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Volume 87, 1959
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Centrophorus plunketi Waite, 1910 is referred to Scymnodon. The lower teeth are centroscymnoid (oblique and without a symmetrical median tooth) but the upper teeth are scymnodoid (longer midway out along either side of jaw than at centre of mouth) as are the dermal denticles. Juvenile dermal denticles of S. plunketi have acutely trifid, concave, ridgeless blades; intermediates, concave with a shallow median longitudinal ridge; adult denticles, obtuse, trifid and strongly ridged. Centroscymnus waitei (Thompson, 1930) known only as juveniles, is recognised as S. plunketi. S. plunketi is abundant in New Zealand waters beyond 300 fathoms, and appears to be a bottom-dwelling, schooling species. Its liver has a high oil content (84 g. oil/100 g. fresh liver) but is low in Vitamin A (370 I.U./g. oil).

The deep-water spiny dogfish described from New Zealand waters by Waite (1910) as Centrophorus plunketi has been variously placed in Centrophorus, Scymnodon and Centroscymnus by most later authors; while Fowler (1933, p. 239) proposes for it a new subgenus Proscymnodon which Whitley 1934, p. 199) regarded as a genus. These divergent opinions reflect the need for further examination of this species, which is now known to be abundant in New Zealand beyond 300 fathoms.

Consideration of recently collected adult material necessitates the transfer of the Plunket shark to the g. Scymnodon, and examination of juveniles and a half-grown specimen which were originally regarded as Centroscymnus waitei (Thompson, 1930) shows that Thompson's species is a synonym of S. plunketi. It is now definite that juvenile and adult dermal denticles of S. plunketi differ strongly in form. Denticle changes of this kind have not previously been reported in Scymnodon.

In their recent account of the Squaloidea, Bigelow and Schroeder (1957, p. 25) restate the generic criteria of Centrophorus, Centroscymnus and Scymnodon (the last including the Proscymnodon of Whitley which does not now warrant generic distinction), using as primary characteristics the shape of the posterior (inner) corner of the pectoral fin, and the size and shape of the teeth. Centrophorus is immediately distinct from Scymnodon and Centroscymnus in that the posterior pectoral corner is more or less extended (exceptionally it is angular in the juveniles of C. squamosus). The diagnosis for Scymnodon is that the posterior pectoral corner is rounded; the upper teeth midway along either side of jaw noticeably longer than those toward centre of the mouth; and the lower teeth erect and symmetrical along the median sector of the mouth, only weakly oblique towards the angles, and with a symmetrical median tooth. In Centroscymnus the posterior pectoral corner is rounded; the upper teeth midway along either side of the jaw scarcely longer than those at the centre of the mouth; and the lower teeth oblique and without a symmetrical median tooth.

The Plunket shark has a rounded posterior pectoral corner, and upper teeth like those of other species of Scymnodon; but its lower teeth are markedly oblique

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along the whole jaw, and there is no symmetrical median tooth (Text-fig. 2). In these lower teeth characters it corresponds with the condition in Centroscymnus. However, as has already been shown by Garrick (in press) the lower teeth are not always reliable criteria even in Centroscymnus, and where there is confusion from reference to the upper and lower teeth this can be resolved in all except the most juvenile specimens by examination of the dermal denticles. On the Plunket shark these are ovoid, tridentate, and have a median longitudinal ridge extending along the entire length of the blade (Text fig. 3), leaving no doubt that the species is referable to Scymnodon. Such denticles contrast with those of Centroscymnus which are more circular in outline, have an uninterrupted circular concavity at the anterior end, and when a median longitudinal ridge is present, it is restricted to the posterior half or two-thirds of the blade, behind the concavity.

Although the obliqueness of the lower teeth of S. plunketi is without generic significance, it does set this species clearly apart from the three other species of Scymnodon currently recognised. In this respect it is as unusual a feature as the transverse ridging which occurs in addition to the usual longitudinal ridging on the dermal denticles of S. obscurus (Vaillant, 1888), but is not found on those of S. ringens Bocage and Capello, 1864, or S. squamulosus (Gunther, 1877), (or, for that matter, S. plunketi).

Compared with S. ringens (figured in Bigelow & Schroeder, 1957, Fig. 13, A-E) and S. squamulosus (in Gunther, 1887, Pl. 2, Fig B), S. plunketi more resembles S. ringens in its overall heavy-bodied form and short snout. However, it differs from S. ringens in having a well-marked concavity in the lower posterior margin of the caudal fin; the posterior tip of the second dorsal fin reaching only as far back as the level of the hypural origin; shorter gill-openings; and in a much greater interspace between the tip of pectoral fin, when laid back, and the level of origin of the 1st dorsal spine (almost equal to the interspace between the eye and the first gill-opening in adult specimens, though much shorter than this in juveniles).

Differences between S. plunketi and S. squamulosus include a much blunter and shorter snout in the former (the preoral length noticeably less than the width of the mouth in S. plunketi, but much more than this width in S. squamulosus); and much larger first dorsal and pectoral fins in S. plunketi which by comparison make those of S. squamulosus appear markedly slender.

It was suggested by Thompson (1930, p. 278) that the species he described as Centrophorus waitei from a single juvenile specimen taken in deep water off Kaikoura might prove to be a growth stage of S. plunketi. This view has not been accepted completely by later authors mainly because the dermal denticles are markedly different Waite (1914) had previously described late embryos of S. plunketi, which Thompson does not mention having available for comparison, and they cannot now be found in the Canterbury Museum. Direct comparison of Thompson's specimen with Waite's identified embryos of S. plunketi is therefore not possible, and no other embryos or juveniles of the latter species have been described. As a consequence Thompson's species has been retained, though Whitley (1940, p. 143) referred it to Proscymnodon, but later (1956, p. 398), following Garrick (1955) to Centroscymnus. Bigelow and Schroeder (1948, p. 451, footnote 8) considered it to be a Scymnodon, but later (1957, pp. 88–96) also transfer it to Centroscymnus and reaffirm Thompson's suggestion that it may be a young stage of S. plunketi. Garrick (1955) gave a full description and illustration of the type of C. waitei, and, chiefly on the nature of the dermal denticles which have strongly concave blades and lack longitudinal ridges, regarded it as a Centroscymnus.

Since Garrick's (1955) study, further juvenile specimens seemingly C. waitei, have become available. These include two male embryos in the Dominion Museum collections, 336 and 348 mm long (Dom. Mus. No. 763) labelled “C. plunketi”,

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from Cook Strait, 1929; and one female 523 mm long (Dom. Mus. No. 2637) long-lined from 500 fathoms off Kaikoura by Mr. R. Baxter in November, 1955. These all agree closely with the type of C. waitei except that their caudal fins have a more or less truncate terminal lobe and a definite subterminal notch (Text-fig. 1, A). In this respect they resemble other species of Centroscymnus and Scymnodon, and hence the unusually pointed caudal fin of the type of C. waitei can no longer be regarded as a distinctive feature of this species. The proportional dimensions of the 523 mm specimen are more or less intermediate between those of the type of C. waitei (318 mm) and those of two adult specimens of C. plunketi (1197 mm and 1417 mm), as in Table I, so that no specific distinction is apparent. Such variation as is present can be recognised as change with growth, the adults having proportionately longer bodies, and shorter heads and tails relative to the juveniles. Equivalent variation has been reported for species of Centrophorus and Centroscymnus (Garrick, 1959; in press).

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Table I.—Proportional Dimensions in per cent. of Total Length.
Scymnodon plunketi Centroscymnus waitei
1,417 mm N.Z. ♀ 1,197 mm N.Z. ♂ 523 mm N.Z. ♀ 318 mm Type ♂
Trunk at pectoral origin:
Breadth 14.4 13.3 12.2 12.9
Height 11.3 10.1 9.7 8.2
Snout length in front of:
Outer nostrils 1.1 0.8 0.8 1.6
Mouth 5.4 5.0 7.5 6.9
Eye: horizontal diameter 3.9 3.7 5.2 5.0
Mouth: breadth 6.7 6.5 6.7 8.2
Nostrils: breadth between inner corners 3.0 2.9 3.8 3.5
Preoral clefts: breadth between inner corners 5.1 4.2 5.0 4.7
Gill-opening lengths:
1st 1.6 1.8 1.5 1.6
5th 1.4 1.6 1.3 1.3
1st dorsal fin:
Vertical height 3.6 3.2 3.1 2.5
Length of base measured from origin of spine 4.8 4.1 4.2 4.1
2nd dorsal fin:
Vertical height 4.3 4.0 4.2 3.5
Length of base measured from origin of spine 5.6 5.2 5.3 5.3
Caudal fin:
Upper margin 17.2 18.4 23.8 23.8
Lower ant. margin 11.1 10.4 14.6 11.6
Pectoral fin:
Ant. margin 13.4 11.8 15.0 12.9
Distal margin 7.6 7.1 6.0
Pelvic fin: ant. margin 8.2 6.6 6.3 5.0
Distance from snout to:
Eye 3.5 3.2 4.2 4.1
1st gill opening 14.8 13.1 15.9 16.0
5th gill opening 17.8 16.7 18.9 19.8
1st dorsal spine 36.8 36.7 37.0 34.3
2nd dorsal spine 68.6 67.8 62.2 60.7
Upper caudal 82.0 81.5 76.5 75.0
Pectoral 17.9 16.8 19.0 20.1
Pelvic 60.7 60.0 54.5 52.0
Interspace between:
2nd dorsal and caudal 8.0 8.5 9.6 8.5
Pelvic and subcaudal 10.6 12.2 10.1 13.2
Distance from origin to origin:
Pectorals and pelvic 43.3 42.8 36.8 31.7
Pelvic and subcaudal 17.6 19.6 15.8 18.8
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The distinction between the dermal denticles of the juvenile C. waitei and those of adult S. plunketi is very misleading. The usual practice of examining denticles from the anterior half of the trunk when applied to the 523 mm specimen is equally misleading, for the majority of the denticles in this region are as in the type of C. waitei—i.e., they are well spaced, with strongly tridentate, concave, ridgeless blades rising steeply from the skin. But examination of the posterior half of the trunk shows in addition slightly larger denticles of another form which are newly erupted,* and have a shallow median longitudinal ridge extending along the whole length of the blade (Text fig. 1, B). The presence of this complete median ridge means that the denticles cannot be those of a Centroscymnus but instead are referable to Scymnodon. Moreover, their shape and sculpture is such that they could reasonably be regarded as intermediate to the final form of the ovoid, tridentate, ridged blades of S. plunketi (Text-fig. 1, C-E).

The above leaves no grounds on which to maintain C. waitei as separate from S. plunketi. If further confirmation is required, it is provided by the teeth. The upper teeth of C. waitei are like those of other species of Scymnodon, including S. plunketi, in that they are longest midway out along each side of the jaw; but the lower teeth series share with S. plunketi the peculiar feature of being oblique at the centre of the mouth and lacking a symmetrical median tooth.

The dermal denticle changes noted above (Text-fig. 1, C-E) are the first to be recorded from a species of Scymnodon, although equally remarkable changes are known from some species of Centrophorus and Centroscymnus. However the changes in S. plunketi are novel in that no other cases are known where ridgeless juvenile denticles are replaced by ridged intermediate and adult denticles. In Centroscymnus coelolepis and C. owstonn the reverse occurs, strongly ridged juvenile and intermediate denticles being replaced by adult denticles which lack ridges or at least have them greatly reduced.

All the known New Zealand catches of S. plunketi are from Cook Strait, Karkoura and Banks Peninsula, but it is likely that line-fishing in deep-water elsewhere on our coasts would show that the species is not restricted to these areas. The depth range so far known is 120–780 fathoms and probably approximates the full range of the species, for it is rarely taken by commercial line-fishermen in 150 fathoms and shallower, and has not yet been caught on lines set deeper than 1,000 fathoms in Cook Strait. The south-eastern Australian records of the species (the only other region where it is known) give it an even narrower range of 260–450 fathoms, as shown by Cowper & Downie (1957, Fig. 12) who record 17 specimens from 7 stations, the greatest single catch of 7 specimens being long-lined in 360–400 fathoms. An indication of its abundance in 300–400 fathoms in New Zealand is shown by the catches of a commercial line-fisherman, Mr. R. Baxter, of Oaro, Kaikoura, who within a month caught sufficient specimens to yield 3,300lbs of liver. The livers average 81b to 101b per shark, so that some 300 to 400 specimens were taken within the period.

It now seems highly probable that S. plunketi is a schooling species of the deeper waters, at least on occasions, the schools segregating not only according to size but also to sex. This is shown by New Zealand catches of December, 1955, when 13 males, 1090 mm to 1198 mm long were taken in 450 fathoms, and July 1957, when 5 females 1,290 mm to 1,320 mm were caught in 600 fathoms. The Australian records also indicate that it is a bottom-dwelling species, for although it was taken there on seven occasions by the use of conventional long-lines, it was not caught at all during an extensive drop-lining programme in the same depths and region.

[Footnote] * This agrees with what has been described in Centroscymnus owstonii, where it has been shown by Garrick (in press) that replacement denticles which are of a new form, first appear on the caudal peduncle.

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Picture icon

Text-Fig. 1—Fig. A—Centroscymnus waitei, tail of embryo male, 348 mm (Dom. Mus. No. 763). Fig. B—C. waitei, female, 523 mm (Dom. Mus. No. 2637), juvenile and intermediate dermal denticles from the flank of the caudal peduncle. Figs. C-E—Scymnodon plunketi, external views and cross-sections of the blades of juvenile, intermediate and adult denticles respectively (Figs G and D. from 523 mm female, E from 1,417 mm female), the cross-sections through a-b in each case

The New Zealand catches (where information is available) have all been from long-lines, and on the few times when drop-lines have been used in depths of 300 fathoms or more, no S. plunketi have been taken.

To date there has been no commercial exploitation of S. plunketi other than isolated catches for liver such as mentioned above. The livers have a high oil content, as shown by the analysis of a New Zealand specimen taken in December, 1956, which had 84 g. oil/100 g. fresh liver; but the vitamin A content is low, being only 370 I.U./g. oil, compared with the common figure of 30,000 I.U./g. for the School shark, Galeorhinus australis. Similar figures are given by Cowper & Downie (1957, Table VI) for an Australian specimen of S. plunketi

Cephalopod beaks and bony fish remains are the only stomach contents found in New Zealand specimens of these sharks. Of 18 specimens dissected, all of them long-lined, 12 had empty stomachs, presumably having vomited their contents. Four contained fish remains only; 1, cephalopod beaks only; and 1, both fish remains

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and cephalopod beaks. Five of the same 18 specimens were infested with the large trematode Otodistomum plunketi Fyfe, 1953; mostly only 2 or 3 of these flukes were present, though one large female contained 29. The majority of the sharks also had larval tapeworms in the coelom, and a monogenetic trematode on the skin, the latter usually very numerous.

Little is known of the reproductive cycle or developmental stages other than the information in Waite (1914) and Phillipps (1946) on two gravid specimens. They show that the eggs are very large and numerous, up to 36 embryos being produced in one litter. In Waite's specimen, the 36 embryos were 165 mm long and attached to globular yolk-sacs 66 mm in diameter. Phillipps' specimen contained at least two embryos (Dom. Mus. No. 763) which are 336 mm and 348 mm long and evidently not far off birth, for their yolk sacs are almost completely resorbed. Thompson's (1930) specimen (as C. waitei) is the smallest free living one that has been recorded, and is 318 mm long; it was taken in August, 1928. The smallest mature male known is about 1,000 mm long. No mature female less than 1,290 mm is so far known; this and three others up to 1,320 mm taken in July, 1957, all contained numerous ova 7 mm to 20 mm diameter in each ovary. A 1,630 mm female caught in November, 1956, contained numerous ova of 5 mm diameter in each ovary (ovary length 230 mm), while a 1,700 mm female taken at the same time had only 8 ova of 20 mm diameter in each ovary (ovary length 100 mm). The small size of the ova in both of the above groups of females when compared with Waite's and Phillipps' records of gravid females taken in July and December respectively, and Thompson's recently-born juvenile taken in August, suggests that the breeding season is not well defined, or that S. plunketi breeds only every second year.

Scymnodon plunketi (Waite, 1910).

Centrophorus waitei Thompson, 1930.