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Volume 88, 1960-61
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Studies on New Zealand Elasmobranchii.—Part X
The Genus Echinorhinus, with an Account of a Second Species,
E. cookei Pietschmann, 1928, from New Zealand Waters*

[Received by Editor, September 14, 1959.]

Abstract

Echinorhinus cookei Pietschmann, 1928, is recorded from a 1980 mm male here designated as neotype E. cookei is uniformly covered with numereous small, bucklerlike dermal denticles, up to 4 mm diam. with strongly indented, angular bases and coarsely ridged spines; in contrast the denticles of E. brucus are sparse, irregularly distributed, up to 15 mm diam if solitary but 35 mm if compound, and have entiremargined bases and finely ridged spines. Three New Zealand juveniles of E. cookei are markedly slender, with one-cusped teeth as in Squalus. The lateral line of E. cookei is an open furrow supported by incomplete, transverse skeletal rings with their free ends projecting as spines; the adult furrow is bridged at irregular intervals by skin. A Californian record of E. brucus is shown to be E. cookei. Previous New Zealand records of E. brucus cannot be confirmed, but E. brucus is known for New Zealand by a mounted skin of a Dunedin specimen in the Otago Museum.

The genus Echinorhinus has been known from New Zealand waters since 1884, with five specimens recorded in the literature as E. spinosus Blainville, 1825, E. mccoyi Whitley, 1931 or E. brucus (Bonnaterre, 1788). Of these E. spinosus and E. mccoyi are currently recognised as synonyms of E. brucus, the cosmopolitan Bramble Shark. Despite these records, only fragmentary Echinorhinus material was held in New Zealand museums, the most complete exhibit being a mounted skin of a specimen about 1,420 mm long in the Otago Museum and labelled “Dunedin. April, 1887”. This specimen is not in the literature and there is no further information on it.

In view of the above it was of considerable interest when two Cook Strait fishermen, Messrs. A. Dellabarca and W. Hickman, of the line-boat “Calabria” brought in a Bramble Shark taken in 40–50 fathoms in Palliser Bay on April 20, 1959. This shark, a male, 1,980 mm long, was obviously referable to Echinorhinus but lacked the large, spine-bearing bucklers, up to 15 mm diameter or more on a specimen of equal size, which, irregularly and rather sparsely distributed over the body, are characteristic of E. brucus. Instead the Palliser Bay shark had a uniform covering of small bucklers, not larger than 4 mm diam. and mostly less, and much more numerous than those of E. brucus. Examination of the literature shows that this shark is identifiable as E. cookei Pietschmann, 1928, a species recorded only from the type taken off Hawaii.

The status of E. cookei has not been clear-cut, apparently because Pietschmann (1928, 1930) did not give his reasons for separating it from E. brucus. Also, Fowler (1941, p. 287) who examined the type of E. cookei, reports that he “cannot find that Echinorhinus cookei is other than a variant of this species” (E. brucus). Bigelow & Schroeder (1948, p. 527) leave the question open, stating that “Final conclusions must await critical comparison of adequate series of specimens”.

[Footnote] * This study has been assisted by a grant from the Research Grants Committee of the University of New Zealand.

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The distinction of E. cookei from E. brucus is definite and striking, at least between sub-adult or adult specimens, but so far can be based only on the bucklerlike dermal denticles (Plate 7, Figs. A-D). There appear to be no significant differences in proportional dimensions,* in external morphology, or in details of the teeth. However, the differences in the denticles lie not only in their relative size as mentioned above, though this alone is sufficient, but also in the shape and sculpture of the denticle bases (entire-margined or nearly so, and with rather fine radial ridging in E. brucus, but with strongly indented margins and coarse ridges (Text-fig. 2, E-H) in E. cookei), the presence in E. brucus of compound denticles up to 35 mm long, as a result of fusion of adjacent denticle bases, while such fused denticles are not a feature of E. cookei; and there is the tendency in E. brucus to retain noticeable-sized denticles on the underside of the snout and around the mouth even on large specimens, while in the type of E. cookei and in my specimen this area is virtually smooth, only a few minute denticles being present. For information on the last-mentioned feature in E. brucus I. am indebted to Dr. Denys E. Tucker, who examined five Mediterranean, Eastern North Atlantic and South African specimens, 910 mm to about 2,150 mm long, in the British Museum.

Comparison of my specimen with Pietschmann's (1930, p. 3) account of the type of E. cookei, a male 2,033 mm long and hence comparable to mine, shows close correspondence between them. Agreement in the dermal denticles is confirmed by enlargements from the negatives of the photographs used by Pietschmann in his Plate 1, A. and B; for these enlargements I. am indebted to Dr. Edwin H. Bryan, jun, Bernice P. Bishop Museum. With respect to the proportional dimensions of the type, it should be noted that Pietschmann gives the length of caudal as 5.2 in total length, while in my specimen the caudal is 3.8 if measured from subcaudal origin, and 4.2 if measured from upper caudal. Similarly other given dimensions involving the caudal indicate a shorter tail in the type than in my specimen, though this is not supported by reference to Pietschmann's photographs of the type. It would seem that Pietschmann either used a method of measurement different from that currently employed, or that he failed to make allowance for the undue upturning of the tail of his specimen as shown in his photograph. Some other of Pietschmann's dimensions also call for comment. His distance “from base of pectoral to tip of snout” as 5.2 in head is obviously in error, while “distance of nostrils” as 2.4 in head, and “depth of mouth (distance of middle of rear border from a line connecting both angles of mouth) 4.1 in breadth of mouth” do not match with his very good photograph of the underside of the head where the same dimensions appear to be about 5.0 and 2.8 respectively.

In my specimen these dimensions are 4.0 and 2.3 Unfortunately the type of E. cookei no longer exists, so that no check can be made on its dimensions, but from Pietschmann's account it can be said that other than in these apparently erroneous differences the type and my specimen agree closely in all respects.

The presence of E. cookei in regions as far separated as the tropical North Pacific (Hawaii) and the temperate South Pacific (New Zealand) suggests that it is a wide-ranging species likely to be encountered throughout the temperate and tropical Pacific as a whole, if not outside this ocean as well. This is supported by the recent capture of a 2,126 mm specimen off Peru (information kindly supplied in advance of publication by Miss N. Chirichigno), and also by the record of a Bramble Shark, 1,960 mm long, taken in 50–55 fathoms off California and reported by Hubbs & Clark (1945) as E. brucus, though it is in fact E. cookei. The photographs of

[Footnote] * E. obesus Smith, 1849, long regarded as a synonym of E. brucus, is a much more heavybodied fish than E. cookei (or for that matter E. brucus also) if Smith's figure (Pl. 1) is accurate. It is shown as having large, sparsely distributed bucklers like those of E. brucus.

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Figs A-B Echinorhinus cookei Pietschmann, neotype (Dom Mus No. 2744) male, 1,980 mm, New Zealand Fig A, Trunk between pectoral and pelvic fins showing small dermal denticles White areas on trunk are due to damage during capture Fig B, Closeup of denticles from above lateral line on trunk
Photos F. O'Leary, Dominion Museusm
Figs C-D Echinorhinus brucus (Bonnaterre), 1,420 mm long mounted skin in Otago Museum, New Zealand Fig C, Trunk between pectoral and pelvic fins showing large bucklers Note buckler spines projecting from profile of trunk Fig D, Close-up of denticles, mostly below lateral line, showing compound bucklers at left of photogroph
Photos. R. R. Forster, Otago Museum
N. B. The dried skin of Figs. C-D makes the denticles shape and size more obvious than in Figs. A-B, where the specimen is formalin preserved

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the Californian specimen in Hubbs & Clark's account indicate a lack of the large bucklers characteristic of E. brucus. This was confirmed through the kindness of Dr. Ernest A. Lachner, who examined the Californian specimen (deposited in the United States National Museum, Cat. No. 130667), and supplied a small representative sample of skin from high on the flank. The sample shows complete agreement in the dermal denticles with my specimen, the bucklers being mostly about 2 mm to 3 mm diam. and having strongly indented bases and prominent radial ridges. Dr. Lachner also reports that there are tiny scattered denticles on the underside of the head and snout, so that these regions are not entirely smooth (though more nearly so than in E. brucus) which is likewise in accord with my specimen. The dimensions of the Californian specimen as given in Hubbs and Clark's account parallel mine except for longer gill-openings, and the preocular length greater than the preoral—in my specimen the anterior margin of the eye is just in front of the level of the mouth. But I. do not regard these differences as warranting specific distinction.

The above-mentioned confusion of E. cookei with E. brucus poses the question of how many other reports of Bramble sharks as E. brucus (or its synonyms E. spinosus and E. mccoyi) are based on E. cookei. The diagnosis of E. brucus in many accounts is not sufficient to distinguish E. cookei. As an example, although Gunther (1870, p. 428) in his generic diagnosis of Echinorhinus gives “skin with scattered large round tubercles”, his diagnosis of E. spinosus is “each tubercle with a small spine in the centre”; the use of the word “small” in this context is likely to cause misinterpretation as to the size of the bucklers as a whole. The same can be said of Jordan & Evermann's (1896) account, where identical wording is employed. The question is unlikely to be answered for many, perhaps most records, unless they are fully descriptive of the denticles, or substantiated by illustrations, specimens, or fragments of specimens showing representative samples of denticles. Consequent on this, of the five New Zealand records of E. brucus, none can with certainty be confirmed as this species. These records are summarised below.

1884. Parker, T. J. (Trans. N. Z. Inst., 16: 280–281). Specimen caught off Dunedin, July, 1883, identified as E. spinosus from mutilated remains (jaws and tail). No description or illustration. (The jaws and tail cannot now be found in the Otago Museum where they were originally deposited.)

1886. Hamilton, A. (Trans. N. Z. Inst., 18: 135). Records, in footnote, specimen of E. spinosus taken at Napier, Hawke's Bay, in September, 1885.

1913. Waite, E. R. (Rec. Cant. Mus., II (1): 17) Identifies from photograph as E. brucus, a 2,567 mm specimen washed ashore at Opotiki, Bay of Plenty. No description or illustration.

1928. Phillipps, W. J. (N.Z. Journ. Sci. & Tech. X. (4): 221–222). 1,575 mm specimen recorded from Cook Strait as E. brucus Describes colour as rich brown with ovate black spots; denticles as circular bony tubercles with a sharp spine in centre.

1946. Phillipps, W. J. (Dom. Mus. Rec. in Zool., I. (2): 19–20). Reports jaws of specimen recently trawled off Hawke's Bay as E. mccoyi.

The black spots on Phillipps' (1928) specimen suggest E. brucus rather than E. cookei which is so far known only as uniformly coloured (some specimens of E. brucus are likewise reputed to lack such spots), while the description of the denticles as “circular bony tubercles” would also point to E. brucus. But these can scarcely be classed as definitive criteria, and in the absence of supporting specimens the presence of E. brucus in New Zealand waters seems to depend on the mounted skin (Pl. 7, C–D) in the Otago Museum which, as mentioned above, has not previously been recorded.

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With regard to E. cookei, two occurrences* in New Zealand other than the Palliser Bay specimen described here can be placed on record. The first of these is of a large specimen, reputedly about 4,000 mm (13 feet) long, taken in 35 fathoms off Shag Point, near Mocraki, towards the end of 1953, by Mr. R. Welsh. The photograph I. have of this specimen, which is the only evidence now available, shows clearly a uniform denticulation and absence of large bucklers. The reported length of this specimen appears to be a record for the genus—E. brucus is not known longer than about 3,000 mm—and from the photograph seems substantially correct in comparison with the size of the people surrounding the specimen. The second additional record is based on three juvenile specimens, 445 mm to 472 mm long, held in the collections of the Department of Zoology, Victoria University of Wellington. These juveniles in which the yolk-sac scars are just healed, were trawled in Cook Strait in either 1946 or 1947, and presented to the University by Mr. F. Abernethy. They show the same form of dermal denticles (Text-fig. 3, C) as the adult, but with two sizes of denticles present, of which the smaller are 0.5 mm to 0.8 mm diam and cover the whole body, including the underside of head and snout where they are almost as numerous as elsewhere. The larger denticles are 1.5 mm to 2.0 mm diam. (about half the size of adult E. cookei denticles) and are newly erupted or erupting; these are present mainly on the upper half of the trunk, above the lateral line; their distribution and spacing agree well with the adult. Despite these similarities the identification of the juveniles a E. cookei is made with some degree of caution, for the smallest denticles on the adult E. brucus in the Otago Museum have the same facies as those of my juveniles and hence also of adult E. cookei—i e, they have strongly indented, angular bases and coarse radial ridges (pers. comm. from Dr. R. R. Forster, Director, Otago Museum). Although the denticulation of juvenile E. brucus is not known, it can be inferred from Dr. Forster's observation that the small denticles on his adult are of juvenile from; hence such juvenile denticles will be common to both species, but will mostly be replaced by larger, circular-based denticles in sub-adult E. brucus. On these grounds the identification of my juveniles is necessarily tentative, except that the number, spacing and distribution of their larger denticles falls so closely into line with that of adult E. cookei, without suggesting the sparse and irregular distribution of E. brucus Further support for the belief that the juveniles are E. cookei is provided by Dr. Denys E. Tucker's examination of five specimens of E. brucus in the British Museum He reports (pers. comm). “as we proceed to larger specimens, it becomes evident that the definitive denticulation must be laid down at a relatively early age” His data give the size of large bucklers on a 910 mm male as up to 17 mm diam., while on a 1,690 mm male “the large denticles show no increase in absolute size and are rather more widely spaced”. This means that if my juveniles are E. brucus rather than E. cookei they would, in doubling their length (to reach the length of Tucker's smallest specimen) have to acquire a full array of adult denticles with diameters up to eight times that of their present 2.0 mm denticles. This seems unlikely.

The slenderness of the juveniles compared with the adult E. cookei is striking, especially along the posterior half of the trunk and the peduncle (Text-fig. 3A) where the depth of the body is relatively only two-thirds that of the adult. In other regions, including the head and the tail, the difference is less marked but still obvious. Relative longitudinal proportions are more nearly alike although the juveniles have rather longer heads and tails, as would be expected.

[Footnote] * Since the preparation of this account, another specimen (female, Dom. Mus. No. 2826) comparable in size to the Palliser Bay specimen, has been taken in western Cook strait. It was lined by W. T. Mc Manaway on October 10, 1959.

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Differences between the juvenile and adult teeth are also remarkable, though apparently paralleled in E. brucus. The juvenile teeth, both upper and lower, are one-cusped (Text-fig. 3, D-E), the cusps strongly reflexed laterally so as to point almost horizontally. They are closely similar to those of Squalus in appearance but rather wider in relation to their height. The adult teeth are more erect, and have four or five cusps, there being one or two minor cusps on each side of the major cusp (Text-fig. 2, I-J). Behind the upper teeth, inside the mouth, is a prominent maxillary velum, fimbriated along its posterior margin, and with a narrow band of many fleshy papillae immediately internal to the teeth (Text-fig. 2, C). It resembles that of many batoids and is an unusual feature for sharks.

The lateral line of juveniles and adult is open for most of its length, as it is in E. brucus, forming a conspicuous white furrow from above the second gill-opening rearwards. In cross-section the upper edge of the furrow slightly overhangs the lower edge, especially in the adult. Each edge of the furrow is armed with a row of pointed spines, their tips directed rearwards; this also agrees with the condition described for E. brucus in Bigelow & Schroeder (1957, p. 134) Closer examination shows that the spines are not discrete units but are the free ends of regularly placed, incomplete skeletal rings, lying inside transversely across the furrow and supporting its walls (Text-fig. 1, A-C); presumably these rings, which have not previously been described, are derived from dermal denticles. They are unique not only for the Squaloidea but for all other true sharks as well (excluding here the Chimaerae

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Text-fig. 1.—Fig. A, semidiagrammatic view of lateral line from trunk of 445 mm juvenile Echinorhinus cookei (Dom. Mus. No. 2792) showing ring-like skeletal elements (SR) with pointed tips bordering the edges of the open furrow Fig. B, skeletal elements of lateral line from same specimen and position as Fig. A, but drawn from dried skin Fig. C, semidiagrammatic view of lateral line from trunk of 1,980 mm Echinorhinus cookei (neotype, Dom. Mus. No. 2774) New Zealand, showing similar features as Fig. A, though skeletal rings (SR) may have bifid tips, the furrow is more nearly closed and is bridged at intervals by skin (BR).

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where similar dermal rings are recorded for Chimaera which has an open lateral line). In the juveniles the rings are simple and readily visible; in the adult they frequently have bifid tips and are obscured to some extent by the more nearly approximated edges of the furrow. The adult lateral line differs further from the juvenile in that it is bridged at irregular but fairly close intervals along its length by narrow cross-connections of skin (Text-fig 1, C), still, however, leaving much more of the furrow open than covered. Careful inspection along the whole length of the lateral line suggests that this incomplete bridging is the normal condition. In this respect my interpretation differs from that of Bigelow & Schroeder, who assume that in E. brucus the canal is normally closed, at least as far back as the pelvic fins, but is opened by damage or wear during capture and subsequent handling. Anterior to the second gill-opening the lateral line is completely closed except for a row of pores as is usual in most other sharks and all other squaloids. It is of particular interest that in this forward region the lateral line does not contain skeletal rings. Rings appear to be one kind of functional modification for support of open lateral line canals. In Chlamydoselachus and Notorhynchus, the only other true sharks with open lateral line canals, the edges of the furrow are margined by a row of somewhat overhanging dermal denticles, whose bases, at least in Notorhynchus, and apparently also in Chlamydoselachus (Garman, 1885, p. 7, Pl. 6, Fig. 10) do not extend far into or across the furrow.

Genus Echinorhinus Blainville, 1816

Teeth similar in upper and lower jaws, with 1 cusp in juveniles but 3 to 7 cusps in older specimens, the middle cusp of each tooth much the largest and so strongly oblique that the inner margin of adjacent middle cusps form an almost continuous cutting edge along the jaw; dermal denticles in the form of tubercles or shields, either simple, with central spine, or compound, with two or more spines, distributed uniformly or in groups; caudal without precaudal pits, the axis moderately raised, the fin deeper below the axis than above, its tip pointed without subterminal notch, oral clefts short, restricted to angles of mouth, the preoral cleft not pouched towards midline; lateral line an open furrow along sides of body, its walls supported by incomplete skeletal rings whose free ends project as spines along edges of furrow, bridged at irregular intervals by narrow cross-connections of skin; spiracles minute; origin of first dorsal over bases of pelvics, and far behind midlength of trunk; gill-openings large; (Adapted from Bigelow & Schroeder, 1948, p. 526, 1957, p. 134)

Echinorhinus cookei Pietschman, 1928.

Echinorhinus brucus (not Bonnaterre). Hubbs, C. L. and Clark, F. N., 1945. Calif. Fish and Game 31 (2):64–67.

The type of E. cookei is now lost, having disintegrated following dessication; no fragments are known to have been ratained (pers. comm. from Dr. Edwin H. Bryan, jun., Curator of Collections, Bernice P. Bishop Museum, Hawaii). I. am reluctant to designate a neotype, but in view of the confusion there has been between E. cookei and E. brucus, and the failure of many accounts of E. brucus to provide sufficient detail to exclude E. cookei, a reference specimen is necessary and warranted I thereby designate the 1,980 mm male described below from Cook Strait, New Zealand, as the neotype of E. cookei. The neotype is deposited in the Dominion Museum, Wellington, New Zealand, where it bears the label “Neotype. Echinothinus cookei Pietschmann, male 1,980 mm T.L., from 40–50 fathoms Palliser Bay, April, 1959. Coll. A Dellabarca & W. Hickman. Dom. Mus. No. 2774”

Study Material

(a) Adult: Male, 1,980 mm T. L. (Neotype, Dom. Mus. No. 2774) lined from 40–50 fathoms, Palliser Bay (Cook Strait, New Zealand), Aprial, 1959.

(b) Juveniles: Female, 450 mm T. L., male, 445 mm T. L. (Dom. Mus. No. 2792) and male, 472 mm T. L. trawled from Cook Strait, New Zealand, about 1946 or 1947 and held in the Zoology Department, Victoria University of Wellington.

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Text-fig. 2—Echinorhinus cookei Pietschmann, 1,980 mm male (neotype, Dom. Mus. No. 2774) New Zealand. Fig. A, lateral view and insets of transverse sections of snout and peduncle. Fig. B, ventral view of head. Fig. C, maxillary velum (medial half, left side) showing fleshy papillae behind teeth, and fimbriated posterior margin. Fig. D, right nostril. Fig. E, dermal denticles from high on side, halfway along trunk, drawn from dried skin. Fig. F, external view of denticle. Figs. G-H, lateral views of recurved and erect denticles. Figs. I-J, upper and lower teeth, right side. cl, level of cloaca.

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Description

Proportional Dimensions in per cent. of Total Length.
Adult (Neotype Dom. Mus. No. 2774) Juveniles (Dom. Mus. No. 2792)
1,980mm 472mm 445mm
Trunk at pectoral origin: Breadth 13.9 12.5 14.4
Height 13.1 11.2 11.7
Snout length in front of: Outer nostrils 4.8 5.9 6.4
Mouth 7.0 9.1 9.9
Eye: Horizontal diameter 1.8 3.2 3.8
Mouth: Breadth 9.8 12.5 12.6
Height 4.3 6.4 6.2
Nostrils: Distance between inner ends 5.3 5.7 5.8
Preoral clefts: Length 1.1 1.4 1.6
Gill-opening lengths: First 2.9 3.6 4.3
Fifth 5.7 4.9 6.3
First dorsal fin: Vertical height 3.8 3.4 2.8
Length of base 6.2 3.8 3.4
Second dorsal fin: Vertical height 4.1 3.0 3.4
Lenght of base 5.4 4.0 3.6
Caudal fin: Upper margin 23.0 26.9 27.0
Lower anterior margin 12.6 12.9 11.7
Pectoral fin: Length anterior margin 11.7 10.8 11.5
Breadth 7.1 5.5 5.8
Distance from snout to: Eye 6.6 8.5 9.2
1st gill-opening 20.1 22.0 23.0
5th gill-opening 25.5 27.5 28.1
Pectoral origin 26.0 28.4 29.5
1st dorsal origin 57.3 58.5 59.0
2nd dorsal origin 67.2 65.4 66.0
Pelvic origin 55.7 54.0 54.0
Upper caudal origin 76.4 73.0 72.0
Interspace between: 1st dorsal base and 2nd dorsal origin 4.3 3.8 4.0
2nd dorsal base and upper caudal
origin 4.1 2.6 2.2
Pelvic base and subcaudal 7.0 6.5
Distance from origin to origin of: Pectoral and pelivic 29.3 25.4 24.7
Pelvic and subcaudal 18.4 16.0

(a) Adult (Text-fig. 1, C, Text-fig. 2, A–J, Plate 7, A–B)

Head depressed, small-eyed, snout pointed. trunk stout, sub-cylindrical anteriorly, markedly compressed posteriorly. Dorsal and ventral profiles of trunk almost parallel. Height of trunk at origion of pectorals about one-sixth of its length to subcaudal origin. Length of body measured to cloaca 63.3% of total length. Caudal peduncle strongly compressed and deep, its depth almost twice its width and equal to snout length in front of eye. No precaudal pits or lateral keels on the peduncle.

Dermal denticles in the form of erect or slightly retrorse thorns with prominent radial ridges extending down on to multiangled bases. The perimeter of each base is deeply indented between adjacent radial ridges so that the numerous angles and ridges are strongly defined. Each denticle carries only one thorn which is sharply pointed, and ridged along all or most of its length. All denticles are solitary, adjacent bases never fused though occasionally contiguous. Diameter of denticle bases mostly 2–3 mm or less, infrequently 4 mm.

Denticles obvious over whole of body excepting underside of snout and around mouth (which regions are essentially smooth, though a few minute denticles are present), small areas at axils of fins, and distal margins of fins. Denticles everywhere wide spaced but more or less uniformly distributed. Largest denticles along dorsal surface of trunk, especially above lateral line; towards ventral surface, and on to head and tail, denticles gradually decrease in size. Several sizes of denticles present, particularly on dorsum of trunk, where minute denticles of uniform size are dispersed throughout larger denticles of various sizes.

Lateral line conspicuous from above 2nd gill-opening to tip of tail as a white furrow with closely approximated edges, the upper slightly overhanging the lower. It is crossed at irregular intervals along its length by narrow, thin, membranous bridges, though considerably more of it is open than is covered Inside the furrow and supporting its walls, are numerous, regularly-placed skeletal elements in the form of incomplete rings, presumably derived from

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Text-fig. 3.—Echinorhinus cookei, juvenile male 445 mm (Dom. Mus. No. 2792) New Zealand. Fig. A, lateral view and insets of transverse sections of snout and peduncle Fig. B, ventral view of head. Fig. C, dermal denticles and lateral line from halfway along trunk, drawn from dried skin Figs. D-E, upper and lower teeth, right side cl, level of cloaca; sp, spiracle.

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dermal denticles. The free ends of these rings are pointed (many are bifurcated also) and project slightly rearwards, forming a fringe or palisade of spines along each edge of the furrow. Anterior to the 2nd gill-opening the lateral line is continued on to the head by a row of lateral line pores.

Head measured to 1st gill-opening 5.0 in total length. Head depressed, flat above and below, and wedge-shaped in profile. Least fleshy interorbital distance 2.0 in head. Snout length measured to eye, 3.0 in head; snout contour bluntly pointed at snout tip; a poorly defined dorsolateral edge along each side of snout from eye to snout tip. Eye small, scarcely longer than high, its horizontal diameter 3.7 in length of snout, its anterior margin just in front of level of mouth. Spiracle minute, placed almost 2½ eye-lengths behind eye. Gill-openings large, nearly vertical, concave, and in a horizontal series anterior to pectoral base. Lengths of gill-openings increasing greatly from 1st to 5th, the 5th almost twice the 1st and 1.4 in the snout length. Interspaces between the gill-openings decreasing rearwards. Nostrils large, transverse, placed two-thirds of distance back from snout tip to mouth. Interspace between nostrils about equal to their distance from snout tip. Each nostril with a large, ovoid, median aperture and a smaller, circular, lateral aperture, the two apertures separated by triangular, nasal flaps. The medial aperture margined in front and behind by a membranous band; the anterior part of this band, which is twice the height of the posterior, is continued laterally to the pointed tip of the rearward-directed, anterior nasal flap. Posterior nasal flap fleshy and internal to the anterior flap. Mouth a high crescent, its height about half its width, and the latter about half the length of the head measured to 1st gill-opening. Preoral clefts short and shallow, only two-thirds the length of the posterior, labial furrow, and the latter about ¼ of distance from angle of mouth to symphysis of lower jaw.

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Teeth 12 - 10/11 - 11, small, similar in the upper and lower jaws. Each tooth with a subrectangular base about four times as long as high, and carrying a large, laterally-directed major cusp flanked by one or two minor cusps on each side. In the upper teeth the medial margin of the major cusp is convex, and there are two minor cusps on each flank, those on the median flank being directed medially, those on the lateral flank directed laterally. In the lower teeth the major cusp is slightly more erect, its medial margin almost straight, and there is only one minor cusp on the lateral flank, though two on the median. All the cusps are sharply pointed and smooth-edged Bases of adjacent teeth only slightly separated except at symphysis of lower jaw where there is a large median interspace. Teeth at centre of mouth slightly smaller than those midway out, though towards angles of mouth the teeth again decrease in size. One row of functional teeth in each jaw.

Inside the mouth behind the upper teeth is a prominent maxillary velum, parallel to the jaw, some 45 mm deep at the midline, and tapering uniformly on each side as it extends to the angles of the mouth. Posterior edge of velum fimbriated, while anteriorly there is a band of many fleshy papillae occupying about ¼ of the depth of the velum.

Both dorsal fins for back on the trunk, small, sub-equal in size, brush-shaped, their apices and posterior free corners rounded, their margins almost straight. 1st dorsal origin over anterior third of pelvic base, insertion just behind middle of pelvic base. Height of 1st dorsal 1.6 in the length of its base, and just less than height of 2nd dorsal. Posterior free corner of 1st dorsal over insertion of pelvic base, and just anterior to origin of 2nd dorsal Interspace between 1st and 2nd dorsals ⅘ of the 1st dorsal base. Posterior free tip of 2nd dorsal just anterior to epiural origin. Caudal measured from hypural origin 3.8 in total length. Caudal scythe-shaped, without a subterminal notch. Epiural lobe moderately developed, hypural lobe deep. Margin of epiural straight for most of its length, convex distally, length of epiural margin twice that of hypural. Anterior margin of hypural weakly convex, apex broadly rounded, and posterior margin concave. Tip of tail bluntly pointed Pectorals lobate, about 1½ times as long as broad, and with long bases, anterior and posterior margins of pectorals convex, distal margins weakly concave; corners broadly rounded Length of anterior pectoral margin 1.1 in distance from rear edge of eye to 1st gill opening. Pelvics very large, triangular, length of their bases equal to length of anterior pectoral margin. Anterior and distal margins straight, apex broadly rounded, and posterior free tip sharply pointed. Claspers tapered, rather slender, extending little behind the posterior free tip of pelvic.

Colour (fresh). Overall greyish-brown, distal margins of fins black; underside of snout and around mouth white, iris black, outer margin of eyeball iridescent greenish-blue.

(b) Juvenile. (Text-fig. 1, A-B Text-fig. 3, A-E)

Description as for adult male, except as noted below; based mainly on 445 mm male Trunk and peduncle slender; depth of peduncle 2.0 in snout length anterior to eye. Length of body measured to cloaca 59.6% of total length.

Dermal denticles of two sizes, the smaller 0.5 mm to 0.8 mm diam across the base, the larger 1.5 mm to almost 20 mm diam. The smaller denticles are close set, and cover the entire fish except for small areas at axils of fins, along distal margins of fins, and a narrow

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band along the lower lip. The larger denticles are present chiefly on the dorsal surface of trunk and head, and are well-spaced but more or less regularly distributed, their greatest concentration being above the lateral line between head and 1st dorsal fin; below the lateral line they are less numerous, as they are on the head and snout, on the peduncle and along the proximal part of the epiural lobe of the caudal. These large denticles are newly erupted or erupting, evidently at a fairly rapid rate for the 472 mm male shows considerably more of them than the slightly smaller 445 mm male.

Lateral line a conspicuous white furrow from the 2nd gill-opening rearwards, where it is completely open, without membranous bridges and with its upper and lower edges more separated than in the adult. The supporting skeletal rings have simple (not bifurcated) ends. Lateral line pores on head and snout very prominent.

Head measured to 1st gill-opening 4.4 in total length. Snout length measured to eye 2.4 in head. Eye large, its horizontal diameter 2.4 in length of snout. Spiracle minute, placed little more than an eye length behind eye. Length of 1st gill-opening 1.4 in length of 5th.

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Teeth 11 - 10/11 - 9 in the 445 mm male, 12 - 11/10 - 10 in the 472 mm male, one-cusped and similar in the upper and lower jaws. Each tooth with an ovoid to subrectangular base about twice as long as high, and carrying a sharply-pointed, smooth-edged blade-like cusp. The cusps are enamelled only on their margins, and are so strongly reflexed as to lie horizontal; hence their median margins form a continuous cutting edge.

Dorsal fins with convex anterior margins. First dorsal origin just anterior to middle of pelvic base or over anterior third, insertion over or just anterior to end of pelvic base. Height of first dorsal 1.3 in its base. Posterior free corner of 1st dorsal over middle of posterior (inner) margin of pelvic Caudal measured from hypural origin 3.3 in total length. Distal margins of pectorals straight or slightly convex. Length of anterior pectoral margin equal to distance from rear edge of eye to 2nd gill-opening.

Length of pelvic base 1.4 in length of anterior pectoral margin. Claspers reaching only two-thirds of distance from pelvic insertion to posterior free tip of pelvic.

Echinorhinus brucus (Bonnaterre, 1788)

As mentioned above, because of possible confusion between this species and E. cookei, its presence in New Zealand waters cannot be confirmed from any of the five previous records, but instead depends only on the unrecorded mounted skin, 1,420 mm long, in the Otago Museum (Plate 7, C-D). No further data other than its label, “Dunedin, April, 1887”, are available: its condition and probable distortion do not warrant it being used for obtaining dimensions.

The recognition of the Otago Museum specimen as E. brucus rather than E. mccoyi (in which species Phillipps placed his 1946 record of a New Zealand specimen) depends mainly on the recent accounts in Bigelow & Schroeder (1948, p. 527; 1957, p. 135) where they describe sufficient variation in E. brucus to provide for the differences cited by Whitley (1931, p. 311) as the basis for his proposed new subgenus and species E. (Rubusqualus) mccoyi. Phillipps (1946, p. 20) holds that the gill-openings are longer in E. mccoyi than in E. brucus; but these seem subject to considerable variation. Thus while the lengths of the 1st and 5th gillopenings in the type of E. mccoyi are 2.9% and 6.4% of the total length respectively, they are 3.6% and 6.4% in a Mediterranean specimen, and 2.3% and 5.4% in a South African specimen of E. brucus (pers. comm. from Dr. Denys E. Tucker on dimensions of specimens in the British Museum).

The above leaves no grounds on which to separate and Australasian species E. mccoyi from the cosmopolitan species E. brucus. But detailed accounts and dimensions of E. brucus from any localities, including the North Atlantic as well as the Australasian region, are still needed if the currently believed cosmopolitan status of E. brucus is to be assured.

Summary

(i) A. second species of Bramble Shark, Echinorhinus cookei Pietschmann, 1928, is recorded and described from a 1,980 mm male recently caught in Cook Strait, New Zealand. E. cookei has been known only from the type taken off Hawaii, and is generally regarded as synonymous with the cosmopolitan E. brucus.

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(ii) E. cookei is not separable from E. brucus in its dimensions, external morphology or teeth, but is strikingly different in its dermal denticles which are numerous, uniformly distributed, small (not more than 4.0 mm basal diam.) and have deeply indented, angular bases and prominently ridged spins. In contrast, the buckler-like denticles of E. brucus are sparse, irregularly distributed, large (up to 15 mm or more basal diam. in solitary denticles but reaching to 35 mm in compound bucklers) with entire-margined bases and finely ridged spines. Adult E. cookei are also almost smooth under the snout and around the mouth, while in E. brucus these areas retain noticeable-sized denticles.

(iii) The Bramble Shark recorded off California by Hubbs & Clark (1945) as E. brucus is shown to be E. cookei. This confusion between the species, due to inadequacies in the earlier accounts of E. brucus, suggests the need for examination of other records of E. brucus. In this respect none of the five records of E. brucus from New Zealand are sufficiently definitive to allow confirmation as to species; but E. brucus is established as a member of the New Zealand fauna by the mounted skin of a Dunedin specimen in the Otago Museum.

(iv) Three juvenile specimens of E. cookei from Cook Strait are also described. These are markedly slender in comparison to the adult, and have one-cusped teeth as in Squalus. Their dermal denticles agree in relative size, spacing and distribution with those of the adult.

(v) The lateral line of juvenile and adult E. cookei is an open furrow from above the 2nd gill-opening rearwards. Its walls are supported by incomplete skeletal rings whose free ends project rearwards as spines along the edges of the furrow. In the adult the furrow is bridged at irregular intervals by narrow cross-connections of skin.

(vi) As the type of E. cookei is no longer in existence, the 1,980 mm male (Dom. Mus No. 2774) from Cook Strait is designated as the neotype of this species.

Acknowledgements

Various people have assisted with this study; some are acknowledged throughout the text, but others I. would like to thank are Dr. R. R. Forster, Otago Museum, for the photographs of E. brucus, and Mr. F. O'Leary, Dominion Museum for those of E. cookei used in this account, those members of the Dominion Museum staff who assisted with the preservation and handling of the New Zealand adult specimen of E. cookei, and lastly Professor L. R. Richardson, Department of Zoology of this University, for continuing encouragement, suggestions and advice.

Literature Cited

Bigelow, H. B., and Schroeder, W. C., 1948. “Fishes of the Western North Atlantic, Part I” Mem. Sears Found. Mar. Research, No. 1, Part 1, pp. 59–576, Text-figs. 6–105.

Bigelow, H. B., and Schroeder, W. C.— 1957. “A Study of the Sharks of the Suborder Squaloidea,” Bull. Mus. Comp. Zool., 117 (1). 1–150, Pls. 1–4, Figs.

Blainville, H. M. D. DE, 1816. “Prodrome d'une nouvelle distribution systematique du regne animal.” Bull. Sci. Soc. Philomatique, Paris, pp. 105–124.

Blainville, H. M. D. DE 1825 “Vertebres, classe 5, Poissons”. In Viellot, Desmarest,… Faune Francaise… Vol. 1, Part 3, 96 pp., 120 plates.

Bonnaterre, P. J., 1788. “Ichthyologie”, in Tab. Encyc. Method des trois Regnes de la nature Paris lvi + 215 pp., Pls. A, B. + 1–100.

Fowler, H. W., 1941. “The Fishes of the Groups Elasmobranchii, Holocephali, Isospondyli, and Ostariophysi… Contributions to the biology of the Philippine Archipelago and adjacent regions.” U. S. Nat. Mus. Bull. 100, Vol. 13, × + 879 pp., 30 text-figs.

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Garman, S., 1885. “Chlamydoselachus anguineus Garm.—a living species of the Cladodont Shark.” Bull. Mus. Comp. Zool., 12 (1): 1–35, 20 Pls.

Gunther, A., 1870. “Catalogue of the Fishes in the British Museum,” Vol. 8, xxv + 549 pp.

Hamilton, A., 1886. “Note on a large sunfish (Orthagoriscus mola, L) recently captured at Napier, Hawke's Bay.” Trans. N. Z. Inst., 18: 135–136.

Hubbs, C. L., and Clark, F. N., 1945. “Occurrence of the Bramble Shark in California.” Calif. Fish & Game, 31 (0): 64–67, Figs. 16–17.

Jordan, D. S., and Evermann, B. W., 1896. “The Fishes of North and Middle America”.

Bull. U. S. Nat. Mus., 47, Part I, lx + 1,240 pp.

Parker, T. J., 1884. “On the Occurrence of the Spinous Shark (Echinorhinus spinosus) in New Zealand waters,” Trans. N. Z. Inst. 16: 280–281.

Phillipps, W. J., 1928 “Sharks of New Zealand; No. 2” N. Z. Journ. Sci. & Tech., × (4): 221–226, Figs.

Phillipps, W. J.— 1946 “Sharks of New Zealand” Dom. Mus. Rec. Zool. 1 (2): 5–20, Figs.

Pietschmann, V., 1928. “Neue Fish-arten aus dem Pacifischen Ozean.” Anz. Akad. Wien., 65. 297–298.

Pietschmann, V.— 1930. “Remarks on Pacific Fishes,” Bull. Bernice P. Bishop Mus., 73: 1–24, 4 Pls.

Smith, A, 1849. “Illustrations of the Zoology of South Africa.” London, Vol. 7, Pisces, 31 Pls.

Waite, E. R., 1913. “Results of an examination of some drawings of New Zealand fishes,”.

Rec. Cant. Mus. II (1): 17–22.

Whitley, G. P., 1931. “New names for Australian fishes.” Australian Zoologist 6 (4): 310–334.

J. A. F. Garrick

M. Sc.,
Department of Zoology,
Victoria University of Wellington,
P. O. Box 196,
Wellington, New Zealand.