Arhynchobatidae

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 : 100-104

Arhynchobatidae (softnose skates)

Bathyraja spp. (includes Bathyraja sp. , B. inter-

rupta, B. kincaidi , B. kincaidi / interrupta , B.

mariposa, B. taranetzi , B. violacea ) ( fig. 76)

The analysis yielded a cluster comprised of 19 specimens whose identities are uncertain. The range of pairwise differences among these specimens was 0–24, with an average of pairwise differences of 7.6. These specimens were collected from a diversity of localities in the eastern North Pacific Ocean, including the Gulf of Alaska, Bering Sea, and Strait of Juan de Fuca. Their specific designations, assigned by their original collectors, varied wildly and include Bathyraja sp. , B. interrupta , B. kincaidi , B. kincaidi / interrupta , B. mariposa , B. taranetzi , and B. violacea . The original field identifications have been retained for all specimens, regardless of current thoughts on generic placements and specific synonymies, because their identities of these specimens could not be confirmed. This draws attention to the need for voucher specimens and/or images to confirm species designations and problems created when they are unavailable. It should be noted that only three of these samples come from museum specimens (i.e., B. interrupta GN 6689 5 UW 111883, B. violacea GN 6707 5 UW 48757, and B. mariposa GN 6694 5 UW 47201). In addition, as indicated in figure 76, at least some images are available for another eight specimens. We suspect that the mixed nature of this cluster may reflect either extensive hybridization among closely related forms with different color patterns or complex polymorphism that has hindered accurate identification in the field.

Bathyraja minispinosa (smallthorn skate) ( fig. 76)

The three specimens of this North Pacific species included here were collected from the eastern North Pacific. All three are deposited at the University of Washington (i.e., GN6696 5 UW 47049, GN6697 5 UW 117948, GN6698 5 UW 117950). The range of pairwise differences among the specimens comprising this cluster was 0–4, with an average of 2.7. These grouped most closely with the specimens comprising the Bathyraja spp. cluster; the average of the pairwise differences between specimens of B. minispinosa and those in the Bathyraja spp. cluster was 24.1.

Bathyraja pallida (pallid skate) ( fig. 76)

Four samples from skates identified by their collectors as B. pallida were included in the analysis. Two of these specimens are deposited in museums (GN4668 5 MCZ 167975 and GN5039 5 VIMS 11758). The analysis yielded a single cluster comprised of these four specimens, however, the range of pairwise differences among these specimens was 4–21, with an average of 17.3. We note, however, that while B. pallida is known only from the Bay of Biscay, the specimens included here come from a diversity of localities that include the western Atlantic off New England, the mid-Atlantic, and the Barents Sea. Further investigation of the identity of skates from these regions is required.

Bathyraja shuntovi (narrownose skate) ( fig. 76)

A single specimen of this New Zealand endemic was included in the analysis. This specimen clustered most closely with those of Bathyraja pallida . The average of the pairwise differences between those comprising the latter cluster and the specimen of B. shuntovi was30.5.

Bathyraja trachura (roughtail skate) ( fig. 76)

The specimen of this eastern North Pacific endemic included in the analysis was collected from the Gulf of Alaska.

Bathyraja aleutica (Aleutian skate) fig. 76)

This cluster consisted of a specimen from the Gulf of Alaska and one from the Bering Sea; they differed from one another by 10 bases. The identity of these specimens was confirmed by D. Ebert (personal commun.) following examination of the images of GN5722.

Rhinoraja macloviana (Patagonian skate) ( fig. 76)

Three specimens of this southeastern South American endemic skate, all from the Falkland Islands, were included in the analysis. The range of pairwise differences among the specimens in this cluster was 0–1.

Rhinoraja albomaculata (whitedotted skate)

( fig. 76)

Three specimens of this southern South American endemic species, all collected from the Falkland Islands, were included here. The range of pairwise differences among these specimens was 1–4, with an average of 2.7; they clustered most closely with specimens of R. macloviana , with an average pairwise difference among specimens of 9.7. We note that this was one of several species of Rhinoraja that grouped among the clusters of species of Bathyraja .

Bathyraja brachyurops (broadnose skate) ( fig. 76)

The four specimens of this species included in the analysis were found to comprise a single cluster with a range of pairwise differences among specimens of 0–10, and an average of pairwise differences of 7. All four of these specimens came from the Falkland Islands and thus represent a southeastern part of the distribution of this species, which is known from Chile to northern Argentina.

Rhinoraja magellanica (Magellan skate) ( fig. 76)

A single specimen of this species, collected from the Falkland Islands, was included. This southern South American endemic clustered along with but outside the specimens of B. brachyurops , R. macloviana , and R. albomaculata .

Bathyraja sp. 1 ( fig. 76)

Three unidentified Bathyraja specimens from the Falkland Islands clustered together. The range of pairwise differences among specimens in this cluster was 0–5, with an average of 3.3.

Bathyraja griseocauda (graytail skate) ( fig. 76)

The analysis included two specimens, both from the Falkland Islands, of this southern South American and Antarctic endemic. These specimens differed from one another by a single base. They grouped most closely with the specimens of Bathyraja sp. 1 . The average of the pairwise differences between specimens of what we believe are two distinct species was 22.2.

Bathyraja scaphiops (cuphead skate) ( fig. 76)

The analysis included two specimens of this southwestern Atlantic endemic. These specimens differed from one another by two bases.

Bathyraja cf. taranetzi ( fig. 76)

The analysis included a single specimen collected from the Bering Sea that was originally identified as B. taranetzi , a species currently known only from the Kuril Islands in the western North Pacific. This specimen grouped most closely with the Bathyraja scaphiops cluster; the average of the pairwise differences between these two species was 42. Several other specimens also identified by their collectors as B. taranetzi clustered elsewhere in the analysis (see above). Thus, we have given this specimen the tentative designation of B. cf. taranetzi . The grouping of this specimen among Bathyraja species is noteworthy in the context of the generic distinctions between Rhinoraja and Bathyraja .

Rhinoraja multispinis (multispine skate) ( fig. 76)

Three specimens of this South American endemic species, all collected from the Falkland Islands, were included in the analysis. All three specimens had identical sequences.

Bathyraja parmifera (Alaska skate) ( fig. 76)

Eight specimens tentatively identified as B. parmifera , two of which are deposited at the University of Washington (GN6699 5 UW 111889 and GN6721 5 UW 117306), were included in the analysis. These specimens were all collected from localities in the eastern North Pacific Ocean, including the Bering Sea, and are generally representative of the distribution of this species. The range of pairwise differences among these specimens was 0–12, with an average of 3. The upper end of this range was accounted for by the second specimen in the UW collection. Given that some of the specimens in this cluster were originally identified as B. aleutica , and none of the specimens in this cluster have been firmly established to represent B. parmifera , the identity of this cluster remains to be confirmed. Our results are consistent with those of Spies et al. (2011) and suggest that this species should be recognized as a member of the generic group level taxon Arctoraja .

Bathyraja smirnovi (golden skate) ( fig. 76)

A single specimen of this western North Pacific Ocean species was included. It clustered most closely with the specimens of B. parmifera . The average of the pairwise differences between specimens of these two species was 17.5. Our results are consistent with those of Spies et al. (2011) and suggest that this species should also be recognized as a member of the generic group level taxon Arctoraja .

Bathyraja bergi (bottom skate) ( fig. 76)

A single specimen of this western North Pacific species was included in the analysis. It clustered along with but outside those of B. parmifera and B. smirnovi ; the averages of the pairwise differences between this species and the specimens comprising the clusters of each of the latter two species were 45.5 and 40, respectively. Our analysis suggests that it would also be interesting to explore the generic placement of this species relative to Arctoraja species.

Bathyraja maculata (whiteblotched skate) ( fig. 76)

The analysis included four specimens of this Bering Sea endemic. The range of pairwise differences among these specimens was 1–5, with an average of 3.8.

Brochiraja asperula (prickly deepsea skate) com-

plex ( fig. 77)

The analysis yielded a group that included two clusters, one of which was comprised of 19 specimens of the New Zealand endemic, Brochiraja asperula ; 13 of these came from the Museum of New Zealand, Te Papa Tongarewa (GN6803 5 NMNZ P.040505, GN6804 5 NMNZ P.040506, GN6809 5 NMNZ P.040512, GN6811 5 NMNZ P.040514, GN6812 5 NMNZ P.040515, GN6820 5 NMNZ P.041493, GN6818 5 NMNZ P.041491, GN6795 5 NMNZ P.040465, GN6798 5 NMNZ P.040467,

GN6801 5 NMNZ P.040469, GN6802 5 NMNZ P.040470, GN6821 5 NMNZ P.041494, and GN6817 5 NMNZ P.041490). These specimens were collected from a diversity of localities throughout the waters of New Zealand, including the Chatham Islands, Chatham Rise, Campbell Plateau, and the South Island. The range of pairwise differences among the 19 specimens in this cluster was 0–2, with an average of 0.3. However, the analysis yielded a second smaller cluster in this group comprised of two specimens also in the National Museum of New Zealand, Te Papa Tongarewa (GN6807 5 NMNZ P.040510 and GN6808 5 NMNZ P.040511), collected off Westland on New Zealand’s South Island. The sequences of these two specimens differed from one another by a single base and were somewhat divergent relative to those comprising the specimens of the first cluster. The average of the pairwise differences between specimens of these two clusters was 27.7. The specimens in the smaller cluster have been referred to as Brochiraja cf. asperula .

Brochiraja spinifera (spiny deepsea skate) ( fig. 77)

Sixteen specimens of this New Zealand endemic, all collected from localities in and around New Zealand (e.g., North Island, South Island, Stewart Island, Campbell Island, Auckland Islands, Chatham Rise), were found to comprise essentially a single cluster. A total of 12 of these are from the Museum of New Zealand, Te Papa Tongarewa (GN6794 5 NMNZ P.040464, GN6826 5 NMNZ P.041752, GN6822 5 NMNZ P.041748, GN6827 5 NMNZ P.041753, GN6805 5 NMNZ P.040508, GN6836 5 NMNZ P.045193, GN6806 5 NMNZ P.040509, GN6810 5 NMNZ P.040513, GN6823 5 NMNZ P.041749, GN6824 5 NMNZ P.041750, GN6797 5 NMNZ P.040466, and GN6828 5 NMNZ P.041754). The range of pairwise differences among the 16 specimens in this cluster was 0–13, with an average of 6.1. While there was some evidence of substructure within this cluster, this remains to be explored in more detail. These specimens have all been referred to here as B. spinifera , but this identity needs to be confirmed. This cluster grouped most closely with the specimens of B. asperula and B. cf. asperula . The average of the pairwise differences between specimens of B. spinifera and those of B. asperula was 25.3, and those of B. cf. asperula 31.1.

Brochiraja leviveneta (smooth blue skate) ( fig. 77)

Three specimens of this recently described New Zealand endemic (see Last and McEachran, 2006) were included in the analysis. All three specimens are deposited in the Museum of New Zealand, Te Papa Tongarewa (GN6835 5 NMNZ P.045192, GN6815 5 NMNZ P.040676, and GN6830 5 NMNZ P.041985). The range of pairwise differences among these three specimens was 4–12, with an average of 9.3.

Brochiraja albilabiata (whitelipped skate) ( fig. 77)

This relatively newly described New Zealand endemic (see Last and McEachran, 2006) was represented by a single specimen from the Museum of New Zealand, Te Papa Tongarewa (GN6833 5 NMNZ P.042691). This specimen grouped most closely with the specimens of B. leviveneta . The average of the pairwise differences between specimens of these two species was 28.7.

Insentiraja subtilispinosa (velvet skate) ( fig. 77)

A specimen of this species from the Australian National Fish Collection (GN4629 5 ANFC H 6417-03), collected from Western Australia, was included in the analysis. This specimen clustered outside those of the Brochiraja species. The generic placement of this species follows Last and Stevens (2009).

Notoraja azurea (blue skate) ( fig. 77)

A paratype (GN4631 5 ANFC H 6409- 02) of this recently described Tasmanian endemic (see McEachran and Last, 2008) was included in the analysis. This specimen clustered with the specimen of Insentiraja subtilispinosa , but the sequences of these two species differed by 34 bases.

Brochiraja microspinifera (small prickly skate)

( fig. 77)

The single specimen of this relatively recently described New Zealand endemic (see Last and McEachran, 2006) was includ- ed in the analysis. This specimen is in the Museum of New Zealand, Te Papa Tongarewa (GN6816 5 NMNZ P.041321). It was found to belong to a cluster consisting of the other Brochiraja species , but also of the specimens of Insentiraja subtilispinosa and Notoraja azurea .

Pavoraja nitida (peacock skate) ( fig. 77)

The analysis included three specimens of this Australian endemic species. These specimens differed from one another by 0–4 bases, with an average of pairwise differences of 2.7.

Pavoraja alleni (Allens skate) ( fig. 77)

This Australian endemic was represented by a single specimen from the Australian National Fish Collection (GN4632 5 ANFC H 6419-03). This specimen clustered with those of its congener, P. nitida . The average of the pairwise differences between specimens of these two species was 53.7.

Irolita waitii (southern round skate) ( fig. 77)

The analysis included a single specimen of this southwestern Australian endemic species (GN4630 5 ANFC H 6350-01). This sample clustered along with, but outside all of the specimens representing species of Brochiraja , Insentiraja , Notoraja , and Pavoraja .

Psammobatis sp. ( fig. 77)

In total, 10 specimens of an as yet unidentified species of Psammobatis , all collected from the Falkland Islands, were included in the analysis. They were found to comprise a single cluster and the range of pairwise differences among these specimens was 0–6, with an average of 2.2. It is likely this represents one of the seven Psammobatis species occurring in the western South Atlantic.

Sympterygia bonapartii (smallnose fanskate)

( fig. 77)

Two specimens of this southern South American endemic were included in the analysis. Both specimens came from Argentina and were identical in sequence.

Arhynchobatis asperrimus (longtailed skate)

( fig. 77)

One specimen of this New Zealand endemic was included in the analysis. This specimen is in the Museum of New Zealand, Te Papa Tongarewa (GN6831 5 NMNZ P.042403). It grouped most closely with the specimens of S. bonapartii and the average of the pairwise differences between specimens of these two species was 130.

Sympterygia acuta (bignose fanskate) ( fig. 77)

The two specimens of this southeastern South American species included in the analysis were both collected from Argentina, and are deposited in the Instituto Nacional de Investigación y Desarrollo Pesquero (GN4726 5 INIDEP T 0405 and GN4727 5 INIDEP T 0407). These differed from one another by six bases. They grouped most closely with but outside the specimens of S. bonapartii and A. asperrimus , which suggests that the distinctions between Sympterygia and the currently monotypic Arhynchobatis needs to be explored in more detail. The average of the pairwise differences between S. acuta and S. bonapartii was 129.5; the average of the pairwise differences between S. acuta and A. asperrimus was 137.

Atlantoraja cyclophora (eyespot skate) ( fig. 77)

Three specimens of this species, which is known from Argentina and Brazil, were included in the analysis. These were all collected from Argentina and are deposited in the Instituto Nacional de Investigación y Desarrollo Pesquero (GN4717 5 INIDEP T 0469, GN4718 5 INIDEP T 0471, and GN4719 5 INIDEP T 0474). They were found to comprise a single cluster; the range of pairwise differences among these specimens was 1–4, with an average of 2.7.

Atlantoraja platana (La Plata skate) ( fig. 77)

The analysis included two specimens of this southwestern Atlantic species, both collected from Argentina and both from the Instituto Nacional de Investigación y Desarrollo Pesquero (GN4721 5 INIDEP T 0425 and GN4722 5 INIDEP T 0426). They differed from one another by one base. They grouped most closely with the specimens of A. cyclophora and the average of the pairwise differences between these two species was 38.8.

Atlantoraja castelnaui (spotback skate) ( fig. 77)

The analysis included a single specimen of this southeastern South American species (GN4720 5 INIDEP T 0406). It clustered along with but outside the specimens of its two congeners. The average of the pairwise differences between A. castalnaui and A. cyclophora was 90, and the average of the pairwise differences between A castalnaui and A. platana was 89.5.

Rioraja agassizii (Rio skate) ( fig. 77)

This monotypic genus was represented by two specimens in the analysis, both collected from Argentina and deposited in the Instituto Nacional de Investigación y Desarrollo Pesquero (GN4724 5 INIDEP T 0404 and GN4725 5 INIDEP T 0408). These specimens thus represent a central element of the distribution of this species, which has been reported from throughout much of the eastern coast of South America. These specimens differed from one another by 10 bases and they grouped most closely with the three species of Atlantoraja .