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 : 42-46

Scyliorhinidae (catsharks): group 1

Apristurus melanoasper (black roughscale cat-

shark) complex ( fig. 28)

The analysis included 10 specimens initially identified as Apristurus melanoasper ; two from the western North Atlantic, three from Australia, and five from New Zealand. The analysis yielded two clusters, one consisting of the two Atlantic specimens (which differed from one another by three bases, one of which was deposited in the Yale Peabody Museum [GN1076 5 YPM ICH.010136]) and a second cluster comprised of the eight specimens from Australia and New Zealand (all of which were identical in sequence). The average of the pairwise differences between members of these two clusters was 19.5. Given that A. melanoasper was described from the North Atlantic, we have given the specimens in the Atlantic cluster the designation of A. melanoasper . Although recent work extended the known distribution of this species to include Australia and New Zealand ( Nakaya et al., 2008), the results in this paper do not fully support this decision, so the Australasian cluster has been given the provisional designation of Apristurus cf. melanoasper . Several of the samples in the latter cluster came from specimens in the Australian National Fish Collection (GN4868 5 ANFC H 1391-01 and GN4869 5 ANFC H 1391-03), and several from specimens in the Museum of New Zealand, Te Papa Tongarewa (GN6723 5 NMNZ P.041310, GN6738 5 NMNZ P.042336, GN6740 5 NMNZ P.042569, GN6754 5 NMNZ P.045140). Detailed taxonomic revision of this complex is required in the future.

Apristurus brunneus (brown catshark) ( fig. 28)

The two specimens of this species were collected from California and thus generally represent the northern, but not the potential southern hemisphere elements of this species, which has been reported from Panama, Ecuador, and Peru. These specimens clustered together, independently from the other species of Apristurus included in the analysis ; pairwise difference between these two specimens was three bases.

Apristurus laurussonii ( Iceland catshark) ( fig. 28)

A total of seven specimens, all collected from the North Sea off the coast of Scotland, were included in the analysis. These specimens represent only one of the eastern components of the rather disjunct distribution of this species, which also includes isolated localities off northwestern Africa, Iceland, Massachusetts, and the Gulf of Mexico. The seven specimens were found to comprise a single cluster. The range in pairwise differences seen among these specimens was 0–3, with a mean of 1.4.

Apristurus cf. sinensis (South China catshark)

( fig. 28)

A total of six samples taken from specimens collected from New Zealand, mostly from the Museum of New Zealand, Te Papa Tongarewa (GN6728 5 NMNZ P.042126, GN6749 5 NMNZ P.045142, GN6757 5 NMNZ TMP004690, GN6745 5 NMNZ P.044309, and GN6752 5 NMNZ P.045139), were included in the analysis. They yielded a single cluster with a range in pairwise differences among specimens of 0–3 (with an average difference of 1). These specimens were originally identified as A. sinensis by Kazuhiro Nakaya. However, no specimens from near the type locality of this species (South China Sea) were included in this analysis. Given that a thorough taxonomic revision of this complex is required, the New Zealand form is provisionally referred to here as A. cf. sinensis .

Apristurus sp. 1 ( fig. 28)

This sample, collected from Western Australia, came from a museum specimen (GN4863 5 ANFC H 6411-02). It clustered most closely with A. cf. sinensis ; the range in pairwise differences between these two taxa was 21.2. This result suggests that it represents either an undescribed species, or a known species that was not otherwise represented in the analysis.

Apristrus platyrhynchus (spatulasnout catshark)

( fig. 28)

The analysis included three specimens collected from deepwater near Bass Strait, Australia. It yielded a single cluster with a pairwise difference of 1–2. Although recent taxonomic work on this species extended its known distribution to include Australia ( Kawauchi et al., 2008), it would be ideal to obtain specimens from the type locality for this species, i.e., Japan, for comparison with the Australian specimens. Thus, the specific designation used here is provisional until such time as its identity can be examined in more detail.

Apristurus sp. 2 ( fig. 28)

This specimen was collected from near Bass Strait, Australia, and clustered most closely with A. platyrhynchus ; the range in pairwise differences between these two taxa is 33.3. This result suggests that it too represents either an undescribed species, or a known species that was not otherwise represented in the analysis.

Apristurus exsanguis (flaccid catshark) ( fig. 28)

The analysis included nine samples, five of which came from specimens at the Museum of New Zealand, Te Papa Tongarewa (GN6753 5 NMNZ P.045130, GN6743 5 NMNZ TMP004689, GN6736 5 NMNZ P.042520, GN6733 5 NMNZ P.042176, and GN6732 5 NMNZ P.042519). These specimens are representative of the distribution of this species, which appears to be endemic to New Zealand. Essentially a single cluster resulted from the analysis. The range in pairwise differences within the cluster was 0–4, with an average of 1.9.

Apristurus macrorhynchus (flathead catshark)

( fig. 28)

Our sample included a total of four samples all taken from specimens in the University of Michigan Museum of Zoology (GN1013, GN1014, GN1015, and GN1016 5 UMMZ 231973), and all of which were collected from Taiwan. These specimens are generally representative of the distribution of this species, which is known only from Taiwan and southern Japan. The analysis yielded a single cluster; the range in pairwise differences among specimens within this cluster was 2–6; the average of pairwise differences among specimens was 4.7.

Apristurus ampliceps (roughskin catshark) complex

and Apristurus manis ( fig. 28)

The relatively newly described species A. ampliceps from Australia and New Zealand (see Sasahara et al., 2008), was represented by eight New Zealand samples, all of which came from specimens deposited in the Museum of New Zealand, Te Papa Tongarewa. Also included in the analysis was a single specimen from the Atlantic Ocean, tentatively identified as Apristurus manis . The analysis yielded two clusters, one consisting solely of five New Zealand specimens of A. ampliceps (with a range of pairwise differences among specimens of 0–3 and an average of 1.6), the other consisting of three specimens from New Zealand (with a range of pairwise differences among these three specimens of 4–6 and an average of 5.5) and the specimen of A. manis . The average of the pairwise differences between members of the two clusters of A. ampliceps (excluding the A. manis specimen) was 21.5 suggesting that conspecificity of specimens in these two clusters is doubtful. However, it is unclear which of the two clusters represents the true A. ampliceps . Similarly, the identity of the A. manis specimen remains to be confirmed. Until such time as this subgroup of Apristurus can be examined in more detail, we have designated specimens comprising the first cluster as A. ampliceps 1 and those comprising the second cluster as A. ampliceps 2. We have, however, provisionally retained the designation A. manis for the Atlantic specimen in the second cluster. The average of pairwise differences between A. manis and A. ampliceps 1 was 27.2; the average of pairwise differences between A. manis and A. ampliceps 2 was 20.7. All five specimens of Apristurus ampliceps 1 were vouchered (GN6726 5 NMNZ P.041688, GN6724 5 NMNZ P.041689, GN6725 5 NMNZ P.041994, GN6735 5 NMNZ P.042385, and GN6751 5 NMNZ P.045206); this is also the case for the three specimens of Apristurus ampliceps 2 (GN6727 5 NMNZ P.041993, GN6742 5 NMNZ TMP004687, GN6744 5 NMNZ TMP004691).

Apristurus profundorum (deepwater catshark)

( fig. 28)

Our material included two specimens that we have tentatively identified as A. profundorum . Both specimens were collected from the western North Atlantic but neither is vouchered or represented by images and thus the identity of this cluster remains to be verified; these specimens differed by 2.0.

Galeus sauteri (blacktip sawtail catshark) ( fig. 28)

The five specimens of this species includ- ed in our analysis were all collected from Taiwan and thus represent the center of the distribution of this species, which is also known from southern Japan and the Philippines. All five samples came from specimens deposited in the University of Michigan Museum of Zoology (GN1021, GN1022, and GN1023 5 UMMZ 231966 and GN991 and GN993 5 UMMZ 231974). The analysis yielded a single cluster, with a range of pairwise difference among specimens of 0–5, with an average difference of 3.2 bases. It is interesting to note that this cluster occurred well outside those containing the other species of Galeus included in the analysis (see fig. 29). The average pairwise differences between these were as follows: 173 (between G. sauteri and G. melastomus ), 154.4 (between G. sauteri and G. arae ), 149.4 (between G. sauteri and G. murinus ), and 166.0 (between G. sauteri and G. polli ).

Parmaturus xaniurus (filetail catshark) ( fig. 28)

The five specimens of Parmaturus xaniurus included in the analysis are generally representative of the distribution of this species given that they were collected from Monterey Bay, California, and this species occurs only in the eastern Pacific from California to Mexico. The analysis yielded a single cluster. The range in pairwise differences among members of this cluster was 1–5, with an average of 3.4.

Haploblepharus edwardsii (puffadder shyshark)

( fig. 29)

The analysis included 19 specimens, all collected from South Africa and identified as Haploblepharus edwardsii . The range in pairwise differences among specimens within this cluster were 0–12, with an average of 4.4. This identification requires confirmation.

Halaelurus buergeri (blackspotted catshark) ( fig. 29)

The analysis included five specimens from the Philippines (GN2219 5 JPAG 005, GN2220 5 JPAG 008, GN2222 5 JPAG 115, GN2234 5 JPAG 114, and GN2252 5 RSE 003), examined by Compagno et al. (2005b). It yielded a single cluster; the range in pairwise differences among specimens was 0–4; the average was 2. Compagno et al. (2005b) noted that, while these specimens resembled Halaelurus buergeri , they differed somewhat in body shape and spotting pattern from those collected elsewhere (e.g., Taiwan, Indonesia, and Japan) and thus gave their specimens the designation H. cf. buergeri . However, as a result of work conducted in conjunction with the description of Halaelurus maculosus from Indonesia by White et al. (2007b), W.W. now considers these Philippine specimens to represent H. buergeri .

Halaelurus sellus (speckled catshark) ( fig. 29)

This species was represented in the analysis by only a single specimen collected from Western Australia. This specimen was deposit- ed in the Australian National Fish Collection (GN4893 5 ANFC H 6367-01). It was identified by P.L. as H. sellus , a species recently described from northwestern Australia by White et al. (2007b). This specimen clustered most closely with those of H. maculosus ; the average of the pairwise differences between specimens of these two species was 60.6.

Halaelurus natalensis (tiger catshark) ( fig. 29)

The analysis included three specimens of this South African endemic species. The range in pairwise differences among specimens in this cluster was 3–5, with an average of 4.

Halaelurus lineatus (lined catshark) ( fig. 29)

Both of the specimens of this species included in the analysis were collected from KwaZulu-Natal in South Africa and identified by Geremy Cliff. They were found to comprise a single cluster; their sequences differed from one another by 4. These specimens are representative of the relatively limited distribution of this species. This species clustered most closely with H. natalensis . The average of the pairwise differences between these two species was 80.3.

Holohalaelurus regani (Izak catshark) ( fig. 29)

All 17 specimens of this species included in the analysis were collected off the coast of Western Cape Province and the western regions of Eastern Cape Province, South Africa. The range in pairwise differences among specimens was 0–6, with an average of 2. Our analysis did not include specimens, formerly considered as the ‘‘Natal’’ form of this species, which were described as Holohalaelurus favus by Human et al. (2006).

Galeus melastomus (blackmouth catshark) ( fig. 29)

A total of eight specimens identified as this species were included in this analysis. These were collected from a diversity of localities in the eastern Atlantic including Ireland, Scotland, and Madeira and thus represent the northwestern elements of the distribution of this species. The sample from Madeira was taken from a specimen in the Museu de História Natural e Aquáio in Funchal (GN6627 5 MMF 36798). The analysis yielded a single cluster ; the range in pairwise differences among specimens in this cluster was 0–4; the average of pairwise differences was 2.

Galeus polli (African sawtail catshark) ( fig. 29)

The six specimens of this species, all collected from western South Africa were included. These represent only a small portion of the distribution of this species, which occurs throughout the west coast of Africa, and also in the Mediterranean Sea. The range in pairwise differences among specimens of this species was 0–3. This species clustered most closely with G. melastomus . The average of the pairwise differences between these two species was 51.4.

Galeus arae (roughtail catshark) ( fig. 29)

The analysis included a single Atlantic specimen identified provisionally as this species by Jose Castro. It was divergent from all other included Galeus species. For example, the average pairwise differences between the species with which it clustered most closely were 72.8 ( A. melastomus ) and 81.5 ( A. polli ).

Galeus murinus (mouse catshark) ( fig. 29)

Both specimens of this species included in the analysis were collected from the central to eastern Atlantic; one of these was identified by Neils Roar Hareide. The specimens clustered together independent of the other species of Galeus . The sequences of these two specimens differed by 2.

Apristurus species (second major cluster) ( fig. 30)

Seven additional specimens belonging to the genus Apristurus were included in the analysis. These clustered together, but well outside the 56 specimens considered to represent,14 other species of Apristurus (see fig. 28), questioning the integrity of this genus as currently circumscribed.

Three of the seven specimens in this cluster were identified as one of the long-snouted catsharks, Apristurus australis (pinocchio catshark). These specimens are representative of the known distribution of this Australian endemic species as they were collected from Tasmania and Western Australia. Both samples from Western Australia came from museum specimens (GN4877 5 ANFC H 2573-01 and GN4878 5 ANFC H 2600-04) and clustered with a specimen from Tasmania. The range of pairwise differences among these three specimens was 2–8 (with an average of 6).

Also included in the analysis was a single specimen from New Zealand. While this specimen clustered most closely with the specimens identified as Apristurus australis , the average of the pairwise differences between it and the specimens in the latter subcluster was 62.3. Thus, we have given this specimen the distinct designation Apristurus sp. 3 as it may represent an undescribed species. Although there are no photographs or retained specimens of this catshark, it could be another long-snout species; it could be the long-snouted Apristurus sp. A of Paulin et al. (1989), which has also been informally referred to as A. cf. herklotsi in unpublished checklists of New Zealand fishes. This cluster is most likely referable to the A. longicephalus group defined by Nakaya and Sato (1999).

Two specimens collected from the northeastern Atlantic clustered together (with a sequence difference of 5), independently from all four of the above specimens. The average of the pairwise differences between these Atlantic specimens and those of A. australis was 153.8, and between the Atlantic specimens and Apristurus sp. 3 was 161.5. Thus, specimens in the Atlantic cluster have been given the provisional designation Apristurus sp. 4 ; it is possible they also represent an undescribed species. Clustering with, but well outside these two specimens was a single specimen from the University of Kansas Ichthyology Collection (GN2533 5 KUI 29258), collected from California and identified as Apristurus kampae . The average of the pairwise differences between the specimen of A. kampae and the two specimens of Apristurus sp. 4 was 69.

Asymbolus rubiginosus (orange spotted catshark)

( fig. 31)

Three specimens from Ken Graham, identified as Asymbolus rubiginosus , from off New South Wales, Australia, were included in the analysis. These specimens clustered together and had a range in pairwise differences of 0– 4, with an average of 2.7.

Asymbolus parvus (dwarf catshark) ( fig. 31)

Both samples of this species included in the analysis came from museum specimens (GN4879 5 ANFC H 6415-01 and GN4880 5 ANFC H 6415-02) collected from Western Australia. These specimens are representative of the distribution of this species, which is endemic to Western Australia. The sequences of these specimens differed from one another by 4. The specimens of this species grouped most closely with those of A. rubiginosus ; the average of the pairwise differences between these two species was 35.

Asymbolus analis (gray spotted catshark) ( fig. 31)

All five specimens of this Australian endemic species included in the analysis were collected in the central part of its range. The analysis yielded a single tight cluster with a range in pairwise differences among specimens of 0–2.

complex ( fig. 31)

The nine specimens included in the analysis are generally representative of the distribution of this Australian endemic; six come from southeastern Australia and three from Western Australia. The analysis yielded two distinct clusters: one consisting of specimens from southeastern Australia and one consisting of specimens from Western Australia. The range in pairwise differences among specimens within the southeastern cluster was 0–6, with an average of 2.8; the range for the Western Australian cluster was 1–2, with an average of 1.3. The average of the pairwise differences between clusters was 19.3 suggesting that, as noted by Gledhill et al. (2008), regional variation occurs within this species. In recognition of this variation, we have given the specimens from Western Australia the designation Figaro cf. boardmani , reserving the designation Figaro boardmani for those in the southeastern cluster. The three samples of F. cf. boardmani were taken from specimens in the Australian National Fish Collection (GN4890 5 ANFC H 6414-07, GN4891 5 ANFC H 6414-08, and GN4892 5 ANFC H 6414-10). Taxonomic revision of this species complex is currently being undertaken by researchers at the Australian National Fish Collection.

Bythaelurus dawsoni ( New Zealand catshark)

( fig. 31)

Six specimens of this New Zealand endemic were included in the analysis and they yielded a single tight cluster. The range in pairwise differences among specimens in the cluster was 0–1. The specimens from which five of these six samples were taken are deposited in the Museum of New Zealand, Te Papa Tongarewa (GN6731 5 NMNZ P.042162, GN6737 5 NMNZ P.042731, GN6739 5 NMNZ P.042162, GN6746 5 NMNZ P.044374, and GN6748 5 NMNZ P.044375).