Scyliorhinidae, T.N.Gill, 1862

Elasmobranch, Its Implications For Global, Parasitology, Diversity And, Naylor, G. J. P., Sc, Caira, J. N., Ct, Jensen, K., Ks, Rosana, K. A. M., Fl, White, W. T., Csiro, Tas, Last, P. R., Csiro & Tas, 2012, A Dna Sequence-Based Approach To The Identification Of Shark And Ray Species And Its Implications For Global Elasmobranch Diversity And Parasitology, Bulletin of the American Museum of Natural History 2012 (367), pp. 1-262 : 47-50

Scyliorhinidae (catsharks): group 2

Atelomycterus marmoratus (coral catshark)

( fig. 33)

In total, 10 specimens of this species were included in the analysis: nine were collected from Malaysian Borneo and one from the Philippines. These specimens represent eastern elements of the distribution of this species, which extends as far west as Pakistan. The analysis yielded a single cluster. The range in pairwise differences within the cluster was 0–11, with an average of pairwise differences of 3.8. There was some evidence of geographic structure within the cluster in that the spe- cimens from Borneo grouped together. The average of the pairwise differences between specimens in the Borneo cluster and the specimen from the Philippines was 9.9. We note that the sample from the Philippines, which came from a museum specimen (GN2235 5 JPAG 044 ), was treated by Compagno et al. (2005b) ; one of the specimens from Borneo was deposited in the IPPS ( GN3705 5 IPPS BO495 ) .

Aulohalaelurus labiosus (blackspotted catshark)

( fig. 33)

The three specimens of this species were all collected from Western Australia and are thus representative of the distribution of this Western Australian endemic. Two samples came from specimens deposited in the Western Australian Museum (GN2268 5 WAM P 31670-001 and GN2269 5 WAM P 31671- 001). The analysis yielded a single cluster. The range in pairwise differences among specimens in this cluster was 4–7, with an average of 5.3.

Atelomycterus marnkalha (eastern banded catshark)

( fig. 33)

All three specimens of this species were collected from the Torres Strait, off Australia, and represent the central elements of its distribution, which extends eastward to New Guinea. All three samples were taken from specimens in the Australian National Fish Collection (GN4881 5 ANFC H 6144-01, GN4882 5 ANFC H 6145-01, and GN4883 5 ANFC H 6146-01). The analysis yielded a single tight cluster, with a range in pairwise differences among members of this cluster of 2–4, and an average of 2.7. It was interesting that this species clustered with Aulohalaelurus labiosus . The average of the pairwise differences between these two species was 158.3 and between A. marnkalha and its congener A. marmoratus 155.6.

Parmaturus sp. ( fig. 33)

Six samples, all collected from New Zealand and deposited in the Museum of New Zealand, Te Papa Tongarewa (GN6734 5 NMNZ P.045528, GN6730 5 NMNZ P.042517, GN6741 5 NMNZ P.044582, GN6747 5 NMNZ P.044583, GN6750 5 NMNZ P.044578, and GN6755 5 NMNZ P.042524), were identified only to genus. The

analysis yielded a single cluster with the range

in pairwise differences among specimens being 0–3, with an average of 1.2. This taxon grouped well away from the cluster comprised of its congener, Parmaturus xaniurus (see fig. 28); the average of pairwise differences between these two species was 190.5. This suggests that the identity of these specimens needs to be examined more closely, both at the specific and generic levels. The only other species of Parmaturus currently known from New Zealand is P. macmillani . This taxon is currently under taxonomic investigation by P.L., Bernard Séret, and Keiichi Sato.

Schroederichthys bivius (narrowmouth catshark)

( fig. 33)

Both specimens of this species were collected from Argentina and thus represent the eastern elements of the distribution of this species, which extends throughout much of coastal Chile and Argentina. The sequences of these two specimens were identical.

Poroderma species

Given they exhibited the least amount of intrageneric variation seen in the analysis, a haplotype map was generated for the two sympatric species of Poroderma . This map ( fig. 90) shows two relatively tight haplotype clusters, with no overlap between species, suggesting that although divergence is low the species are distinct.

Poroderma pantherinum (leopard catshark) ( fig. 34)

In total, 16 specimens of this species were included in the analysis. These specimens came from a diversity of localities throughout South Africa and thus represent much of the distribution of the species. The analysis yielded a single cluster. The range in pairwise differences among specimens was 0–6, with an average of 3.

Poroderma africanum (striped catshark) ( fig. 34)

The 12 specimens of this species were collected from South Africa and thus are representative of the distribution of this South African endemic. The analysis yielded a single cluster, which was tightly allied with the cluster of specimens of P. pantherinum . The range in pairwise differences among specimens in the P. africanum cluster was 0–2, with an average of 1.3. The

average of the pairwise differences between

specimens of the two Poroderma species was 6.3.

Scyliorhinus capensis (yellowspotted catshark)

( fig. 34)

A total of 15 specimens of this southern Africa endemic were included. The range of pairwise differences among these specimens was 0–2; the average was 0.6.

Scyliorhinus canicula (smallspotted catshark)

( fig. 34)

The single specimen of this species was collected from the eastern Atlantic. It grouped most closely with the specimens of the southern Africa endemic S. capensis . The average of the pairwise differences between specimens of these two species was 57.5.

Scyliorhinus retifer (chain catshark) ( fig. 34)

Also included in the analysis were three specimens preliminarily identified as Scyliorhinus retifer . One of these came from a specimen in the University of Kansas Ichthyology Collection (GN2530 5 KUI 26984) that was collected from the mid-Atlantic bight. The others were collected from the northwestern Atlantic and Gulf of Mexico. The analysis yielded a single, divergent cluster. However, the range in pairwise differences among these specimens was substantial, at 9–22, and the average was 14.7.

Scyliorhinus stellaris (nursehound) ( fig. 34)

A single specimen of S. stellari s, collected from the eastern Atlantic, was included. This specimen grouped along with, but outside its congeners. The average pairwise difference between the specimen of S. stellaris and those in the S. retifer cluster was 73.3, between S. stellaris and the specimen of S. canicula 73, and between S. stellaris and S. capensis 66.5.

Cephaloscyllium variegatum (saddled swellshark)

( fig. 34)

The analysis included six specimens of this species from Australia, one of which was included among the other material examined by Last and White (2008c) in the original description of this species (GN4889 5 ANFC H 3580-01). The range in pairwise differences among specimens in this cluster was 0–2, with an average of 0.9.

Cephaloscyllium albipinnum (whitefin swellshark)

( fig. 34)

The three samples of this species included in the analysis were taken from the holotype (GN4887 5 ANFC H 5314-11), a paratype (GN4888 5 ANFC H 5940-01), and an additional examined specimen (GN4886 5 ANFC H 3588-01) included in the original description of this relatively new species from New South Wales and Tasmania, Australia (see Last et al., 2008b). The range in pairwise differences among these three specimens was 1–2, with an average of 1.3.

Cephaloscyllium hiscosellum (Australian reticulate

swellshark) ( fig. 34)

The specimen of this species included in our analysis was a paratype of this Western Australian species (GN4884 5 ANFC H 6419- 01), described by White and Ebert (2008). It clustered outside but along with the specimens of C. albipinnum . The average of the pairwise differences between this specimen and those of C. albipinnum was 44.3.

Cephaloscyllium umbratile (Japanese swellshark)

( fig. 34)

The four samples for this species included in the analysis come from specimens in the University of Michigan Museum of Zoology (GN981 5 UMMZ 231960 and GN982, GN1017, and GN1018 5 UMMZ 231967) originally identified as C. isabellum . These specimens were all collected from Taiwan. However, Schaaf-Da Silva and Ebert (2008) clarified the status of C. umbratile as a valid species having been previously placed in synonymy with C. isabellum . Thus, the specimens included here have been provisionally identified as C. umbratile . As such, they represent the southern portion of the western North Pacific distribution of this species. The range in pairwise differences among specimens of was 0–4, with an average of 2.7. Schaaf-Da Silva and Ebert (2008) also described two new species of Cephaloscyllium from Taiwan, so the identity of these specimens should be confirmed.

Cephaloscyllium sp. 1 (Philippine swellshark)

( fig. 34)

Our analysis included two specimens of Cephaloscyllium collected from the Philippines, both of which were treated by Com- pagno et al. (2005b) (GN4361 5 JPAG 231 and GN4352 5 JPAG 232) as Cephaloscyllium sp. 1 . These specimens clustered together and their sequences differed from one anoth- er by three bases. However, this species needs to be critically compared with C. pardelotum and C. maculatum both described by Schaaf- DaSilva and Ebert (2008) from Taiwan, as well as with C. pictum , recently described from Indonesia by Last et al. (2008c). These specimens clustered mostly closely with specimens of C. umbratile . However, the average of the pairwise differences between these two clusters was 47.8.

Cephaloscyllium laticeps (Australian swellshark)

( fig. 34)

In total, six specimens, all collected from southeastern Australia were included in the analysis. One of these samples was taken from a museum specimen (GN4885 5 ANFC H 3581-01). The range in pairwise differences among specimens in this cluster was 0–5, with an average of 1.9.

Cephaloscyllium ventriosum (swellshark) ( fig. 34)

One sample of this species, taken from a specimen in the University of Kansas Ichthyology Collection (GN2529 5 KUI 28129) collected from California, was included in the analysis. This specimen was grouped with all other Cephaloscyllium species , but was by far the most divergent species. The average of the pairwise differences between this specimen and those of the six other Cephaloscyllium species was 109.

LAMNIFORMES (mackerel sharks) Lamnidae (mackerel sharks)

Isurus oxyrinchus (shortfin mako) ( fig. 35)

In total, 24 specimens of I. oxyrinchus were included in the analysis, which yielded a welldefined cluster. The range in pairwise pdifferences among specimens was 0–21, with an average of 10.6. This cluster consisted of two fairly well-defined subclusters, one of which exhibited more sequence heterogeneity than the other; this is also reflected in the haplotype map for phenotype ( fig. 91A). However, as illustrated by the haplotype map of geography ( fig. 91B) there was no apparent geographic pattern to these subclusters, each of which was comprised of specimens from the western North Atlantic, Gulf of Mexico, California, Gulf of California, and Taiwan; one subcluster also included a specimen from South Africa and the other also included a specimen from Vietnam. The range in pairwise differences among specimens in the subcluster including the specimen from South Africa was 0–4, with an average of 2.4. The range in pairwise differences among specimens in the other subcluster was 0–14, with an average of 6.8. The average of the pairwise differences between specimens in the two subclusters was 17.2. In the absence of morphological or geographic data to support this distinction, we have tentatively identified all 24 members of the cluster as I. oxyrinchus . Nonetheless, these results are interesting in light of previous workers (e.g., Moreno and Morón, 1992; Heist et al., 1996; Schrey and Heist, 2003), all of who provided evidence of population structure among makos globally.

Isurus paucus (longfin mako) ( fig. 35)

A total of six specimens of this species were included in the analysis, four from the western North Atlantic, one from the Caribbean Sea, and one from the Gulf of Mexico. These specimens comprised a single cluster ; the range in pairwise differences among specimens in this cluster was 0–8, with an average of 3.3. The average of the pairwise differences between the six specimens of this species and the 24 specimens of I. oxyrinchus was 110.3.

Carcharodon carcharias (great white shark)

( fig. 35)

The 17 specimens of this species included in the analysis were collected from South Africa, the western North Atlantic, California, and South Australia. The analysis yielded a single cluster, with two subclusters. The range in pairwise differences among all 17 specimens was 0–20, with an average of 4.2. One subcluster was comprised of the specimens from South Africa and the western North Atlantic, the other comprised of specimens from California and South Australia (i.e., from localities in the Pacific Ocean). The range in pairwise differences among specimens in the former subcluster was 0–2; the two specimens in the latter subcluster differed from one another by seven bases. On average, pairwise difference between specimens of the two subclusters was 16.3. This intriguing result, which is consistent at least in part with the findings of Pardini et al. (2001) and Jorgensen et al. (2009), suggests that some consideration should be given to the potential lack of conspecificity of C. carcharias from the Atlantic and Indian oceans, and those from the Pacific Ocean.

Lamna nasus (porbeagle shark) ( fig. 35)

The analysis of 10 specimens yielded a single relatively divergent cluster with some evidence of two subclusters within. The range in pairwise differences among all 10 specimens was 0–15, with an average of 6.8. One subcluster consisted of the two specimens from Tasmania, both vouchered (GN2261 5 AMS I32756-002 and GN2262 5 AMS I32756-001), and the other consisted of specimens collected from the northeastern and northwestern Atlantic. The two specimens from Tasmania differed from one another by nine bases. The range in pairwise differences among specimens in the other subcluster was 0–8. The average of the pairwise differences between specimens from the Tasmanian subcluster and those from the Atlantic Ocean subcluster was 14. These results suggest that some consideration should be given to the possibility of the existence of taxonomic variation among porbeagles between ocean basins.

Lamna ditropis (salmon shark) ( fig. 35)

Two specimens of this north Pacific– dwelling species, both from Japan, were included. They differed by 4. The average of the pairwise differences between L. ditropis and L. nasus was 68.