Parascylliidae, 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 : 55-59

Parascylliidae (collared carpetsharks)

Parascyllium collare (collared carpetshark)

( fig. 39)

A total of six specimens of this Australian endemic, collected from Victoria and New South Wales, were included in the analysis. They were found to comprise a tight cluster. The range in pairwise differences among specimens was 0–2, with an average of pairwise differences of 0.9.

HETERODONTIFORMES (bullhead sharks) Heterodontidae (bullhead sharks)

Heterodontus mexicanus (Mexican hornshark)

( fig. 40)

The 22 samples of this species included in the analysis were all collected from the Gulf of California, and thus represent only the more northern elements of an eastern Pacific distribution of this species. Three of these specimens were deposited in the Texas Cooperative Wildlife Collection (GN5178 5 TCWC 7572.01, GN5224 5 TCWC 7576.02, and GN5231 5 TCWC 7583.01) and two in the Instituto de Biologia, Universidad Nacional Autonóma de México (GN5390 5 IBUNAM PE9509 and GN5397 5 IBUNAM PE9511). The range in pairwise differences among these specimens was 0–6, with an average of 1.8.

Heterodontus portusjacksoni (Port Jackson shark)

( fig. 40)

The seven specimens of this species includ- ed in the analysis were collected from Australia in New South Wales and Western Australia and thus span the western and eastern elements of the Australian distribution of this species. The sequences of all seven specimens were identical. Three of these specimens are deposited in the Australian National Fish Collection (GN4841 5 ANFC H 6340-01, GN4842 5 ANFC H 6354-09, and GN4843 5 ANFC H 6354-11).

Heterodontus zebra (zebra bullhead shark) com-

plex ( fig. 40)

A total of three specimens originally identified as H. zebra were included in the analysis. These were collected from Australia, Malaysian Borneo, and Japan. The sample from Australia, taken from a specimen in the Australian National Fish Collection (GN4844 5 ANFC H 6581-01), was found to cluster with the specimens of H. portusjacksoni , away from the other two specimens identified as H. zebra . Whereas the average of the pairwise differences between the Australian specimen and those in the H. portusjacksoni cluster was 13, the average of the pairwise differences between the Australian specimen and those of H. zebra from Japan and Malaysian Borneo was 24. Specimens from Japan and Malaysian Borneo differed from one another by two bases. The average of the pairwise differences between the two specimens from Japan and Malaysian Borneo and those of H. portusjacksoni was 19. These results suggest that the specimen from Australia probably represents an undescribed species distinct from both H. zebra and H. portusjacksoni . Given that the type locality of H. zebra is China, we have referred to specimens from Japan and Malaysian Borneo as H. zebra , and used the designation H. cf. zebra for the Australian form. A taxonomic revision of this group is currently being undertaken by P.L. and W.W.

Heterodontus galeatus (crested bullhead shark)

( fig. 40)

The analysis included four specimens of this eastern Australian endemic, all collected from New South Wales, Australia. The sequences of these four specimens were identical. The average of their pairwise differences from H. portusjacksoni was 59, and from H. zebra was 64.

Heterodontus francisci (horn shark) ( fig. 40)

The nine specimens of this species included in the analysis were all collected from the Gulf of California, and thus represent only the more northern elements of the disjunct eastern Pacific distribution of this species. Two of these specimens were deposited in the Texas Cooperative Wildlife Collection (GN5225 5 TCWC 7576.01 and GN5254 5 TCWC 7567.06). The range in pairwise differences among these specimens was 0–1. The average of the pairwise differences between this species and the other included described species were as follows: between H. mexicanus was 82.9, between H. portusjacksoni was 76.3, between H. zebra was 74.4, between H. cf. zebra was 77.3, and between H. galeatus was 95.3.

SQUALIFORMES (dogfish sharks) Squalidae (dogfish sharks)

Squalus species

The genus Squalus has recently undergone intensive scrutiny and revision with, for example, 11 new species being described in 2007 alone (e.g., see Last et al., 2007d), and with multiple species having been resurrected in the last several years (e.g., see Last et al., 2007d; Ebert et al., 2010). In our treatment of species in this genus, we have attempted to follow this new taxonomy as closely as possible.

Squalus acanthias (piked dogfish) ( fig. 41)

The analysis included a total of 176 specimens of this species, collected from the western North Atlantic off several states including Rhode Island and Maryland, the western South Pacific off New Zealand, Tasmania, and the eastern South Pacific off Chile. The three specimens from Tasmania were vouchered (GN4956 5 ANFC H 2921- 03, GN4957 5 ANFC H 4266-01, and GN4958 5 ANFC H 4876-01). The analysis yielded a single cluster with a range in pairwise differences among specimens of only 0–10, and an average of 2.2. This relatively low variation is one of the most striking results of our analysis given the small size of this essentially benthic species, which belongs to a genus that otherwise shows rather extensive regional diversification.

Squalus suckleyi (spotted spiny dogfish) ( fig. 41)

A total of eight specimens, collected from the eastern North Pacific were included in the analysis. These specimens clustered independently from the specimens of S. acanthias . The specimens of S. suckleyi had a range of pairwise differences of 2–15, with an average pairwise difference of 6.5. The average of the pairwise differences between the specimens of S. suckleyi and those of the S. acanthias was 11.9. This result is consistent with the work of Ward et al. (2007), Hauser (2009), and Verissimo et al. (2010) regarding the distinct nature of specimens from the North Pacific, and thus also supports the resurrection of S. suckleyi for specimens previously identified as S. acanthias collected from localities in the North Pacific as implemented by Ebert et al. (2010).

Squalus sp. ( fig. 42)

In total, eight specimens collected from the western South Atlantic off the coast of Brazil were included in the analysis. The range in pairwise differences among these specimens was 0–7, with an average of 3.1. These specimens have been referred to here preliminarily as Squalus sp. This identity remains to be explored further in the context of the species known to occur off the coast of Brazil (e.g., see Gadig, 2001).

Squalus cf. mitsukurii (shortspine spurdog) ( fig. 42)

Six specimens, originally identified as the Squalus mitsukurii , a species described from Japan, all collected from South Africa, were included in the analysis. The range in pairwise differences among these specimens was 0–2, with an average of 0.9. Conversations with Dave Ebert lead us to believe these specimens represent an undescribed species, which resembles but is distinct from S. mitsukurii . Thus, we have designated this specimen as S. cf. mitsukurii at this time. A taxonomic treatment of this species is currently underway by Dave Ebert. This species clustered with Squalus sp. , Squalus cubensis , and Squalus montalabani . The average of the pairwise differences between S. cf. mitsukurii and these three species was 8.5, 13, and 15 bases, respectively.

Squalus cubensis (Cuban dogfish) ( fig. 42)

Two specimens, collected by John Morrissey from Jamaica and identified as S. cubensis , were included in the analysis. These specimens differed from one another by five bases. They represent the more northern elements of the range of this species, which is distributed as far south as the Falkland Islands. The average of the pairwise differences between S. cubensis , and Squalus sp. , also from the western Atlantic Ocean, was 12.

Squalus montalbani (Philippine spurdog) ( fig. 42)

The analysis included two samples of this recently resurrected (see Last et al., 2007c) Indo-Pacific species, both of which were collected from voucher specimens deposited in the Australian National Fish Collection (GN4981 5 ANFC H 2609-07 and GN4982 5 ANFC H 4623-05). These specimens differed from one another by one base.

Squalus chloroculus (greeneye spurdog) ( fig. 42)

Four specimens of this relatively newly described (see Last et al., 2007c) Australian endemic species, collected from Western Australia, Victoria, and Tasmania, and thus from throughout much of its range, were included in the analysis. These samples all came from specimens deposited in the Australian National Fish Collection (GN4962 5 ANFC H 2564-24, GN4963 5 ANFC H 4775-01, GN4964 5 ANFC H 594-01, and GN4980 5 ANFC H 2606-06). The range in pairwise differences among these specimens was 0–3, with an average of 1.5.

Squalus grahami (eastern longnose spurdog)

( fig. 42)

The two specimens of this newly described, Australian endemic species (see White et al., 2007c) included in the analysis consisted of one paratype (GN4973 5 ANFC H 4682-02 ) and one voucher (GN4972 5 ANFC H 4623- 03 ) in the Australian National Fish Collection. Both specimens were collected from New South Wales, Australia ; they were identical in sequence.

Squalus edmundsi (Edmund’s spurdog) ( fig. 42)

Three specimens of this newly described species (see White et al., 2007c) were included

in the analysis. They consisted of a paratype

(GN4969 5 ANFC H 2605-05) and two voucher specimens (GN4968 5 ANFC H 2591-17 and GN4970 5 ANFC H 3969-15 ) in the Australian National Fish Collection. All three specimens were collected from Western Australia and thus represent the more southern elements of the distribution of this species, which also occurs in Indonesia and Malaysia. These specimens clustered together ; the range in pairwise differences among specimens was 1–4, with an average of 2.7.

Squalus cf. megalops ( fig. 42)

A total of 17 specimens, all from southern Africa, and originally identified as S. megalops , were included in the analysis. The range of pairwise differences among these specimens was 0–7, with an average of 2.9. These specimens comprised a cluster distinct from the Australian endemic S. megalops . The average of the pairwise differences among specimens of this cluster and those of S. megalops was 13.2. At this time, these specimens are considered to represent a distinct, and possibly undescribed species, and have been given the designation Squalus cf. megalops . They clustered most closely with S. brevirostris . The average of the pairwise differences between specimens of S. cf. megalops and S. brevirostris was 10.1.

Squalus brevirostris (Japanese shortnose spurdog)

( fig. 42)

Two samples of this species, both taken from specimens deposited in the Kagoshima University Museum (GN4996 5 KAUM I 186 and GN4995 5 KAUM I 187), were included in the analysis. Both specimens came from the waters off Japan and thus represent the more northern elements of the distribution of this species, which extends to the South China Sea. These two specimens were identical in sequence and clustered most closely with the specimens of S. cf. megalops .

Squalus megalops (shortnose spurdog) ( fig. 42)

The analysis included seven specimens of this species, all but one of which is deposited in the Australian National Fish Collection (GN4974 5 ANFC H 2605-08, GN4975 5 ANFC H 3762-01, GN4978 5 ANFC H

6581-23, GN4976 5 ANFC H 4649-05,

GN4977 5 ANFC H 6581-22, and GN4979 5 ANFC H 6581-24). These specimens were collected from Western Australia, Victoria, and New South Wales and thus represent much of the distribution of this species, which is now considered an Australian endemic (see Last and Stevens, 2009). The analysis yielded a single cluster of these specimens. The range in pairwise differences among specimens in this cluster was 0–6, with an average of 1.9.

Squalus formosus ( Taiwan spurdog) ( fig. 42)

Three specimens of this recently described species ( White and Iglésias, 2011) from Taiwan were included in the analysis. The range in pairwise differences among these specimens was 0–2, with an average of 1.3. All three specimens are deposited in the University of Michigan Museum of Zoology (GN969 5 UMMZ 231956 and GN976 and GN980 5 UMMZ 231963). These specimens clustered most closely with the specimens of S. albifrons . The average of the pairwise differences between specimens in these two clusters was 29.3. Ward et al. (2007) also found that specimens of this species (as ‘‘sp. Taiwan highfin’’) clustered most closely with S. albifrons from eastern Australia.

Squalus albifrons (eastern highfin spurdog)

( fig. 42)

The three specimens of this newly described (see Last et al., 2007e) eastern Australian endemic consist of a holotype (GN4960 5 ANFC H 4627-01), a paratype (GN4959 5 ANFC H 3589-01), and a voucher (GN4961 5 ANFC H 4704-01), all deposited in the Australian National Fish Collection. The range in pairwise differences among these specimens was 1–2, with an average of 1.3.

Squalus japonicus (Japanese spurdog) ( fig. 42)

All three specimens of this species included in the analysis were collected from Taiwan and thus represent only the more northern elements of the distribution of this species. One of the specimens was deposited in the University of Michigan Museum of Zoology (GN975 5 UMMZ 231962). All three specimens were identical in sequence.

Squalus nasutus (western longnose spurdog)

( fig. 42)

The two specimens of this newly described species (see Last et al., 2007b) included in the analysis were collected from Western Australia. One of these was deposited in the Australian National Fish Collection (GN4983 5 ANFC H 6413-01). These specimens differed from one another by one base. They clustered most closely to the S. japonicus cluster, as was also seen by Ward et al. (2007). The range of pairwise differences between specimens in these two species was 9.5.

Squalus crassispinus (fatspine spurdog) ( fig. 42)

The analysis included three specimens of this newly described (see Last et al., 2007a) species, two of which are paratypes (GN4965 5 ANFC H 4649-03 and GN4966 5 ANFC H 4649-04 ) and one a voucher (GN4967 5 ANFC H 6412-01 ), all deposited in the Australian National Fish Collection. The analysis yielded a single cluster ; the range in pairwise differences among these specimens was 1–8, with an average of 5.3.

Cirrhigaleus australis (southern Mandarin dogfish)

( fig. 42)

The two specimens of this newly described species (see White et al., 2007a) included in the analysis consisted of the holotype (GN4944 5 ANFC H 5789-01) from Tasmania, Australia, and a voucher collected from New Zealand. These specimens differed by four bases. We believe this result supports the suggestion of White et al. (2007a) that C. australis occurs in New Zealand.

Cirrhigaleus asper (roughskin spurdog) ( fig. 42)

A single specimen of C. asper collected off Florida was included in the analysis. This specimen clustered with the two specimens of C. australis . The average of the pairwise differences between this specimen and those comprising the cluster of C. australis was 58.5.