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Description of a unique catshark egg capsule (Chondrichthyes: Scyliorhinidae) from the North West Shelf, Western Australia.

INTRODUCTION

Recent taxonomic revisions, species descriptions and nomenclatural changes to the Australian chondrichthyofauna (eg. Huveneers 2006; Last et al. 2006, 2007, 2008a-c; Yearsley & Last 2006; Last & Gledhill 2007; Jacobsen & Bennett 2007, 2009; Last & Chidlow 2008; Last & Stevens 2009) prompted a review of the chondrichthyan reference collection at the Western Australian Museum (WAM) to update or correct the nomenclature and species assignments of its specimens. During the course of that review, a number of interesting taxonomic finds arose. One of these was the discovery of several lots of a unique egg capsule belonging to a species of shark from the family Scyliorhinidae (catsharks).

The egg capsules discovered in the WAM collection were identifiable as scyliorhinid egg capsules due to their size, overall shape, and presence of an embryo associated with one of the egg capsules. However, it was immediately apparent that they represented an undescribed egg capsule, being unlike any other egg capsule yet described for a chondrichthyan fish. The character unique to these novel egg capsules was the presence of well developed ridges, T-shaped in cross section, that run longitudinally along the length of the egg capsules, with the egg capsules bearing very short horns and tendrils.

The unique characteristics of the egg capsules made it initially impossible to place them within a genus. However, an examination of the only embryo associated with the egg capsules provisionally identifies the species as a member of the genus Apristurus Garman, 1913. Although species of Apristurus are known to occur in the region, the egg capsules were recovered from localities from which specimens of Apristurus are not yet known (Last & Stevens 2009). Further, the gross morphology of the embryo does not agree with the Apristurus candidate species that are known to occur closest to the localities of the egg capsules and may therefore represent an undescribed species. This contribution describes these novel egg capsules and the associated embryo, and compares them to the other catshark genera, and members of Apristurus, known to occur in Australian waters.

MATERIALS AND METHODS

Morphometrics and meristics: Measurements of the dimensions of the egg capsule followed Gomes & de Carvalho (1995) with terminology adapted by Ebert et al. (2006) and Flammang et al. (2007), with the addition of egg capsule depth (ECD), the greatest transverse depth of the egg capsule.

Morphometric measures of the embryo follow Compagno (1984, 2001) with scyliorhinid adaptations from Human (2006). Total length of the embryo was taken via direct measurement, and a limited number of morphometrics were estimated to the nearest 0.5 mm from stereomicroscope photographs of the embryo and calibrated scale bar. With the proposed placement of the embryo within the genus Apristurus, morphometrics were again estimated from the stereomicroscope photographs using the methodology of Nakaya et al. (2008a).
Table I. Weight, morphometric measures (mm), and measurements
as percentage of ECL (in brackets) of the egg capsules.
Abbreviations - Wt, egg capsule empty dry weight (g); ECL,
egg capsule length; ABW, anterior border width; ACW, anterior
case width; WCW, minimum waist width; PCW, posterior case width;
PBW, posterior border width; and ECD, maximum depth.

     P.28064-011  P.28097-005  P.30047-001

Wt          1.23         1.56         1.51

ECL           48           50           57

ABW    19 (39.6)    12 (24.0)  13.5 (23.7)

ACW    19 (39.6)  18.5 (37.0)  19.5 (34.2)

WCW    18 (37.5)  18.5 (37.0)    18 (31.6)

PCW    22 (45.8)  21.5 (43.0)    21 (36.8)

PBW     6 (12.5)    4.5 (9.0)    2.5 (4.4)

ECD    15 (31.3)    16 (32.0)    16 (28.1)



Photographs of the egg capsules were taken using a SLR digital camera. The embryo was mounted on a pin and photographs were taken using a Leica MZ16A stereomicroscope with a 0.63x lens, and Leica DFC 500 camera. The scale bar was calibrated using vernier callipers and Leica Application Suite ver.3.6.0 immediately prior to the photographs being taken.

Study material: All lots are deposited in the Western Australian Museum (WAM) ichthyological reference collection: WAM P.28064-011, single empty egg capsule, collected by N. N. Sinclair and P.F. Berry aboard R.V. Courageous scampi survey, 17 August 1983 from 450-452 m at 18[degrees]01'S 118[degrees]13'E, approx. 100 km SW of Rowley Shoals, Western Australia; WAM P.28097-005, single empty egg capsule, collected by N. N. Sinclair and P. F. Berry aboard R.V. Courageous scampi survey, 22 August 1983 from 410-414 m at 18[degrees]15'S 118[degrees]02'E, approx. 180 km SW of Rowley Shoals, Western Australia; WAM P.30047-001, single egg capsule with associated embryo, collected during a CSIRO scampi survey, February/March 1989, North West Shelf, Western Australia, with no further details.

RESULTS

Egg capsule description: The weights and morphometric dimensions of the egg capsules are given in Table I, and photographs of each of the egg capsules are provided in Fig. 1. The egg capsules have an overall shape typical of scyliorhinid sharks (Ivanov 1987; Gomes & de Carvalho 1995; Ebert et al. 2006; Flammang et al. 2007), with a quadrangular outline rounded at the posterior end, posterior and anterior portions of the capsule separated by a waist, anterior horns, and posterior tendrils.

[FIGURE 1 OMITTED]

Although present in these egg capsules, the waist is only marginally narrower than the anterior and posterior case widths and is poorly defined. The egg capsules possess very short horns on either side of a relatively broad anterior margin. Only WAM P.28097-005 possessed what appeared to be a full complement of the tendrils, which had otherwise snapped off partially or totally on the other egg capsules. A single tendril extends from each side of the egg capsule. The tendrils were relatively short compared to other scyliorhinid egg capsules and very thick at their origins, but tapered quickly and tightly coiled. The lateral margins of the egg capsule turn abruptly medially at the posterior of the egg capsule, allowing for a very short posterior margin. The tendrils curve around tightly at the posterior, leaving little space between the tendrils and the posterior margin.

The ridges on the egg capsules are well developed, more so than in any other chondrichthyan egg capsule so far described. The ridges run longitudinally along the length of the egg capsule. The ridges are T-shaped in cross section (Fig. 2) and obscured the location of both the anterior and posterior fissures, which could not be located on any of the egg capsules. Each egg capsule had a lateral ridge on each of their sides, however, the number of ridges on the egg capsule surfaces varied between egg capsules, and each surface. WAM P.28064-011 possessed five ridges on one surface and six on the other, WAM P.28097-005 possessed six and seven ridges on each of its surfaces, respectively, and WAM P.30047-001 also possessed six and seven ridges, respectively.

[FIGURE 2 OMITTED]

Embryo description: The embryo (WAM P.30047-001) was approximately 27 mm TL and 0.2 g, attached by a yolk stalk to a yolk sac measuring 14 mm x 13 mm x 9 mm and weighing 1.2 g (Fig. 3). Estimated morphometrics of the embryo are provided in Table II. There were few differences between the two morphometric methodologies, although several additional morphometrics were included following the Nakaya et al. (2008a) methodology.

[FIGURE 3 OMITTED]

[FIGURE 4 OMITTED]

[FIGURE 5 OMITTED]

[FIGURE 6 OMITTED]
Table II. Morphometric measures, and proportions as
percentage of total length, of the embryo (WAM P.30047-001)
estimated from photographs (Figs 4-6) taken using a
stereomicroscope and calibrated scale bar. Two sets of
measures are provided using the methodologies of Compagno (1984, 2001)
and Human (2006), compared to Nakaya etal. (2008a), respectively.
[dagger] Morphometric character number of Nakaya et al.
for equivalent (or near equivalent) measurement. * Total
Length was taken via direct measurement of the embryo. ?
indicates that the corresponding morphometric measurement
could not be estimated from the photographs using the methodology
of Nakaya et al. (2008a). t Morphometric character of Nakaya
et al. (2008a).

Morphometric         Per Compagno          Per
                       & Human           Nakaya
                                         et al.

                          mm         %     [double   mm     %
                                         dagger]

Total length *                 27              1    27

Snout to 1st dorsal            15  55.6        5    15  55.6
origin

Interdorsal space               1   3.7       34     1   3.7

2nd dorsal-caudal             1.5   5.6        -
space

Snout to anal                15.5  57.4        9     ?     ?
origin

Snout to pelvic              12.5  46.3        7  12.5  46.3
origin

Snout to pectoral               8  29.6        6   8.5  31.5
origin

Snout to vent                13.5  50.0        8  13.5  50.0
length

Pectoral to pelvic            3.5  13.0       37     3  11.1
space

Pelvic to anal                0.5   1.9        -
space

Anal to caudal                2.5   9.3        -
space

Head width at                   5  18.5        -
posterior margin of
orbit

Interorbital space            4.5  16.7       30   4.5  16.7

Snout to 1st gill             8.5  31.5       11   8.5  31.5
slit

Snout to spiracle               7  25.9       12     7  25.9

Snout to orbit                  5  18.5       13     5  18.5

Eye length                    1.5   5.6       26   1.5   5.6

Snout to mouth                4.5  16.7       16   4.5  16.7

Snout to nares                  3  11.1        -

Mouth length                    1   3.7       22     1   3.7

Mouth width                   1.5   5.6       21   1.5   5.6

Nostril width                 1.5   5.6        -

Inner internares                2   7.4       23     2   7.4
space

Outer internares                4  14.8        -
space
Pectoral length               3.5  13.0        -

Pectoral anterior             2.5   9.3       52     3  11.1
margin

Pectoral posterior              3  11.1       53     ?     ?
margin

1st dorsal length               3  11.1       43   3.5  13.0

1st dorsal base                 3  11.1       44     3  11.1

1st dorsal height               1   3.7       45     ?     ?

Pelvic length                   3  11.1       57     3  11.1

Pelvic height                 1.5   5.6        -

2nd dorsal length               4  14.8       47     4  14.8

2nd dorsal base               2.5   9.3       48   2.5   9.3

2nd dorsal height               1   3.7       49   1.5   5.6

Anal length                     4  14.8       62     ?     ?

Anal height                     1   3.7       65     ?     ?

t PreDl-insertion                              4    17  63.0
length

t Pre-outer nostril                           14     3  11.1
length

t Pre-inner nostril                           15   3.5  13.0
length

t Nostril length                              28   1.5   5.6

t Nostril-mouth                               29     1   3.7
space

t D1-D2 insertions                            36     4  14.8

t P1-P2 origins                               39     4  14.8

t P1 width                                    55     3  11.1



The embryo (Figs. 4-6) was characterised by possessing a greatly depressed head and flat snout that is pointed and wedge-shaped in lateral view, with simple nasal flaps and no nasoral grooves, an arched mouth with well developed upper and labial furrows, body not compressed, spindle or tadpole shaped, a large first dorsal fin proceeded closely by a larger second dorsal fin, moderately sized, paddle shaped pectoral fins, very large pelvic fins and anal fin and a moderately-sized caudal fin with no crest of denticles. The mouth of the embryo was wide open.

DISCUSSION

Whereas the taxonomy of Australian sharks and rays are relatively well known, due in large part to the taxonomic works mentioned above, descriptions and species identification of egg capsules for oviparous chondrichthyans are less well known and are lacking in the literature, despite their potential use as taxonomic characters (see Gomes & de Carvalho 1995; Flammang et al. 2007). While recent species descriptions have included examples of egg capsules where known, egg capsules remain unknown for many Australian oviparous chondrichthyans. Likewise, descriptions of scyliorhinid egg capsules are limited (Springer 1979; Gomes & de Carvalho 1995; Ebert et al. 2006; Flammang et al. 2007).

The embryo is apparently in early to mid development judging by the presence of external gill filaments protruding through the gill slits and spiracles. Given the lack of information available regarding hatching times for Apristurus (and other scyliorhinids in general), the embryo's age can not be determined, but development of Apristurus embryos take 24-27 months in 5[degrees] C water based on observations of conspecifics at the Monterey Bay Aquarium (Flammang 2005; Flammang et al. 2007). Jones & Geen (1977) removed complete egg capsules from freshly caught Apristurus brunneus and maintained them in 10[degrees]C water. After a period of 71/2 months, the embryos were approx. 30 mm (about the size of the current embryo), after 14 months were 55-60 mm long, and are free swimming at approx. 70-80 mm.

Although the author concedes that morphological features of an embryo at this stage of development are far from diagnostic for any particular species, the gross morphology of the embryo was developed well enough that there is little doubt that the shark belongs to the family Scyliorhinidae. Through a process of elimination of all other possible scyliorhinid genera known to occur in Australia, the combination of the above characters observable in the gross morphology suggest that the embryo belongs to the genus Apristurus Garman, 1913.

The definition of the genus Apristurus is a catshark of the Scyliorhinid subfamily Pentanchinae Smith & Radcliffe and tribe Pentanchini Smith & Radcliffe, with a combination of characters including a greatly depressed head and flat snout that is pointed and wedge-shaped in lateral view; nasal flaps simple and not elongate; nasoral grooves absent; mouth arched with well developed upper and lower labial furrows; body soft and relatively stocky, not compressed, spindle or tadpole shaped; two large dorsal fins, with first subequal or smaller than second; pelvic fins that are larger than the second dorsal fin; a large anal fin that is larger than the pelvic and dorsal fins, and is separated from the lower caudal lobe by a notch only; and lacking a distinct crest of denticles on the upper or lower margins of the caudal fin (Nakaya 1975; Springer 1979; Compagno 1988).

The overall gross morphology and proportions of the embryo agree with Apristurus. Of the other scyliorhinid genera present in Australia, the embryo cannot be placed within the genus Atelomycterus because that genus possesses expanded nasal flaps that are fused into a nasal curtain, whereas the nasal flaps are simple on the embryo in question. The embryo is excluded from the genera Figaro, Galeus and Parmaturus because these possess distinct crests of denticles on the upper caudal fin margin (and lower caudal fin margin in Figaro and Parmaturus), whereas no denticle crests are evident on the present embryo. Cephaloscyllium possesses a first dorsal fin that is much larger than the second dorsal fin, and no labial furrows; the opposite is true for the embryo in question. The anal fin is larger than the second dorsal fin in the embryo examined here, which is the opposite of that found in the genera Aulohalaelurus, Bythaelurus and Halaelurus, where the anal fin is subequal to the first dorsal fin. The genus Asymbolus has poorly developed labial furrows, expanded nasal flaps, and dorsal fins that are well separated from each other and the caudal fin, whereas the dorsal fins are close together and narrowly separated from the caudal fin in the embryo.

Egg capsules described for members of the genus Asymbolus have long tendrils, much longer than those observed on the present egg capsules, have fine silky longitudinal striations with long silky filaments emanating from the sides, which are lacking in the current egg capsules and the anterior horns are longer for A. vincenti compared to those observed on the current egg capsules, although the anterior horns are barely developed in A. analis egg capsules (Waite 1906; Whitley 1938). The egg capsules of Atelomycterus are smooth with no striations, elongate with a greatly depressed and tapering posterior margin and the long posterior horns are very close together and touch each other in some instances, producing only short tendrils from the posterior end (Whitley 1939; Compagno & Stevens 1993a; Bor et al. 2003). The Atelmycterus egg capsule illustrated by Whitley (1938) is unlikely to be from that genus. Egg capsules are unknown for Aulohalaelurus, but the genus is presumedly oviparous (Springer 1979; Last & Stevens 2009). Members of the genus Bythaelurus variably display retained oviparity, probably giving birth to live young with egg capsules observed in utero being described as thin, fragile bags (eg. B. lutarius, B. clevai), or are egg layers (eg. B. canescens, B. dawsoni, B. hispidus?) producing tough egg capsules (Bass et al. 1975; Springer 1979; Seret 1987; Francis 2006). The egg capsules of B. canescens and B. dawsoni are very similar in overall form of the current egg capsules, although egg capsules of B. dawsoni only possess fine striations, whereas B. canescens egg capsules do have prominent striations (Springer 1979; Francis 2006). The egg capsules and reproduction remain unknown for the Australian representative, B. incanus (Last & Stevens 2009), however, the embryo in question here would have to undergo dramatic ontogenetic changes prior to hatching for it to be considered a member of Bythaelurus. Although egg capsules remain unknown for most Australian Cephaloscyllium (Last et al. 2008c; Last & White 2008), those that are known are highly variable in form. Cephaloscyllium albipinnum, C. hiscosellum, and C. variegatum possess egg capsules that are similar in overall form with the present egg capsule, however, they have tendrils at their anterior ends and are smooth, as may be the case for C. signourum (Last et al. 2008b; White & Ebert 2008; Last & White 2008). Cephaloscyllium laticeps possesses distinctive and unique egg capsules that are laterally expanded, with long tendrils at both the anterior and posterior ends, and pronounced transverse ridges (Whitley 1938, as a synonym of Parascyllium; Springer 1979; Last et al. 2008b). Egg capsules appear to be unknown for the genus Figaro, despite F. boardmani being reported as being oviparous in the literature (Gledhill et al. 2008; Last & Stevens 2009). Egg capsules are also unknown for the only Australian representative of the genus Galeus, G. gracilis (Compagno & Stevens 1993b; Last & Stevens 2009). Elsewhere, Galeus egg capsules are known to vary greatly, from having a distinct waist and long tendrils at both the anterior and posterior ends, or long tendrils only at the posterior ends, to having egg capsule that closely resemble those of Atelomycterus and having fine or no striations (Nakaya 1975; Soto 2001; Iglesias et al. 2002). In the extreme, Galeus polli is known to have retained oviparity and give birth to live young (Springer 1979; Ebert et al. 2006). Halaelurus sellus, the only Australian representative of that genus, has an egg capsule with a distinct waist, no anterior horns, curved posterior margin with reduced horns and tendrils, and no striations (White et al. 2007). Other members of Halaelurus have similarly shaped egg capsules, but variably have anterior tendrils, with anterior and posterior tendrils varying in length between species, but all of them lack striations (Springer & D'Aubrey 1972; Nakaya 1975; White et al. 2007; Akhilesh et al. 2011). Parmaturus bigus is the only Australian representative of that genus (Last & Stevens 2009). Known from a single female specimen, the female aborted an egg capsule on capture that was presumably discarded because it was not described or illustrated (Seret & Last 2007). Elsewhere, the egg capsules of Parmaturus are elongate, with an indistinct waist, poorly to moderately developed anterior and posterior horns and tendrils (Cox 1963; Flammang 2005).

Chondrichthyan egg capsules with longitudinal striations have been described for species within several scyliorhinid genera, including Apristurus (Nakaya 1975; Springer 1979; Ebert et al. 2006; Flammang et al. 2007; Nakaya et al. 2008b), Asymbolus (Whitley 1938), Bythaelurus (Springer 1979, as a synonym of Halaelurus), Cephaloscyllium (Whitley 1938), Galeus (Nakaya 1975; Springer 1979), Schroederichthys (Springer 1979; Gomes & de Carvalho 1995), and Scyliorhinus (Gomes & de Carvalho 1995), in most cases however, these are described as fine striations. Some Cephaloscyllium bear egg capsules with well developed ridges running transversely across the surface, but these are simple ridges and not T-shaped in cross section (Whitley 1938, as a synonym of Parascyllium; Springer 1979; Last et al. 2008b).

Flammang et al. (2007) list the descriptions available for egg capsules of the genus Apristurus and summarise the state of knowledge, to the time of that study, of egg capsules for the genus. Of the eight Australian representatives of Apristurus, egg capsules are known for A. ampliceps, A. australis, A. longicephalus, A. melanoasper, A. pinguis, and A. platyrhynchus (Nakaya 1975; Ivanov 1987; Nakaya & Sato 2000; Iglesias et al. 2004; Flammang et al. 2007, A. ampliceps as Apristurus sp. D sensu Last & Stevens 1994; Kawauchi et al. 2008; Nakaya et al. 2008b; Sato et al. 2008) and the current egg capsules do not correspond to any of those species. Egg capsules remain unknown for A. bucephalus and A. sinensis (Sasahara et al. 2008; White et al. 2008).

Amongst Apristurus, A. laurussonii, A. macrorhynchus, A. manis, A. melanoasper, A. microps, A. pinguis, A. platyrhynchus, A. saldanha, A. spongiceps have longitudinal striations on their egg capsules (Nakaya 1975; Nakaya & Sato 1998; Iglesias et al. 2004; Ebert et al. 2006, egg capsule assumed to be A. saldanha through process of elimination; Flammang et al. 2007; Kawauchi et al. 2008; Nakaya et al. 2008b), and of these, A. laurussonii and those assumed to belong to A. saldanha appear to have the strongest ridges but are simple in cross section, unlike the egg capsules in question.

Of those species whose egg capsules remain unknown, A. bucephalus is known to occur in the south-west of Western Australia (Last & Stevens 2009), a minimum of approximately 1700 km south of where the egg capsules were collected. In contrast, A. sinensis has been recorded from off central Western Australia and off Ashmore Reef (Last & Stevens 2009), approximately 500 km south and 900 km north, respectively, making it the most likely potential candidate based on distribution.

The known depth range of A. sinensis is 940-1290 m (Last & Stevens 2009), whereas the egg capsules were collected from a depth range of 410-452 m. While it is certainly plausible that A. sinensis may exploit a shallower depth for a nursery, its extensive range (western and eastern Australia, South China Sea, and perhaps New Zealand (Last & Stevens 2009)) would suggest that if it did occur at such depths, that it might be reasonable to expect that specimens would have been captured which would corroborate its occurrence at such depths. However, the assignment of A. sinensis to Australian specimens is only tentative and the A. sinensis group in Australia displays three distinct genetic variations. Therefore data recorded for that species elsewhere may not be applicable to Australian examples, and perhaps even the populations within the Australian subset (White et al. 2008; Last & Stevens 2009).

Further, the gross morphology of the embryo does not agree with that of A. sinensis, which has relatively smaller pectoral and pelvic fins, and possesses a longer interspace between the pectoral fin insertion and pelvic fin origin than what is observed on the embryo. The author recognises that interpretation of the embryos' morphology is limited due to ontogeny and associated allometric growth, therefore cannot be considered definitive. However, it is typical in vertebrates that appendages grow only towards the later stages of development, therefore it is unlikely that the fins of the embryo are going to become proportionately smaller, and may in fact become proportionately larger.

The specific identity of the egg capsules in question could not be resolved in the current investigation. While there are potential described species that are candidates, there are morphological and/or distributional inconsistencies preventing any of these potential candidates from being assigned to the egg capsules based on our current knowledge of those taxa. A further possibility, is that the egg capsules belong to an as yet undescribed species of Apristurus. The morphometrics and gross morphology of the embryo suggest this.

Western Australia, and particularly the North West Shelf, remains a poorly surveyed region compared to other areas of Australia, such as southern Western Australia and central eastern Australia, as examples. It is likely that the known distributions and diversity of Apristurus in Western Australian waters may reflect the relatively low sampling effort that the region has received, and that their diversity and distributions are underestimated. Further surveying of Western Australia is needed, particularly of the North West Shelf, in order to resolve species distributions, taxonomic questions, and may ultimately lead to the discovery of the true taxonomic identity of the egg capsules in question.

ACKNOWLEDGEMENTS

I am indebted to Sue Morrison, acting curator of fishes at WAM, for allowing me access to the fish collection and facilities at WAM, and proposing me as a research associate of the museum. I am grateful to Corey Whisson (WAM) for photographing the embryo. I wish to thank Helen Larson, John Pogonoski, and David Ebert, for constructive comments on an earlier draft of this paper. David Ebert made available Brooke Flammangs' unpublished M.Sc. thesis and provided continuing discussions on Apristurus.

Received: 14 January 2011 - Accepted: 11 May 2011

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Brett A. Human

Research Associate. Department of Aquatic Zoology, Western Australian Museum, Locked bag 49, Welshpool DC, Perth WA 6986, Australia. E-mail: brettahuman@gmail.com
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