p. 199) proposed the new name Centrophorus kaikourae for Thompson's specimen, and it is under this name or as Deania kaikourae that the species is recognised by subsequent authors.

The features of Thompson's specimen leave no doubt that it belongs in the genus Deania rather than Centrophorus, for although it has laterally grooved finspines, a subterminal notch on the lower side of the caudal fin, and one-cusped teeth dissimilar in the upper and lower jaws, these characters are common to both genera; while the rounded inner corner of the pectoral fin, the attenuate head in which the preoral length is greater than the distance from the mouth to the level of the pectoral origin, and the pitch-fork shaped dermal denticles are diagnostic of Deania.

The recognition of Thompson's specimen as D. kaikourae rather than D. calcea appears to rest solely on its greater length of first dorsal base to second dorsal base than is the case in D. calcea, for in no other feature does Thompson's account differ from those of D. calcea. Thompson's specimen can no longer be found in the Canterbury Museum, so his account is the only information available on it.

In Thompson's illustration the length of the second dorsal base (measured from point of emergence of spine to posterior end of base) is 1.5 in that of the first dorsal similarly measured, while in all other species of Deania this proportion varies from 0.9 to 1.3. The difference would seem to give validity to D. kaikourae were it not for the fact that in 9 out of 10 New Zealand specimens of Deania which I have examined, and which are from the same or nearby localities as Thompson's specimen, the proportion lies between 1.0 and 1.3, and only in the tenth is it 1.6 and hence comparable to Thompson's figure.

On this evidence, the separation of D. kaikourae from D. calcea on the lengths of its dorsal fin bases is not tenable, for these are strongly variable features, with Thompson's specimen, like my tenth specimen, lying at one extreme of the variation.

Phillipps (1946, p. 18) states that D. kaikourae has a shorter tail than D. calcea, but I am not able to confirm this in specimens of comparable size, although it should be noted that the tail undergoes considerable change with growth, becoming proportionately shorter as total length increases.

Comparison of New Zealand specimens of Deania with a mature male D. calcea taken south-west of the Faroes (kindly loaned to me through the courtesy of Dr. J. R. Pfaff from the Universitets Zoologiske Museum of Copenhagen) confirms the view that D. kaikourae and D. calcea are conspecific, for I can find no feature in proportional dimensions, external morphology, or details of teeth and dermal denticles on which to separate them. The name D. kaikourae should therefore be relegated to the synonymy of D. calcea.

The New Zealand specimens of D. calcea provide evidence that the North Pacific D. eglantina Jordan & Synder, 1902, is also synonymous with D. calcea. D. eglantina (including D. aciculata (Garman, 1906), D. hystricosa (Garman, 1906) and D. rostrata (Garman, 1906) following Bigelow & Schroeder, 1957, p. 104) can be distinguished from D. calcea only because the few specimens known show marked sexual dimorphism in the shape of the lower teeth. Females of D. eglantina have strongly oblique lower teeth, while the two males which have been described have erect teeth (including a symmetrical median tooth) at the centre of the lower jaw.^*

In contrast, the lower teeth of D. calcea are described as oblique in both sexes, and in the one North Atlantic male which I have seen, there is no symmetrical median tooth. The teeth of New Zealand D. calcea agree with the description of those from the North Atlantic, though they do show some sexual dimorphism, for the females generally have more strongly oblique lower teeth than the males (Text–Fig. 1A, 1B). This dimorphism is not constant in that a few females tend

Text–fig 1—Lower right teeth at centre of jaw from New Zealand specimens of Deania calcea Fig. A—744 mm female with strongly oblique teeth Fig. B—823 mm male with less oblique teeth than the female in Fig. A. Fig. C—835 mm male with symmetrical median tooth and nearly erect paramedian tooth.

towards the less oblique teeth characteristic of the males. Of four New Zealand males examined, one 835 mm long differs markedly from the rest in having erect teeth at the centre of the lower jaw and a symmetrical median tooth (Text–fig. 1C). In these features it matches the condition in D. eglantina as illustrated in Garman (1913, Pl. 12, Fig. 2, as Acanthidium aciculatum). The consequences of this is that in the absence of other characters to separate D. eglantina from D. calcea the possession of erect lower teeth (and a symmetrical median tooth) in the only two male D. eglantina described can no longer be regarded as sufficiently distinctive to warrant specific recognition. D. eglantina (as known from Japan) is therefore proposed as a synonym of D. calcea.

Smith (1949, p. 58, Fig. 49) identifies a South African species of Deania as D. eglantina, but his illustration is of a shark which though generally similar in dimensions to D. calcea has a relatively higher and shorter first dorsal fin, the height about 1.5 in the length of its base measured from the origin of the spine. In D. calcea (now including the Japanese D. eglantina) the first dorsal fin is markedly low and long, its height 3.0 to 4.0 in its base similarly measured (Text–Fig. 3A). This difference in proportions is significant since it cannot be accounted for by change with growth, available data on specimens of D. calcea 300 mm to 1,110 mm long indicating that as total length increases there is very slight decrease in the height of the first dorsal fin relative to its length. Consequently Smith's species cannot be retained in D. eglantina, but instead appears close to, if not conspecific with the Australian D. quadrispinosa (McCulloch, 1915) which has a similarly high and short first dorsal (its height 15 in its base from spine in the illustration of the type, and about 2.0 in an 890 mm paratype, Austral. Mus. No. 1. 13536).

The remaining species of Deania currently recognised are the South African D. natalense (Gilchrist, 1922), the Philippines D. profundorum (Smith and Radcliffe, 1912), and the recently described D. elegans Springer, 1959, from off North Carolina, U. S. A. These also have high first dorsal fins like D. quadrispinosa, but D. natalense and D. profundorum differ from D. quadrispinosa and D. elegans in

that their pectoral fin tips reach to or beyond a vertical through the first dorsal spine. In D. calcea, D. quadrispinosa, Smith's “D. eglantina” and D. elegans the pectoral tips fall far short of the first dorsal spine, the distance varying from ⅓ to ⅔ the length of the pectoral fin. As this distance shows little relative increase with growth in D. calcea (unlike some other squalid species such as Scymnodon plunketi where in juveniles the pectoral tips reach almost to the spine while in adults there is a distance equal to half the pectoral fin length), the difference between D. calcea and the D. natalense-D. profundorum group is diagnostic.

The smaller size at maturity of D. profundorum (Garman, 1913, p. 219 lists a mature male at 440 mm and a female at 590 mm) contrasts with D. calcea where maturity is not yet known at less than 700 mm in either sex. D. natalense is unusual in the shape of the inner corner of its pectoral fins, which Gilchrist (1922, pl. 7,

Text–fig. 2.—Deania calcea, male 823 mm (Dom. Mus. No. 2688) from New Zealand. Fig. A—Group of dermal denticles from high on side at level of 1st dorsal, and including three forms of denticles—viz., I = simple trifid denticle; ii = denticles with a fourth tooth projecting vertically above pedicle at anterior end of median ridge; iii = denticles similar to form ii but having an extra process or tooth on each side arising from the lateral ridges. Fig. B—lateral view of simple trifid denticle (form 1 in Fig. A) Fig. C—lateral view of form ii denticle. Fig. D—Lateral views of form iii denticles showing two shapes of the processes from the lateral ridge. Fig. E—Anterior view of form ii denticle showing fourth tooth at anterior end of median ridge Fig. F—external view of denticles from the flank of a large female (1,030 mm, New Zealand) showing three shapes of the angular processes which are sometimes present on the lateral margins of the median teeth of the largest denticles

steep-sided, sharp-topped ridge along their length. On most denticles there is also a fourth tooth, smaller than the others, projecting from the anterior end of the median ridge, more or less above the pedicle, with its tip directed vertically away from the blade. A few of the larger denticles have in addition to this fourth tooth, two further tooth like processes, one on each side, arising in a similar fashion from the ridges on the lateral teeth. There may also be an angular process, sometimes pointed, on each lateral margin of the median tooth near its posterior end, though these seem to occur only on large denticles of the largest specimens. In all cases, whether the blades have three simple pointed teeth or have some or all of the additional tooth-like processes mentioned above, they arise from erect, elongate, slender pedicles supported on rhomboidal bases. In the smaller simple denticles, the bases are strengthened by four ridges extending on to the pedicle, one from each angle. In the larger and more complex denticles there is one extra ridge and angle on each side of the anterior margin of the base. Denticles from the side of the trunk below the 2nd dorsal fin are mostly of the simple trifid form, those from the interorbital region include both simple and complex forms, but with the latter predominating; those from the upper lip are thin and leaf-shaped with a single median tooth, and either lacking in sculpture or having the ridges greatly reduced; those from the margin of the lower lip are similarly shaped but thick, almost sessile, with shallow heavy ridges; while those from the caudal axis are ovoid to subcircular in shape, with numerous longitudinal ridges converging at the posterior end where the median tooth is small, and the lateral teeth reduced and irregular in size, shape and number.

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Head measured to 1st gill-opening 4.9 to 4.7 in the total length. Head strongly depressed, elongate, and shallowly wedge-shaped in profile so that the snout tip is sharply pointed. The least fleshy interorbital distance 2.9 to 3.0 in the head. Snout long, with a well defined thin edge from the leve of the eye to the snout tip. This edge is at the mid-level of the profile between the eye and the nostril, but much nearer to the dorsal profile between the nostril and the snout tip. Contour of snout from above smoothly and rather broadly rounded at the snout tip, but widening rapidly just in front of the nostrils to give a distinct step. Behind the nostrils the contours run parallel as far as the eyes, where they gradually diverge to reach the maximum width of the head at the level of the 1st gill-opening. Eye large, ovoid, twice as long as high, its horizontal diameter 2.0 to 2.2 in the snout. Spiracle large, subtriangular, placed level with the dorsal margin of the eye, and behind it by a distance just less than its own length. Gill openings vertical, concave, and in a horizontal series anterior to the pectoral base. Lengths of the gill-openings subequal, and 2.7 to 3.0 in the horizontal diameter of the eye. Interspace between the 1st and 2nd gill-openings slightly greater than that between the 2nd and 3rd or 3rd and 4th, and about twice that between the 4th and 5th. Nostrils of moderate size, almost transverse, and placed midway between tip of snout and eye. Interspace between the nostrils 1.3 in the prenasal length. Each nostril subdivided into two apertures, one lateral and subcircular, the other medial and ovoid, by the anterior and posterior nasal flaps which are extensions of the anterior and posterior nasal margins. The anterior nasal flap is bilobed, with one pointed process dividing the whole nostril into its medial and lateral portions, and the second pointed process extending over the medial aperture. The posterior nasal flap is thick, fleshy, and internal to the anterior flap. The medial nasal aperture is margined, both anteriorly and posteriorly, by a naked membranous fold. Mouth slightly arched, its width 2.1 to 2.4 in the preoral length, and the latter 1.9 to 1.8 in the distance from snout tip to pectoral origin. The upper lip with a coarsely scalloped margin, the lower lip smooth. Preoral clefts long, reaching 3/5; to ⅔ of the distance from the corners of the mouth to the upper symphysis, and extended posteriorly as furrows for an equal length.

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Teeth generally similar in both sexes, 14-1-15/14-14 in the 823 mm male, dissimilar in the upper and lower jaws. The upper teeth erect, each with a short awl-shaped, smooth-edged cusp borne on a quadrangular base. The bases of adjacent teeth are in contact but do not overlap each other. Upper teeth at centre of mouth slightly smaller than those on either side of them, while towards the corners of the mouth the teeth also are smaller and in most specimens increasingly oblique. Three or four rows of upper teeth functional at the centre of the mouth, two or three towards the corners. The lower teeth oblique, more so in the females than in the males, each with a smooth-edged, reflexed cusp borne on a subquadrangular base. Each cusp is sharply notched laterally, and overlaps the adjacent cusp so that an almost continuous cutting edge is formed; this is particularly so towards the corners of the mouth where the obliqueness of the cusps increases to the extent where the cusps are almost horizontal. There is no median tooth in the specimens used for this description, though one male examined had a symmetrical median tooth and more or less erect paramedian teeth. The base of each lower tooth slightly overlaps its lateral neighbour. Two rows of lower teeth functional.

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First dorsal markedly long and low, originating smoothly from the dorsal profile at about the level of the axil of the pectoral. Interspace between the 1st dorsal spine and the tip of the pectoral when the latter is adpressed to the side is slightly more than half the length of the pectoral. Height of 1st dorsal 3.2 to 3.3 in that part of its base measured from the origin of the spine to the posterior insertion of the base, and the latter 1.9 to 1.8 in the head. First dorsal spine short, its exposed portion less than half the height of the fin, scarcely curved, and not reaching to the apex of the fin. The anterior margin of the 1st dorsal, posterior to the spine, broadly convex, the apex not prominent, and the distal margin almost straight. The posterior margin free from the base for a distance more than half the length of the base, and the posterior tip sharply pointed. The 2nd dorsal 1.6 to 1.5 times as high as the 1st, but with a shorter base which (measured from the origin of the spine) is 1.2 to 1.3 in that of the 1st similarly measured. Origin of the 2nd dorsal over the middle or rear part of the pelvic base, with the 2nd dorsal spine originating posterior to the latter. The 2nd dorsal spine very long and curved, its exposed portion 1.3 to 1.4 m the height of fin, its tip level with or just higher than the apex. Anterior margin of the 2nd dorsal fin convex, the apex rounded, and the distal margin straight or slightly concave. The posterior free tip pointed and extending to the level of or slightly behind the epiural origin. Caudal measured from the hypural origin 4.6 to 4.7 in the total length, the epiural lobe moderately developed, the hypural lobe rather weak. Height of the epiural lobe 4.5 to 5.0 m its length and 1.3 to 1.4 in the height of the hypural. The epiural margin almost straight, the terminal lobe truncate or slightly convex and separated from the hypural lobe by an obtuse but distinct subterminal notch. Origin of the hypural a little anterior to or below the level of the posterior insertion of the 2nd dorsal. The lower anterior margin of the hypural straight for most of its length but convex towards the apex which is little more than a right angle and not greatly rounded. The posterior hypural margin very shallowly concave. Pectorals lobate, almost twice as long as broad, the length of their anterior margins 1.8 to 2.0 in the head. The anterior margin concave for the proximal 1/5 of its length but straight or slightly convex distally. The posterior and distal margins convex, the posterior corner smoothly rounded in most specimens but slightly angular in some, and the anterior corner rounded. Pelvics small, subtriangular, the anterior and distal margins straight, the apex smoothly and broadly rounded, and the posterior free tip pointed. The claspers ovoid in section, tapered distally and sharply pointed.

Colour: Uniform light grey on most specimens, but a darker brownish grey on some larger specimens.

Etmopterus baxteri Garrick, 1957. Pl. 26, Figs. A, C.

Since the preparation of my original account of Etmopterus baxteri as a new species based on one specimen from 500 fathoms off Kaikoura, eleven more specimens have been taken. Ten of these are from Cook Strait and were caught during deep-water fishing experiments by members of the Department of Zoology, Victoria University of Wellington. The ten include one male 655 mm long, lined from 780 fathoms in November, 1956; and one male 334 mm long and eight females 261 mm to 382 mm long, taken in a trap set in 600 fathoms in July, 1957. The eleventh specimen, a female of 750 mm, was lined from 480 fathoms off Kaikoura in November, 1958, by Mr. R. Baxter, who had previously caught the type in the same locality.

The additional material, especially the smaller specimens, provides a much better basis for comparison of my species with those from other oceans. This is of particular importance as the type of E. baxteri, a female 742 mm long, was the largest known etmopterid at the time of its description. Examination of the new material now shows that in the extensive size-range represented (261 mm to 750 mm) there is considerable change with growth of those relative propotions formerly thought to be of diagnostic value. The effect of such change is that while larger specimens agree with the criteria for E. baxteri, smaller specimens are akin to E. princeps Collett, 1904, a species known only from the North Atlantic. Similarity between these two species was noted in my previous account, but E. baxteri was thought distinct in having a shorter tail region (length from pelvic origin to tip of caudal equal to distance from snout tip to posterior tip of the 1st dorsal fin in E. baxteri, but reaching to midway between 1st dorsal tip and pelvic origin in E. princeps), and a shorter caudal fin (upper caudal margin equal to distance

from snout tip to 2nd gill-opening in E. baxteri but from snout tip to pectoral origin in E. princeps). It is now evident that with increase in total length the tail region in E. baxteri decreases proportionately relative to the head-trunk length, chiefly because of an accelerated growth rate in the middle region of the trunk. As evidence of the latter, the distance from 1st dorsal spine origin to pelvic origin averages 16.8% (15.5%-18.4%) of the total length in four of my small specimens (261 mm to 290 mm long) but is 20.7% in a 655 mm specimen and 22.4% in the type (742 mm). The head length likewise decreases proportionately with increase of the total length, although to a somewhat lesser extent than the decrease of the caudal fin length. This explains the failure of my diagnosis of a shorter caudal fin relative to the head length in E. baxteri than in E. princeps, for while the length of the upper caudal margin is 18.4% of the total length and less than the length of head to 5th gill-opening (20.7%) in the 742 mm type of E. baxteri, it averages 24.6% in the four small specimens (261 mm to 290 mm) mentioned above where it is slightly greater than the head length which averages 23.8%.

Bigelow & Schroeder (1957, p.63) have suggested further dimensional criteria for separating; E. baxteri from E. princeps, their distinctions depending on the length of the interspace between the 1st and 2nd dorsal fins compared with, firstly, the distance from snout tip to pectoral fin, and secondly, the interspace between the 2nd dorsal and caudal. However, as has alredy been shown, the interspace between the dorsals is a region of accelerated later growth, and in my range of material of E. baxteri, both of Bigelow's and Schroeder's alternative characters are covered, and there is smooth intergradation between them.

I find also that my earlier statement that E. baxteri had “a more conspicuous and differently shaped pelvic flank mark; less oblique nostrils; and upper teeth mostly with 7 or 9 cusps rather than the 5 cusps in princeps” no longer holds true, for all of these features are sufficiently variable in my present material to include both E. baxteri and E. princeps. Thus the pelvic flank mark shows some change with growth, its anterior prolongation which extends over the pelvic base being much more attenuate in small specimens than it is in the large holotype. However, it is similarly attenuate in a 750 mm specimen. The number of cusps on the upper teeth increases with growth, small specimens having mostly 3 or 5 cusps per tooth, and larger specimens 5, or occasionally 7 or 9. Excepting the holotype, the upper teeth of all specimens I have examined have lesser cusps diminishing uniformly in size on each side of the major cusp. The holotype is therefore unusual in having a minute lesser cusp sandwiched between the major cusp and a larger lesser cusu.

The dermal denticles which in mature adults are rather stoutly thornlike are more slender in smaller specimens. Naked areas are also more extensive in smaller specimens, as shown by a 261 mm female where the entire ventral surface of the snout and around the mouth are smooth; all the fins are essentially smooth, including the caudal, which has only a few rows of denticles on the axis and one row along the upper margin; and the naked axillary areas are of greater extent. In a 382 mm female the snout tip, the area between the nostrils and part-way back to the mouth, and around the mouth are still smooth, but the 1st dorsal fin, the caudal fin and the paired fins have at least their proximal halves covered with denticles. In a 655 mm male, only the axillary areas, the tip of snout and the second dorsal fin remain naked.

The density of the denticles also appears to increase with growth, larger specimens having relatively more denticles per unit area than smaller specimens. Accompanying this is a change from an obvious linear arrangement of the denticles along the trunk in small specimens to a random arrangement in larger specimens (Plate 26, figs. A, C). The linear arrangement is largely obscured on the trunk of specimens more than 400 mm total length, but persist on the caudal peduncle and tail even in the largest specimens.

Fig. A —Etmopterus baxteri, juvenile female 261 mm from New Zealand.
Fig. B —E. princeps, juvenile male 262 mm (Mus Comp Zool No. 37457) from western North Atlantic.
Fig C —E baxteri, adult male 655 mm from New Zealand
Fig D —E lucifer, showing pelvic flank mark
Photos M. D. King, Victoria University of Wellington

Despite the similarities which are now apparent between E. baxteri and E. princeps, comparison of two sets of specimens of about equal size does show some differences. The most obvious of these is that the second dorsal spine in E. baxteri is strongly curved and long, reaching almost to the apex of the fin in small specimens, less so in large specimens (Plate 26, Figs. A, C). In E. princeps the second dorsal spine is scarcely curved and reaches only about ⅔ of the distance to the apex in small specimens (Plate 26, fig. B). The dermal denticles of E. baxteri are sparser and more regularly disposed than those of E. princeps (Plate 26, figs. A, B). Differences in proportional dimensions between a 261 mm female E. baxteri and a 262 mm male E. princeps (the latter Harv. Mus. Comp. Zool. No. 37457 and kindly supplied by Mr. W. C. Schroeder of Woods Hole Oceanographic Institution) are definite, the specimen of E. baxteri being considerably narrower in the head, and having smaller and generally more slender paired and median fins. However, these differences could not be sustained entrely in a comparison made by Mr. W. C. Schroeder of a slightly larger male specimen of E. baxteri

with comparable specimens of E. princeps. Mr. Schroeder was able to confirm the differences in the 2nd dorsal spine and in the density of the dermal denticles, but although he found the dorsal fins to be somewhat lower in E. baxteri, other dimensional differences were not apparent. From this it is problematical whether the differences noted in my comparison are due to variation or to specific distinction. This can be solved only by further examination and comparison of both species. As an aid to this I am including here (p. 499) proportional dimensions of six specimens of E. baxteri covering the size-range so far known. In the meantime, the differences in the 2nd dorsal spine and in the density of the denticles seem sufficient evidence for retaining E. baxteri as a separate species.

Etmopterus lucifer Jordan and Snyder, 1902. Plate 26, Fig. D.

Through the courtesy of Mr. K. Fraser, Inspector of Fisheries, Tauranga, and Mr. G. Warman, Kaikoura, I received three additional specimens of an Etmopterus in which the dermal denticles are arranged in definite, uniserial, longitudinal rows, and there is a prominent attenuate pelvic flank mark (Plate 26, Fig. D). These specimens are referable to E. abernethyi Garrick, 1957, a species described from two specimens lined from 100–200 fathorns off Kaikoura, but it is now evident that E. abernethyi is a synonym of the Indo-Pacific E. lucifer Jordan and Snyder, 1902. The additional material shows that the differences which I had previously recognised between E. abernethyi and E. lucifer express changes with growth or variation similar to those noted above for E. baxteri, and do not reflect specific distinction. It should be emphasised that since E. lucifer is one of the smallest species of sharks, all specimens recorded being less than 400 mm long, considerable changes with growth can occur with only small increases in total length. The extent of some of these changes with increments of as little as 22.0 mm in total length is discussed below.

Compared with E. lucifer, E. abernethyi was regarded as long tailed, but in five specimens now available, the length of upper caudal margin ranges from 134% of the interspace between pelvic base and subcaudal origin in the smallest (278 mm long) to 83% and 91% in the largest (341 mm and 396 mm long); in the type of E. lucifer (about 300 mm long) it is about 100% Likewise the interspace between pelvic base and subcaudal origin varies from less than the distance from snout tip to first gill-opening in my 278 mm specimen, to more than the distance to pectoral origin in the 396 mm specimen, while in the type of E. lucifer it is equal to the distance to fourth gill-opening. Thirdly, the snout length which I had thought was longer in E. abernethyi than in E. lucifer ranges from 1.5 times the eye diameter in my smaller specimens to 0.9 times in the larger, and thus overlaps the condition in E. lucifer.

It can be seen that in all three criteria given above the dimensions of the 300 mm type of E. lucifer are about intermediate between those of my smallest (278 mm) and largest (396 mm) specimens. The identification of my material as E. lucifer is confirmed by comparison with five specimens of E. lucifer from Kumanado, Japan (kindly supplied by Dr. T. Abe, East Sea Fisheries Research Laboratory, Tokyo, and Dr. R. Ishiyama, Shimonoseki College of Fisheries). I cannot find any constant differences between my material and these five E. lucifer, either in proportional dimensions, external morphology, dermal denticles, teeth or colour pattern. The colour pattern is somewhat variable, and the mid-dorsal pale band which I had thought characteristic of E. abernethyi occurs also on some of the Japanese specimens. The shape of the black, presumably luminescent areas on the ventral surface of the caudal peduncle varies from that shown in my 1957 (p. 183, Fig. 3C) account to that given for E. lucifer in Bigelow, Schroeder and Springer (1953, p. 272), but most of the specimens approach the former.

Oxynotus bruniensis (Ogilby, 1893). Text-fig. 4, A-I; Text-fig. 5, A-F; Text-fig. 6, A-H.

Although the Prickly Dogfish was listed from New Zealand waters as early as 1890 (Hutton, p. 276, as Centrina salviana Risso) on the basis of a specimen in the Christchurch Museum, and further specimens have been reported as Oxynotus bruniensis (Ogilby) by Thomson (1918, p. 5), Phillipps (1946, p. 18) and Parrott (1958, p. 116), no definitive account of a New Zealand specimen has yet appeared.

The photograph in Thomson is presumably of the Wairau Bar specimen which he records, but this is not stated. This photograph was later used by Waite (1921, fig. 32) in his account of South Australian fishes, again without reference, while in Whitley (1940, fig. 154) it is republished and credited to Waite. Parrott's recent photograph (1958, p. 116) appears to be the only illustration definitely attributable to a New Zealand specimen (other than a newspaper photograph referred to by Phillipps, 1946, p. 19) for no other illustrations have appeared except McCulloch's figure of an Australian specimen in Phillipps (1928, fig. 2).

The Prickly Dogfish is not uncommon in New Zealand waters, and nine trawled specimens (including a pregnant female with well formed embryos) taken between 1953 and 1959 have been available for this study. These were trawled from the east coasts of the North and South Islands, from just north of Cook Strait to as far south as Dunedin.

No other localities are yet known for New Zealand, though the species occurs elsewhere off Southern Australia and Tasmania. Seven of the nine specimens were

taken in 64 fathoms to 140 fathoms; one in 25 fathoms; and for one the depth is not known.

Current recognition of the New Zealand species as Oxynotus bruniensis (Ogilby, 1893) is supported by my material which agrees with the description of the type (Ogilby, 1893, p. 62) and with McCulloch's (1914, Pl. 13) excellent illustration of an Australian specimen. Likewise all my specimens fit the key diagnosis for O. bruniensis in Norman (1932, p. 77) and Bigelow & Schroeder (1957, p. 17) even though they range in size from 525 mm to 722 mm long and show some change with growth of those features used in separating O. bruniensis from the eastern North Atlantic O. centrina (Linnaeus, 1758) and O. paradoxus Frade, 1929. In particular the height of the 1st dorsal spine increases relative to that of the fin, being 45% of the fin height in the 525 mm specimen and 55% in the 722 mm specimen; the 2nd dorsal spine and fin show a similar trend from 57% to 67%. Accompanying these changes is a lengthening of the fin bases relative to the fin heights, the fins therefore becoming less slender and more obtuse. The interspace between the 1st and 2nd dorsal fins also increases slightly relative to the length of the 2nd dorsal base, being 80% of the base in the 525 mm specimen and 95% in the 722 mm specimen. As well as these changes with growth, there is also a considerable amount of variation in proportions and fin shapes between specimens of about equal size; similar variation is reported by McCulloch (1914, p. 81) in his Australian material which included six specimens from 585 mm to 600 mm long.

The key diagnosis of O. bruniensis adapted from Norman (1932) and Bigelow & Schroeder (1957) is as follows:—Spiracle subcircular, its vertical diameter less than half the horizontal diameter of eye; 1st dorsal with the spine sloping a little forwards, and supported by radial cartilages; distance from 1st dorsal spine up to apex of 1st dorsal about equal to length of 1st dorsal spine; distance from 2nd dorsal spine tip to apex of 2nd dorsal about equal to or less than length of 2nd dorsal spine; interspace between 1st and 2nd dorsal fins about equal to length of 2nd dorsal base.

In contrast to O. bruniensis, O. centrina has an ovoid to crescentic spiracle higher than half the length of the eye; the 1st dorsal fin without radial cartilages; a lower 1st dorsal fin, the distance from spine tip to apex only ½ to ⅔ as long as the spine; and a longer dorsal interspace which is not less than 1½ times as long as the 2nd dorsal base. In O. paradoxus the 1st dorsal spine slopes rearwards; the 1st and 2nd dorsal fins are very high, the distance from spine tip to apex at least 1½ times the length of the corresponding spine; and the dorsal interspace about twice the length of 2nd dorsal base.

The dermal denticles of my specimens differ according to the size of the specimen they are from in a manner similar to that reported for O. centrina in Bigelow & Schroeder (1957, p. 14). In my smaller specimens the denticles are mainly tridentate, the median tooth much the largest, and the blades directed steeply away from the skin. (Text-fig. 4, D-E.) They resemble, in these features, the denticles of juvenile Scymnodon plunketi and Centrophorus squamosus. In the denticles from larger specimens, the blades are almost vertical to the skin, and there is a stronger longitudinal keel on the posterior face of the blade; this keel usually has one prominent tooth near its distal end (as do the head denticles of juvenile Scymnodon plunketi), so that most of the blades are four-toothed as a whole (Text-fig. 4, A-C). Occasionally there are two or more teeth on the posterior keel, or the keel may be paired, while other variations include an extra lateral tooth on each side of the blade. The median longitudinal ridge on the anterior face of the blade is also thicker than it is on small denticles, and is multi-ridged, especially near its basal end.

Ogilby (1893, p. 63) and Bigelow and Schroeder (1957, p. 17) record the denticles of Australian specimens of O. bruniensis as having mostly five-toothed blades, but this is not supported by my material where the majority of denticles from the sides of the trunk of mature adults are four-toothed. In a personal communication, Mr. W. C. Schroeder informs me that their record is in error, the specimen they examined actually having four-toothed denticles.

Text-fig. 4. —Oxynotus bruniensis. Fig. A —Dermal denticles from high on side below 1st dorsal of 595 mm male, New Zealand. Fig. B— Anterior view of four–toothed denticle from side of 722 mm female, New Zealand Fig. C—Lateral view of same denticle as Fig. B, showing tooth at distal end of posterior median keel (insets show variation in keel teeth on other denticles from same specimen). Figs. D–E— Anterior and lateral views of three-toothed denticle from side of 525 mm male, New Zealand. Figs. F-I—Outlines of denticle bases from 722 mm female showing four–angled form (Fig. F) and more irregular forms (Figs. G-I), where are additional angles and ridges on anterior margin.

Text-fig. 5.—Oxynotus bruniensis, 560 mm male, New Zealand. Fig. A—Functional upper teeth drawn from complete tooth–bearing membrane removed from jaw and pressed flat; stippled teeth indicate course of row on central sector of arch. Fig. B—Functional lower teeth. Fig. C—First right upper tooth. Fig. D—7th right upper tooth. Fig. E—Diagrammatic plan of upper teeth to show the steeply arched rows, and the longitudinal series. Fig. 7—Scymnodon plunketi, 1,417 mm female (Dom. Mus. No. 2636), plan of upper teeth showing broadly arched rows and radially arranged series.

The upper teeth of O. bruniensis are one-cusped, erect and lanceolate, with about four rows. ^* of teeth functional (Text-fig. 5, A).

Viewed in situ, the teeth occupy a forward–arched quadrangular area, wider than long, at the front of the mouth. However, if the angles of the mouth are incised, a narrow band of smaller, irregular, oblique teeth is displayed on each side of the quadrangular area, trailing rearward and placed on the medial face of the upper jaw towards the angle where they evidently have little function. The arrangement of the teeth on the quadrangular area has recently been suggested by Bigelow & Schroeder (1957, p. 14) as sufficient reason for referring Oxynotus to a separate family Oxynotidae, though they give other reasons also (p. 9). They describe the teeth (p. 13) as “in quincuncial arrangement, the functional rows.

successively longer from front to rear, the first row consisting of three teeth only, the number of teeth greater by one in each successive row, and with about six rows functional.” In my material, I find that to make row counts comparable to those of Bigelow & Schroeder, it is necessary to count the teeth in definite transverse rows, as though the jaw were not arched. Also it is necessary to assume that in moving from one transverse row to the next row behind it, the teeth series making up the second row alternate in position with those of the first row. This method of counting the teeth is very practical in Oxynotus where the teeth of such successive rows are sharply delineated in situ. However, it is not a true indication of the arrangement of the teeth which in fact differs very little from that of other squaloid sharks, having several rows of upper teeth functional. If the entire upper teeth membrane is removed from the jaw and laid flat (Text-fig. 5, A, E), the rows of teeth can be seen to follow an arch around the jaw like that of other squaloids (Text-fig. 5 F shows diagrammatically the row arrangement of Scymnodon plunketi for comparison). The arch in Oxynotus is narrower than is common for most sharks and because of this it is difficult to trace the exact course of the rows at the sides of the mouth. In the central sector of the mouth the rows are well defined, and each series is represented in each complete row.

It can be seen that the morphologically most anterior row of teeth is incomplete at the centre of the mouth, but is represented by the 4th series on each side of the median tooth; similarly the second row lacks a median tooth and sometimes the 1st series on either side of the median tooth also. This incompleteness of the anterior rows is matched in the specimen of Scymnodon plunketi figured here (Text-fig. 5 F) and I have observed it also in Centroscymnus owstonii, C. crepidater and Dalatias licha; it appears to be the usual condition for those squaloid sharks where the upper teeth are well spaced. In contrast such species as Squalus acanthias, S. blainvillii and Deania calcea where the teeth bases overlap or are contiguous, the most anterior row is always complete at the centre of the mouth.

Although the rows of upper teeth of O. bruniensis are not easy to follow at the sides of the mouth, the series are very obvious because of the changing tooth shapes in each series from about the 5th outwards. It is noteworthy that these side series run longitudinally (Text-fig. 5 A, E) as do the more central series. In most squaloid sharks the side series are arranged more or less radially to the arch of the jaw (Text-fig. 5 F, Scymnodon plunketi) but in some, and particularly Dalatias licha the series are more nearly longitudinal as in Oxynotus.

In view of the above, it must be accepted that the arrangement of the upper teeth of Oxynotus is not novel in the Squaloidea and does not merit recognition as a familial character. Much the same can also be said of the other characters which have been suggested as diagnostic of the Oxynotidae. For example, the strong ventrolateral muscle ridges along the trunk have counterparts (admittedly not as strongly developed) in most squaloids. Erect, keeled and toothed dermal denticles are found at least in juveniles of Scymnodon plunketi and Centrophorus squamosus. The chief distinction of the oxynotids appear to lie in the extravagance of these and other features (such as their peculiarly high dorsal fins and long, lanceolate pectorals). Other than this extravagance they have no unique character to separate them from the Squalidae. Consequently recognition of them as a separate family Oxynotidae is largely a matter of systematic convenience.

Seven embryos of O. bruniensis found in my 722 mm female are each about 100 mm long. They were taken in December but although well-formed are attached to large globular yolk–sacs 70 mm wide by 60 mm high, and so are a long way off birth. They have external branchial filaments up to 18 mm long (shown here only in the illustration of the ventral view and only on the left side, Text-fig. 6, G-H). The general appearance is much more like that of a “typical” squaloid than of adult Oxynotus, for their bodies are slender, their dorsal fin margins less

of eye 1.5 in snout and 4.6 to 4.4 in head. Spiracle large, ovoid to subcircular, placed just above level of posterior corner of eye and behind it by a distance slightly more than its own length; length of spiracle 3.0 in horizontal diameter of eye. Gill-openings very small, vertical, concave, and in a horizontal series anterior to pectoral base. Lengths of first four gill-openings either subequal or decreasing rearwards, ¼ to ⅔ the horizontal diameter of eye; 5th gill-opening varying from slightly larger to smaller than preceding gill-openings. Interspaces between gill-openings subequal between 1st and 2nd, and 2nd and 3rd, and larger than that between 3rd and 4th, and 4th and 5th. Nostrils very large, close together, oblique, placed well forward on anterior third of snout so that the lateral nasal aperture faces almost directly frontwards. Interspace between nostrils narrow, 3.2 to 2.6 in preoral length. Lateral and medial nasal apertures round, separated from each other chiefly by the thick fleshy posterior nasal flap; medial aperture more than half covered by the extensive, convex-margined anterior nasal flap which projects rearwards; posterior margin of medial aperture bordered by a low membranous band. Mouth small, almost transverse, with thick, fleshy lips which are complexly pleated longitudinally—i.e., across the lips at right angles to the mouth. Preoral clefts long, reaching more than ⅔ of distance from angles of mouth to upper symphysis, and extended rearwards for an equal or shorter distance by abruptly shallowing furrows.

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Teeth 14/5-1-5 in a 525 mm male, 17/5-1-5 in a 560 mm male, dissimilar in upper and lower jaws. Upper teeth narrow, erect and of uniform size except for the last few series on each.

side where the teeth are oblique and diminish in size. In the central sector of mouth there are ten or eleven series of teeth with four or five rows or parts of rows functional; these occupy a slightly arched quadrangular area about twice as wide as long. The lateral two or three series of oblique teeth comprise about three rows which trail rearwards as a narrow band from each side of the quadrangular area, being placed on the medial face of the upper jaw where they cannot be seen without dissection. Each upper tooth from central sector of mouth.

Text-fig. 6—Oxynotus bruniensis. Fig. A—595 mm male, New Zealand, lateral view and insets of transverse sections of snout and peduncle. Figs. B-C—Dorsal and ventral views of head of same. Fig. D—1st dorsal fin of 525 mm male, New Zealand. Fig. E—1st dorsal fin of 722 mm female, New Zealand . Fig. F—Right nostril of 595 mm male. Figs. G-H—104 mm embryo (one of seven from 722 mm, New Zealand female), ventral view of head, and lateral view. N.B.—External gill filaments, up to 18 mm long, are omitted in. Fig.H and shown only on the left side in. Fig.G; the yolk-sac, 70 mm × 60 mm, is also omitted in both figures of the embryo. C = level of cloaca. YS = stalk of yolk-sac.

with a long, sharply-pointed, lanceolate cusp, its edges irregularly and finely serrate, borne on a bifid base. The cusp stands well clear of the base on a rather narrow pedicle which proximally is irregularly striated longitudinally. The bases of adjacent teeth are placed close together or touching but do not overlap each other. Lower teeth with only one row functional, broad, blade-like, erect and symmetrical at the centre of the jaw where there is a median tooth, but slightly oblique towards the angles. Each tooth with a high rectangular base and a triangular cusp; the margins of the cusp are irregularly and finely serrate. Bases of adjacent lower teeth overlapping laterally.

1st dorsal fin subtriangular, large, very high, its height about twice the length of its base measured from origin of exposed spine, and greater than length of head. Origin of 1st dorsal obscure; anterior margin steep in front of spine but steepning even more behind spine where it is increasingly convex towards the apex; apex pointed, less than a right angle; distal margin at first concave from apex downwards but becoming convex halfway along its length; posterior margin short, less than horizontal diameter of eye; posterior free corner right-angled or acute. 1st dorsal spine sloping forward, largely concealed, its short exposed tip above or behind posterior tip of pectoral when latter is adpressed to side; total length of 1st dorsal spine just less than or equal to distance from its tip to apex of 1st dorsal fin. 2nd dorsal similar to 1st but smaller, its height 1.3 to 1.5 in height of 1st dorsal, its base measured from origin of exposed spine 2.0 in base of 1st similarly measured. Origin of 2nd dorsal fairly well defined, almost an eye-length in front of pelvic origin; length of complete base of 2nd dorsal equal to interspace between 1st and 2nd dorsals. Anterior margin of 2nd dorsal behind exposed tip of spine less steep than that of 1st; distal margin more nearly straight. 2nd dorsal spine sloping rearwards, largely concealed, its exposed tip above or just in front of posterior insertion of pelvic base; total length of 2nd dorsal spine equal to distance from its tip to apex of 2nd dorsal fin. Caudal deep and short, its length measured from hypural origin 4.0 to 4.8 in total length; epiural lobe moderately developed, its height about 4.0 in length of its anterior margin and 1.5 in height of hypural, its anterior margin slightly convex; terminal margin straight to convex, long, its length about the length of epiural; subterminal notch shallow and obtuse, but distinct; origin of hypural well anterior to epiural; lower anterior margin of hypural convex, apex a bluntly rounded right-angle, posterior hypural margin straight, convex or slightly sinuous. Pectorals leaf-shaped, about as long as head but narrow, their greatest breadth (near base) approximating half their length; anterior margin straight proximally, becoming convex towards the anterior (outer) corner which is sharply rounded and acute; distal margin almost straight, more than ⅔ as long as anterior margin; posterior margin very short, posterior inner corner very broadly rounded and obtuse. Pelvics small, constricted at base; anterior, distal and posterior margins very slightly convex, anterior and posterior corners bluntly rounded; length of anterior margin about ¼ longer than distal margin, and ⅔ as long as anterior pectoral margin. Claspers in 595 mm male ovoid in section, rapidly tapering to their tips, which reach as far back as hypural origin; for ⅔ of their length they are joined to their respective posterior pelvic margin; subterminally each carries a strong lateral spur.

Colour: Uniform greyish-brown, but with white or translucent areas at tips of dorsal fins and claspers, along distal margins of pectorals and pelvic fins and at apex of hypural lobe of caudal; there is also in most specimens a light-coloured patch on the ventral surface beneath the gill-openings; lips pinkish-white.

Dalatias licha (Bonnaterre, 1788). Text-fig. 7, A-B; Text-fig. 8, A-H.

The Black Shark was first reported from New Zealand by Parker (1883, p. 222) who identified it as the Mediterranean Scymnus lichia, now recognised as Dalatias licha and known from the Atlantic and Pacific, both north and south, as well as the Mediterranean. Whitley (1931, p. 310) proposes the new name Scymnorhinus phillippsi for the Australasian species without indicating why it is specifically distinct except for the statement that “as McCulloch hinted (it) is obviously not conspecific with the French Squalus lichia.” McCulloch (1914, p. 81) identified specimens from the Great Australian Bight as S. licha though he says: “In using that name.

I do so only because the descriptions of the Mediterranean species are so incomplete that, in the absence of typical specimens to compare with, it is impossible to determine whether they are specifically identical or distinct.” Phillipps (1946, p. 19) accepts Whitley's name and regards D. phillippsi as distinct from D. licha in having the mouth placed further back, differently placed fins, a shorter caudal fin, as well as in various other characters.

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Head depressed, the eye large, and the snout remarkably short; trunk slender, subcircular in section anteriorly but slightly compressed posteriorly. Height of trunk at origin of pectorals about 1/9th of its length to origin of caudal. Length of body measured to cloaca 63.5% to 65.8% of the total length. Caudal peduncle without lateral keels or precaudal pits.

Dermal denticles on sides of trunk of moderate size, and loosely spaced so that there are considerable interspaces between them where the skin is exposed. Each denticle has a low,

Text-fig. 7.—Dalatias licha, male 1,132 mm (Dom. Mus. No. 1295) from New Zealand. Fig. A—Group of dermal denticles from high on side at level of 1st dorsal. Fig. B—Lateral view of denticle.

near-horizontal, thornlike blade expanded laterally at its anterior end, and united along the greater part of its undersurface with a rhomboidal base so that a pedicle as such is scarcely differentiated. The external surface of each blade carries three longitudinal ridges which converge posteriorly and meet at the tip of the sharply-pointed median tooth. Anteriorly the margins of the blade fall away steeply towards the base in a series of sharp-edged irregular facets. Denticles from the caudal axis similar to those from the trunk but with more elongate blades; those from the interorbital region have stronger sculpture and more obtusely pointed blades; those from the lower lip stand more erect and are more nearly ovoid than thornlike; while those from the upper lip are ovoid to leaf-shaped and lack ridges except at their anterior ends.

Head measured to 1st gill-opening 7.1 to 6.9 in the total length. Profile of head only slightly tapered to the thick foreshortened snout. The least fleshy interorbital distance 2.7 in the head Snout contour bluntly rounded at the snout tip, widening rapidly at the level of the nostrils, and slowly from there rearwards. Length of snout measured to the eye 5.4 to 4.6 in the head. Eye large, longer than high, its length 1.7 to 1.9 in the preoral length, the latter 3.1 in the head. Spiracle large, transverse, placed just above the eye and behind it by a distance equal to twice its own length. Gill-openings almost vertical, slightly concave, and in a horizontal series anterior to the pectoral base. Lengths of the first four gill-openings subequal and less than the horizontal diameter of the eye; the 5th gill-opening equal in length to the eye. Interspaces between the gill-openings about the same except for that between the 4th and 5th, which is only half that of the others. Nostrils oblique and placed very close to tip of snout. Interspace between nostrils about equal to length of snout measured to eye. Each nostril subdivided into a circular anterolateral aperture and an elongate ovoid posteromedial aperture by the anterior and posterior nasal flaps. The anterior nasal flap broadly triangular, with a bluntly pointed apex and a smoothly rounded medial margin. The posterior nasal flap fleshy and internal to the anterior flap. Mouth only slightly arched, its corners concealed by the smoothly curved, overhanging upper lip which extends posterolaterally on each side as far back as the level of the spiracle. Width of mouth just less than the preoral distance. Preoral clefts not visible externally when the mouth is closed, but opening on the inner face of the upper lip. Internal to the upper lip is a prominent fimbriated layer lying against the upper teeth; no such layer is present on the lower lip.

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Teeth 9-1-9/9-1-9 in a 1,010 mm male, dissimilar in the two jaws. The upper teeth elongate, each with a bifid base and a sharply-pointed, smooth-edged, recurved, awl-shaped cusp; those at the centre of the mouth noticeably smaller and more erect than those on either side, which are moderately oblique; three rows of upper teeth functional. The lower teeth blade-like, each with a high rectangular base and a narrow, strong triangular cusp; the cusps have uniformly serrated edges, the serrations continued downwards onto the lateral shoulder of each cusp where there is an acute notch. At the centre of the mouth there is a symmetrical median lower tooth, but on either side of it, towards the corners, the teeth have cusps which are slightly oblique. The base of each lower tooth overlaps its lateral neighbour. One or two rows of lower teeth functional.

1st dorsal of moderate size, triangular to brush-shaped, originating posterior to tip of pectoral when latter is adpressed to the side by a distance equal to the horizontal diameter of the eye. Length of base equal to the free posterior margin, and also to the height of the fin, and about equal to the preoral length. 2nd dorsal larger than the 1st, its base 25% longer, its shape generally the same except that the posterior free tip is more sharply pointed. Origin of 2nd dorsal about over middle of pelvic base. Caudal measured from the hypural origin 4.4 to 4.3 in the total length; epiural lobe with a straight anterior margin and a broadly rounded apex. Terminal margin convex, and the subterminal notch deeply incised and prominent. Origin of the hypural a little posterior to the midpoint between the rear tip of the 2nd dorsal and the epiural origin; lower anterior margin of hypural about half the length of the epiural margin, slightly convex distally. The apex smoothly rounded and about a right angle, and the posterior margin almost straight. Pectorals rather small, subovate, originating just behind 5th gill-opening. Anterior pectoral margin convex, its length twice the width of the fin and considerably less than the distance from snout tip to 1st gill-opening; posterior and distal margins smoothly continuous, the posterior angle scarcely apparent; anterior angle less than a right angle but broadly rounded. Pelvics large and triangular, their bases one-quarter longer than the 2nd dorsal base. Anterior, posterior and distal margins straight, the apex smoothly rounded, and the posterior free tip sharply pointed.

Colour: Uniform dark brown to black above and below; upper and lower lips, and the distal margins of all the fins, pale-coloured or whitish.