or photographs of H. mirabilis from that area its occurrence
remains unconfirmed (Krefft 1990; Bianchi et al. 1999a,
1999b). Morphologically, H. mirabilis and H. africanus
are very similar in colouration, size, and snout shape, but
H. mirabilis can be distinguished by a pronounced concave
second dorsal fin margin, nearly separating the fin into
anterior and posterior portions, and a larger eye diameter of
35% HDL, compared to 31% HDL in H. africanus. Variations
in snout morphology and colouration have been observed
in H. africanus specimens from the east and west coasts of
South Africa (Compagno 1986; Compagno et al. 1991), but
natural variation and artifacts of preservation yield overlap-
ping morphometrics and do not present sufficiently strong
evidence to separate H. africanus into two species at this
time. Preliminary mitochondrial genetic data suggest that
H. africanus, H. mirabilis and other morphologically similar
species (e.g. H. alberti, H. mitsukurii) are closely related
and may, upon further investigation, prove to be a species
complex (JM Kemper, College of Charleston, USA, pers.
comm.).
Re-examination of literature (Compagno et al. 1991) and
cruise data (DAE unpublished data) indicates the presence
of one or more large-bodied species of Hydrolagus in
southern Africa. A large Hydrolagus species first reported by
Compagno (1986, 1999) and referred to as an undescribed
‘pointynose blue chimaera’ and ‘big black chimaera’ has
since been collected in offshore trawls. This species has
tentatively been designated H. cf. trolli based on its large
size (550–650 mm BDL at maturity), second dorsal fin of
uniform height, blue to purple colouration, and long, pointed
snout (Didier and Séret 2002). Size can easily distinguish
adults of the two species; however, juveniles of H. cf.
trolli may be confused with H. africanus based on similar
colouration, resulting in the misidentification of SAM and
SAIAB material and further taxonomic uncertainty. Further
examination of museum collections, the acquisition of fresh
 African Journal of Marine Science 2015, 37(2): 157–165
specimens and molecular analysis are needed to fully
elucidate the identification of these other Hydrolagus species
from southern African waters.
Review of southern African chimaeroid species
Eight species of chimaera are found in the southern African
region, representing the three families and all six genera
(Ebert and van Hees 2015). Callorhinchus capensis
Duméril, 1865, the St Joseph’s elephantfish, was likely the
first southern African endemic to be given a valid binomial
name (Compagno 1999; Eschmeyer 2015). This species is
characterised by a plough- or hoe-shaped snout, distinct
second dorsal fin, heterocercal caudal fin and silvery colour
with dusky brown markings. It occurs on the continental
shelf to 600 m, but is most commonly found shallower than
250 m (Compagno et al. 1991). This species is endemic to
southern Africa, ranging from Namibia to the KwaZulu-Natal
coast of South Africa (Compagno 1986; Compagno et al.
1989, 1991). In terms of human consumption, Callorhinchus
capensis was long considered a low-quality fish with limited
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value, until a commercial fishery, centred in St Helena Bay
on the west coast of South Africa, developed in the early
1980s and in which landings have remained at approximately
800 tonnes per year (Freer and Griffiths 1993; DAFF 2013).
Compagno et al. (1990) reviewed the family Rhinochimaeri-
dae, commonly referred to as the longnose chimaeras, and
described a new species from the southern African region.
The family can be subdivided into two distinct lineages, the
subfamilies Harriottinae and Rhinochimaerinae.
The Harriottinae comprise two genera, Harriotta Goode &
Bean, 1895 and Neoharriotta Bigelow & Schroeder, 1950,
which can be separated by the presence (in Neoharriotta)
or absence (in Harriotta) of a distinct anal fin (Didier et al.
2012). Harriotta raleighana Goode & Bean, 1895 is one
of the widest ranging chimaeroid species and has been
reported from Namibia to the Western Cape province, South
Africa (Ebert in press). This species has a uniform brown
colour with a long, flattened snout and short, broad pectoral
fins. Neoharriotta pinnata (Schnakenbeck, 1931) is a large-
bodied species (500–600 mm BDL at maturity) described
from Walvis Bay, Namibia, and ranging north to the western
Sahara (Ebert in press). However, recent records from
the Gulf of Aden and off south-eastern India in the Bay of
Bengal indicate this species may be more wide-ranging
than previously thought (Ali et al. 2009; Suresh and Raffi
2012; Ebert 2014). The monogeneric Rhinochimaerinae is
represented by two species: Rhinochimaera atlantica Holt
& Byrne, 1909 and Rhinochimaera africana Compagno,
Stehmann & Ebert, 1990. Members of this lineage have
smooth, blade-like toothplates rather than raised tritors,
tubercles on the dorsal caudal fin and a dorsoventrally
compressed neurocranium (Didier 1995). Rhinochimaera
africana was originally described from southern Africa but is
now known to occur globally, with records from the western,
central and south-eastern Pacific (Compagno et al. 1990;
Didier and Nakaya 1999; Last and Stevens 2009; Angulo et
al. 2014). The species is distinguished from R. atlantica by
its dark colouration, longer, broader, paddle-shaped snout,
and shorter caudal fin (Compagno et al. 1990).
The occurrence of a chimaera species from southern
Africa has been known since the mid 19th century when
163
Duméril (1865) identified a specimen as Chimaera
monstrosa Linnaeus, 1758 from the Cape of Good Hope.
Most subsequent authors agreed with Duméril’s identifica-
tion (see Kemper et al. 2010), until Compagno et al. (1989)
re-evaluated these records and suggested that this may
be a new species, which was later described as Chimaera
notafricana Kemper, Ebert, Compagno & Didier, 2010. This
species can be identified by a combination of the following
characters: pectoral fin, when depressed, reaches the origin
of the pelvic fin base; caudal fin ventral margin terminates
slightly posterior to the caudal fin dorsal margin insertion;
and there is a uniform blackish-brown colouration with dark
blue streaking. Chimaera notafricana has a restricted coastal
distribution from Lüderitz, Namibia, to south of Algoa Bay,
Eastern Cape province, South Africa (Kemper et al. 2010).
The vulnerability of southern African chimaeroids to
expanding deep-water demersal fishing pressure, the
lack of available biological and ecological data and the
taxonomic uncertainty of the genus Hydrolagus are of
concern (DAFF 2013). In 2006, the International Union for
the Conservation of Nature (IUCN) classified N. pinnata,
R. africana, H. africanus and H. trolli as ‘Data Deficient’ on
the Red List of Threatened Species (IUCN 2013). Since
then, little information has been added to the already sparse
knowledge of this order. Improved systematic resolution will
aid in the identification, conservation and management of
this unique and understudied group of cartilaginous fishes
in the southern African region.
Acknowledgements — The following individuals are sincerely
thanked for their assistance and support: Paul Cowley and Roger
Bills (South African Institute for Aquatic Biodiversity), Leonard
Compagno and Michael Bougaart (South African Museum), Gavin
Naylor and Jenny Kemper (College of Charleston), and Rob Leslie
(South African Department of Agriculture, Forestry and Fisheries).
A travel grant provided by the California State University Council on
Ocean Affairs, Science and Technology (COAST) allowed KAW to
present these results at the Sharks International 2014 Conference
in Durban, South Africa, 2–6 June 2014. The Save Our Seas
Foundation, and the South African Institute for Aquatic Biodiversity,
provided funding support to DAE. Additional support was provided
by a National Science Foundation grant (Jaws and Backbone:
Chondrichthyan Phylogeny and a Spine for the Vertebrate Tree of
Life, DEB 1132229).
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Appendix: Key to southern African species

1a. Snout plough-shaped; trunk a pale silver with several dark spots; heterocercal tail; large anal fin preceding caudal fin; male pelvic
claspers unbranched and tube-like without fleshy denticulate tip ……………….……………………………….Callorhinchus capensis
1b. Snout not plough-shaped…...…………………………………………………………................................................………………….…..2
2a. Elongate, spear-shaped snout; male pelvic claspers unbranched, slender rods with denticulate bulbous tip; body even brown with
no distinct markings.……………….....................................................................................................................................................…3
2b. Blunt fleshy snout; male pelvic claspers are branched with fleshy denticulate lobes at the tips .....................................................…..6
3a. Toothplates with smooth shearing blades; tubercles present on dorsal caudal fin; dorsal surface of head not arched about snout
profile ……..……………………………………………..………..…………........................................................................................…….4
3b. Toothplates with raised hypermineralised tritors on surface; tubercles not present on dorsal caudal fin; dorsal surface of head
arched about snout profile .…….…………….………….……...…..………....………............................................................................5
4a. Snout broad and paddle-shaped; even dark brown body colour; caudal tubercle count 40–46……….....…..Rhinochimaera africana
4b. Snout narrow and conical; body colour pale, whitish to grey-brown; caudal tubercle count 19–33…..........Rhinochimaera atlantica
5a. Separate anal fin preceding the ventral lobe of caudal fin ………………………..…............................................Neoharriotta pinnata
5b. No anal fin present……………..………………………..……..……………..…..….................................................Harriotta raleighana
6a. Anal fin present, separated from caudal fin by small notch………………….....…..............................................Chimaera notafricana
6b. Anal fin absent, ventral caudal fin is continuous along entire length…....……………………..............................................……………7
7a. Large-bodied with distinctive pointed snout; dark blue to black in colour; second dorsal fin uniform along entire length;
bifid claspers…….……….………....……………………......................................................................................…Hydrolagus cf. trolli
7b. Medium-bodied; brown to tan in colour; second dorsal fin slightly indented at centre; trifid pelvic claspers; males with lateral patch
of denticles on the prepelvic tenaculum .…………..….……..............................................................................Hydrolagus africanus

Manuscript received June 2014, revised November 2014, accepted January 2015

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