Rajidae

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 : 93-100

Rajidae (skates)

Dipturus innominatus ( New Zealand smooth skate)

( fig. 71)

In total, 11 specimens of this New Zealand endemic were included in the analysis. They were found to comprise a single cluster and the range in pairwise differences among specimens in this cluster was 0–6, with an average of 2.2.

Dipturus gudgeri (bight skate) ( fig. 71)

The differences between sequences of the three specimens of this Australian endemic that were included in the analysis ranged from 1–7, with an average of 4.7. These grouped most closely with the specimens of D. innominatus . The average of the pairwise differences between specimens of these two species was 29.2.

Dipturus sp. 4 ( fig. 71)

The analysis included two skate specimens collected from the Philippines (GN4339 5 BRU 096 and GN4353 5 JPAG 091) that were treated by Compagno et al. (2005b) as Dipturus sp. 4 . These specimens differed from one another by 13 bases. As noted by Compagno et al. (2005b) these specimens may represent an undescribed species in this genus.

Dipturus healdi (Heald’s skate) complex ( fig. 71)

Two specimens, both of which had provisionally been identified as Dipturus healdi , were included in the analysis. These specimens differed from one another by 56 bases. Two morphs referable to D. healdi were discussed by Last et al. (2008d), and a specimen of each morph was included in this analysis. The specimen of the northern morph (GN6788 5 ANFC H 6574-20) is consistent with D. healdi and clusters well away from the southern morph (GN6789 5 ANFC H 6419-04), which is provisionally referred to as Dipturus cf. healdi ; these forms are indisputably separate species. Dipturus healdi clustered most closely with Dipturus sp. 4 , with an average of the pairwise differences between these two taxa of 39.5.

Dipturus tengu (acutenose skate) ( fig. 71)

The analysis included one specimen identified as this species, collected from Taiwan. If this identification is correct, this specimen is representative of a central element of the distribution of this species, which is known to occur from Japan to the Philippines.

Dipturus springeri (roughbelly skate) ( fig. 71)

The four specimens of this species, all collected from South Africa, were identical in sequence. They represent the central southwestern elements of the distribution of this species, which occurs along the coast of Africa from Angola to Somalia as well as Madagascar.

Dipturus australis (Sydney skate) ( fig. 71)

Six specimens of this species, which is endemic to eastern Australia, were included in the analysis. They formed a single cluster. The range of pairwise differences among these specimens was 0–2, with an average of 0.7.

Dipturus cerva (whitespotted skate) ( fig. 71)

Three specimens of this South Australian endemic were included in the analysis. They were identical in sequence.

Dipturus confusus (longnose skate) ( fig. 71)

Two specimens of this recently described (see Last, 2008) southeastern Australian endemic species were included here. They were identical in sequence and clustered most closely with the two specimens of D. cerva . The average of the pairwise differences among specimens of these two Australian endemic species was 18, and between this species and its other Australian congener, D. australis , was 30.3.

Zearaja chilensis (yellownose skate) ( fig. 71)

The eight specimens of this species includ- ed here had a range of pairwise differences of 0–5, with an average of 2; they comprised a single cluster. These specimens all came from a single locality in Chile and thus represent the western part of the distribution of this species, which occurs throughout the coastal regions of southern South America and the Falkland Islands .

Zearaja nasuta ( New Zealand rough skate)

( fig. 71)

Five specimens of this New Zealand endemic were included in the analysis. These specimens had a range of pairwise differences of 1–5, with an average of 3.2. They comprised a cluster that grouped most closely with the specimens of Z. chilensis . The average of the pairwise differences between specimens of these two species was 10.3.

Zearaja flavirostris (Falklands skate) ( fig. 71)

All three specimens identified as this species included in the analysis, were provid- ed by Joost Pompert, of the Falkland Islands Fisheries Department, extending the range of this species from central Chile to the Falkland Islands. They comprised a single cluster, with a range of pairwise differences among specimens of 0–3 and an average of 2. They grouped most closely with the cluster consisting of Z. chilensis and Z. nasuta . The average of the pairwise differences between specimens of Z. flavirostris and those of Z. chilensis was 29.5, and those of Z. nasuta was 33.9. Given these differences, we have recognized Z. flavirostris as a species distinct from Z. chilensis , despite the suggestion that these two species may be synonyms (e.g., Last and Gledhill, 2007).

Dipturus leptocauda (thintail skate) ( fig. 71)

Both specimens identified as this species were collected from the Falkland Islands, extending the range of this species from Brazil and Uruguay south to include the Falkland Islands. The sequences of these specimens were identical to one another.

Dipturus batis (gray skate) and Dipturus oxy-

rhinchus (sharpnose skate) complex ( fig. 71)

The analysis included a total of 12 specimens originally identified as D. batis and two identified as D. oxyrhinchus . These specimens come from several localities in the eastern North Atlantic and were found to comprise a single group, exhibiting considerable internal structure. The largest cluster consisted of eight specimens from the eastern North Atlantic and three from the Azores. The range of pairwise differences among the 11 specimens in this cluster was 0–8, with an average of 4.3. However, there was evidence of structure within this cluster, with six of the specimens from the eastern North Atlantic (identified as D. batis by Bernard Séret) comprising a subcluster distinct from that consisting of the three specimens from the Azores identified locally as D. batis and two specimens from the eastern North Atlantic identified as D. oxyrhinchus , also by Bernard Séret. The average of the pairwise differences among specimens in these two subclusters was 7.5. In order to call attention to this discrepancy, we have retained the original identifications of these specimens. This result suggests that the identity of specimens of D. batis from the Azores needs to be investigated in more detail.

One specimen, also collected from the eastern North Atlantic, grouped outside this subcluster of 11 specimens. The average of the pairwise differences between this specimen and those in the D. batis and D. oxyrhinchus subcluster was 23.7. This specimen has been given the designation Dipturus cf. batis 1. Finally, two specimens, both collected from Norway, grouped outside all 12 other specimens in the cluster overall; these two specimens were identical with one another in sequence. The average of the pairwise differences between these specimens and those in the D. batis and D. oxyrhinchus subcluster was 16.7, and between D. cf. batis 1 was 37. These two specimens have been given the designation Dipturus cf. batis 2.

These results suggest that it is likely these specimens represent several distinct species, all of which bear a morphological resemblance to Dipturus batis . Recent published work by Iglésias et al. (2009) reveals that the D. batis complex likely consists of two species, both of which should be considered as valid, i.e., D. cf. flossada and D. cf. intermedia . Following these designations and morphological characters provided by Iglésias et al. (2009), the specimens with images in the second subcluster (GN5160, GN5165 and GN6546) are consistent with D. cf. intermedia . Thus, this second subcluster, which includes the two D. oxyrhinchus specimens, may represent D. cf. intermedia . There is a possibility that one of the other subclusters represents D. cf. flossada and one represents D. oxyrhinchus . Another species which Iglésias et al. (2009) reported as very close to these species is the Norwegian skate D. nidarosiensis , and it is possible the two specimens from Norway may represent that species. Additional specimens, with designat- ed vouchers and confirmed identifications based on the work of Iglésias et al. (2009), are required to resolve this issue. The lack of specific locality data for our eastern North Atlantic specimens makes it difficult for us to compare our results with those of Griffiths et al. (2010), except to note that, like these authors, we found specimens identified as D. oxyrhinchus to cluster among specimens identified as D. batis .

Dipturus laevis (barndoor skate) ( fig. 71)

The three specimens of this unusually large skate included in the analysis all came from the western North Atlantic and are thus generally representative of the distribution of this species. One of these specimens is in the Harvard Museum of Comparative Zoology MCZ (GN2603 5 MCZ 159202 and TCWC 11021.01). The range of the pairwise differences among these specimens was 0–1. These specimens grouped along with, but outside the 14 specimens in the Dipturus batis and Dipturus oxyrhinchus complex.

Dipturus pullopunctatus (slime skate) ( fig. 71)

Three specimens of this southern African endemic, all collected from South Africa, were included. These specimens were identical in sequence.

Spiniraja whitleyi (Melbourne skate) ( fig. 71)

Two specimens of this southern Australian endemic species were included in the analysis. These differed from one another by two bases and they grouped along with but outside the Dipturus and Zearaja species.

Raja sp. 1 ( fig. 71)

A single specimen collected from Taiwan clustered independently from all specimens of Dipturus , Zearaja , Spiniraja , and Raja . It is possible that this specimen, for which an image is available, represents a known species, but we have been unable to assign a specific identification with confidence at this time; its generic designation also requires confirmation.

Raja rhina (longnose skate) ( fig. 71)

This northeastern Pacific species was represented by 17 specimens ; the analysis yielded a single cluster. The range of pairwise differences among the specimens in this cluster was 0–8, with an average of 1.3. Two of these specimens are deposited in the University of Washington Fish Collection (GN6712 5 UW 47645 and GN6713 5 UW 49457 ). This species grouped well away from the specimens of most of the other Raja species included in the analysis (see figs. 73, 75). This is consistent with McEachran and Dunn (1998) and Compagno (2005b), who listed this species, among others, as comprising a ‘‘ North Pacific Assemblage’ ’ that may represent an undescribed genus .

Raja binoculata (big skate) ( fig. 71)

The analysis included two specimens of this northeastern Pacific species. These specimens differed by six bases. This species, like Raja rhina , grouped well outside specimens of the other Raja species included in the analysis (see figs. 73, 75). This is consistent with McEachran and Dunn (1998) and Compagno (2005b) who listed these two species, among others, as comprising a ‘‘North Pacific Assemblage’’ that may represent an undescribed genus. The average of the pairwise differences between specimens of this species and those of Raja rhina was 111.2.

Okamejei cairae (Borneo sand skate) ( fig. 72)

Thirteen specimens of this recently described species (see Last et al., 2010a) were included in the analysis. These were all collected from Sarawak in Malaysian Borneo, Pontianak in West Kalimantan ( Indonesia), or Vietnam. The specimens from Vietnam extend the northern boundary of the distribution of this species. They comprised a single cluster and the range of pairwise differences among them was 0–4, with an average of 2.1. All nine specimens from Kalimantan were vouchered (GN4730 5 ANFC H 7099-07, GN4731 through GN4735 5 ANFC H 7099-01 through ANFC H 7099- 05, GN4737 5 ANFC H 7099-06, GN4729 5 MZB 17176, and GN4736 5 MZB 17177).

Okamejei cf. porosa ( fig. 72)

Five specimens collected from Japan were included in the analysis. While the range of pairwise differences among these specimens was only 0–8 (with an average of 3.9) there was substantial morphological variation among these specimens (e.g., in color pattern, disc shape, etc.), based on images for four of the five specimens. These specimens, although they also resemble Okamejei meerdervoortii , are provisionally referred to here as Okamejei cf. porosa until their identities can be examined in more detail. These specimens grouped most closely with those of Okamejei cairae . However, the average of the pairwise differences between the two species was 96.

Okamejei hollandi (yellow-spotted skate) ( fig. 72)

A total of 14 specimens, identified as O. hollandi by Last et al. (2010c), were included in the analysis. These specimens were all collected from Pontianak in West Kalimantan ( Indonesia) ; one was vouchered (GN4738 5 ANFC H 7099-08 ). The range of pairwise differences among these 14 specimens was 0– 13, with an average of six bases. The average of their pairwise differences relative to specimens of Okamejei cairae was 106.9 ; the average of their pairwise differences relative to specimens of Okamejei cf. porosa was 107.6.

Raja miraletus (brown skate) complex ( fig. 73)

In total, 23 specimens that were generally consistent with the color pattern of R. miraletus (i.e., in their possession of small dark spots and a pair of distinctive ocelli on their discs) were included. These consisted of 17 specimens from Senegal and six from South Africa. The analysis yielded three distinct clusters. Two of these clusters consisted of specimens from Senegal and the third of specimens from South Africa. The average of the pairwise differences between specimens in the two Senegal clusters was 99.3. Morphologically, specimens of these two clusters differed most conspicuously in coloration; in one form the small dark disc spots were interspersed with numerous white spots, whereas specimens of the other form exhibited few, if any, small white spots. Specimens of the first Senegal cluster exhibiting the latter color pattern were designated R. miraletus . The more heavily spotted specimens comprising the second cluster of specimens from Senegal were given the designation Raja cf. miraletus 2. The range of pairwise differences among specimens in the R. miraletus cluster was 0–7 (with an average of 2) and among specimens in the R. cf. miraletus 2 cluster was 1–4 (with an average of 2.4).

The cluster of specimens from South Africa had a range of pairwise differences of 0–2 (with an average of 1.2). They most closely resembled R. cf. miraletus 2 in color pattern. However, they clustered more closely with the specimens of R. miraletus . These specimens have been given the designation Raja cf. miraletus 1 until such time as the taxonomy of this complex can be examined in more detail. The average of the pairwise differences between R. cf. miraletus 1 and R. cf. miraletus 2 was 85.2 and between R. cf. miraletus 1 and R. miraletus was 49.1. Our results suggest that the differences seen among these three forms may reflect the existence of several distinct species in these regions as was observed by McEachran et al. (1989). However, it is important to note that several known species from one or more of these regions (e.g., R. herwigi and R. ocellifera ) were not included in the analysis.

Raja clavata (thornback skate) ( fig. 73)

Eight of the specimens of this species were collected from the eastern North Atlantic and the Azores ; the ninth specimen came from the Black Sea. The range in pairwise differences among all nine specimens was 0–11, with an average of 6.1. The analysis yielded some evidence of a subcluster comprised of the specimens from the Azores, and the other comprised of the specimens from the eastern North Atlantic and Black Sea. The average of the pairwise differences among specimens in these two subclusters was 9.5. The range within the Azores subcluster was 0–3, and within the North Atlantic subcluster 1–3. These results parallel the situation in the D. batis complex, which provides evidence that the fauna of the Azores may differ somewhat from that of their mainland counterparts. This result is consistent with that of Chevolot et al. (2006) who found regional differentiation between specimens from the European continental shelf and the Azores .

Raja straeleni (biscuit skate) ( fig. 73)

The nine specimens of this species, which occurs throughout much of the western coast of Africa, were all collected off South Africa. The range in pairwise differences among specimens of this species was 0–3, with an average of 1.6. This species grouped most closely with Raja clavata . The average of the pairwise differences between these two species was 23.1.

Raja asterias (Atlantic starry skate) ( fig. 73)

The analysis included three specimens of this species, all collected from Spain and generally representative of the eastern elements of this primarily Mediterranean-dwelling species. The range of pairwise differences among specimens was 1–4, with an average of 2.7. The analysis yielded a single cluster, which grouped most closely with that consisting of R. clavata and R. straeleni . The average of the pairwise differences between R. asterias and R. clavata was 62.1, and between R. asterias and R. straelini was 66.1.

Raja montagui (spotted skate) ( fig. 73)

A single specimen of this species was included here. This specimen was collected from the coast of England and thus represents a northerly element of the distribution of this species, which extends throughout the Mediterranean Sea and northern coast of Africa. It grouped most closely with but outside the Raja clavata , Raja straeleni , and Raja asterias specimens. The average of the pairwise differences between this specimen and those of R. clavata was 66.1, between this specimen and those of R. straeleni 70.3, and between this specimen and those of R. asterias 78.3.

Rajella fyllae (round skate) ( fig. 74)

The analysis included 18 specimens from the eastern North Atlantic (identified by Bernard Séret) and three from Norway. These were found to comprise a single cluster and the range of pairwise differences among these 21 specimens was0–6,withanaverageofpairwisedifferencesof 1.7. These specimens represent the more eastern parts of the distribution of this species, which occurs throughout the entire North Atlantic.

Rajella sp. ( fig. 74)

Two specimens from the Harvard Museum of ComparativeZoology (GN4664 5 MCZ167899 and GN4665 5 MCZ 167900), both collected on the continental shelf off New England, were included. These two specimens were identical in sequence. They grouped most closely with the two specimens of Rajella caudaspinosa ; these two species were, in turn, most closely allied with Rajella fyllae . The average of the pairwise differences between Rajella sp. and R. caudospinosa was 33, and between the specimens of Rajella sp. and those of R. fyllae 33.3. We believe these specimens represent a western Atlantic relative of R. caudospinosa .

Rajella caudaspinosa (munchkin skate) ( fig. 74)

Two specimens of this southern African endemic, all collected from South Africa, were included. These specimens were identical in sequence. They clustered most closely with Rajella sp. As noted above, the average of the pairwise differences between these two species was 33.

Rajella leopardus (leopard skate) ( fig. 74)

Five specimens of this southern African endemic, all collected from South Africa, were included in the analysis. The range in pairwise differences among specimens in this cluster was 0–3, with an average of 1.2.

Rajella kukujevi (mid-Atlantic skate) ( fig. 74)

The four specimens of this species included in the analysis, all collected from the North Atlantic, comprised a single cluster. The range of pairwise differences among these specimens was 0–3, with an average of 1.5. These specimens grouped most closely with those of Rajella leopardus . The average of the pairwise differences between these two species was 49.

Dipturus linteus (sailskate) ( fig. 74)

A total of five specimens identified as this species, all collected from the eastern North Atlantic and off Norway, and thus representing the eastern portions of the distribution of this species, were included in the analysis. The range in pairwise differences among these specimens was 0–1. They comprised a single cluster, which grouped most closely with the clusters of the three Rajella species. This Dipturus species clustered well away from all 15 of its congeners included in the analysis (see fig. 69), lending support to Compagno (2005b) who listed this species as ‘‘ Dipturus ? linteus’’ and noted that its generic placement needs investigating.

Amblyraja radiata (thorny skate) ( fig. 74)

In total, the analysis included 14 specimens identified as this species, one of which is in the Harvard Museum of Comparative Zoology (GN2602 5 MCZ 159184). These were found to comprise a single cluster. Although there was evidence of structure within this cluster, and the range of pairwise differences among the members of this cluster was 0–15, with an average of 8.5, there was no obvious pattern to this variation; all specimens within this cluster have thus been given the same designation. These specimens represent the more eastern part of the distribution of this species, which occurs throughout the North Atlantic.

Amblyraja hyperborea (Arctic skate), Amblyraja

jenseni (Jensen’s skate), Amblyraja badia (broad

skate), and Amblyraja sp. complex ( fig. 74)

The analysis yielded a somewhat confusing cluster of nine specimens, five of which are deposited in museums. This cluster consists of four specimens from the Barents Sea identified as A. hyperborea by Age Hoines, one specimen from California identified as A. badia deposited at the University of Washington (GN6681 5 UW 115021), one specimen from the mid- Atlantic identified as A. jenseni by Chip Cotton, and deposited at Virginia Institute of Marine Sciences (GN5040 5 VIMS 11757), as well as two specimens deposited at the Harvard Museum of Comparative Zoology (MCZ GN4666 5 MCZ 167944 and GN4667 5 MCZ 167945), both identified only as Amblyraja sp. , and a specimen from the Tasman Sea, identified as Amblyraja hyperborea , from the Australian National Fish Collection (GN4650 5 ANFC H 5944-01). These specimens differ somewhat in morphology. The range of pairwise differences among the nine specimens comprising this cluster was 0–11, with an average of 5.7. As none of the specimens comprising this cluster have been definitively identified, we have retained the original identifications as they serve to draw attention to the uncertainty associated with the identity of the members of this cluster.

Amblyraja doellojuradoi (southern thorny skate)

( fig. 74)

Three specimens of this species, all collected from the Falkland Islands, were included in the analysis. They represent the more eastern parts of the distribution of this species, which occurs throughout the southern regions of South America. The range of pairwise differences among specimens was 4–8, with an average of 5.3. These specimens grouped most closely with the A. hyperborea , etc., cluster. The average of the pairwise differences between the specimens of these two clusters was 18.7.

Leucoraja naevus (cuckoo skate) ( fig. 75)

In total, 29 specimens of this species were included in the analysis. These were all collected from localities in the eastern North Atlantic and thus represent the northern portions of the distribution of this species, which has been reported from as far south as Senegal. The range of pairwise differences among the specimens in this species was 0–6, with an average of pairwise differences of 1.5.

Leucoraja wallacei (yellowspot skate) ( fig. 75)

The 12 specimens of this southern African species, all collected from South Africa, were found to comprise a single cluster. The range in pairwise differences among specimens was 0–8, with an average of 3.7. This species grouped most closely with the eastern North Atlantic species Leucoraja naevus ; the average of the pairwise differences between these two species was 37.5.

Leucoraja fullonica (shagreen skate) ( fig. 75)

The analysis included two specimens collected from the Azores and two from the eastern North Atlantic. These specimens thus represent some of the southern and western elements of this species, which occurs as far eastward as Murmansk, Russia. The range in pairwise differences among these specimens was 0–3, with an average of 2. The analysis yielded a single cluster, which grouped most closely with the cluster consisting of L. naevus and L. wallacei . The average of the pairwise differences among specimens of L. fullonica and L. naevus was 72.4, and L. wallacei was 65.4.

Leucoraja erinacea (little skate) ( fig. 75)

In total, four specimens of this western Atlantic species were included in the analysis, which yielded a single cluster with a range of pairwise differences among specimens of 3–8 and an average of pairwise differences of 5.3. One of these specimens is in the University of Kansas Ichthyology Collection (GN2505 5 KUI 26967).

Leucoraja ocellata (winter skate) ( fig. 75)

The two specimens of this western Atlantic species included differed by two bases. They grouped most closely with the specimens of L. erinacea ; the average of the pairwise differences between specimens of these two species was 45.5.

Leucoraja garmani (rosette skate) ( fig. 75)

The single specimen of this species, collected from the western North Atlantic, represents a more northern element of the distribution of this species, which occurs as far south as Venezuela. It grouped outside all four other species of this genus included here. The averages of the pairwise differences between this specimen and its congeners were: L. naevus 130.3, L. wallacei 130.9, L. fullonica 132.5, L. erinacea 142.8, and L. ocellata 143.

Rostroraja alba (white skate) ( fig. 75)

A total of six specimens of this species was included in the analysis. These specimens were all collected from South Africa and thus represent only a central element of the distribution of this species, which has been reported from Great Britain, the Mediterranean Sea, much of the coast of Africa, and the Red Sea. These specimens were identical in sequence. This species grouped most closely with the specimens identified as Raja velezi . The average of the pairwise differences between Ro. alba and Ra. velezi was 86. This grouping is generally consistent with the affinities reported by McEachran and Dunn (1998) and Compagno (2005b) who listed these species, among others, as comprising an ‘‘Amphi-American Assemblage.’’ However, these results suggest that some consideration should be given to Rostroraja as the generic name to apply to the species comprising this entire assemblage.

Raja velezi (rasptail skate) ( fig. 75)

All three specimens of this species were identical in sequence. All three were collected from the Gulf of California and thus represent a northern distribution of this species, which has been reported from as far south as Peru. These specimens grouped most closely with those of Ro. alba .

Raja eglanteria (clearnose skate) ( fig. 75)

Four specimens, identified as R. eglanteria , collected from the western North Atlantic and the Gulf of Mexico off Florida, were included in the analysis. These represent much of the distribution of this species, which occurs from Massachusetts to Florida. The range in pairwise differences among specimens was 0–12, with an average of 8.2. This species grouped with Ra. velezi and Rostroraja alba , well outside specimens of the other Raja species included in the analysis (see figs. 71, 73), which is consistent with McEachran and Dunn (1998) and Compagno (2005b) who listed Ra. eglanteria and Ra. velezi , among other species, as comprising an ‘‘Amphi-American Assemblage’’ that may represent an undescribed genus.

Okamejei jensenae (Philippine ocellate skate)

( fig. 75)

The analysis included two specimens of the recently described Okamejei jensenae (see Last and Lim, 2010) from the Philippines, which were treated by Compagno et al. (2005b) (GN4343 5 BRU 171 and GN4382 5 JPAG 328) as Okamejei sp. 1 . These differed from one another by one base. It is of note that this species grouped most closely with specimens in the R. eglanteria and R. velezi clusters, well away from its congeners (see fig. 70). The generic placement of this species requires additional taxonomic investigation when adult males of this species are collected as clasper cartilages are needed to assign this taxon to a genus.

Malacoraja senta (smooth skate) ( fig. 75)

All three specimens of this species were found to be identical in sequence. These specimens were collected from the western North Atlantic and are largely representative of the distribution of this species, which occurs from Newfoundland to New Jersey.

Neoraja caerulea (blue pygmy skate) ( fig. 75)

Two specimens of this eastern North Atlantic endemic were included in the analysis; they were identical in sequence. They clustered most closely with the specimens of M. senta and the average of the pairwise differences among specimens of these two species was 80.