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Zoology Publications from Victoria University of Wellington—Nos. 49 to 51

Nemesis robusta (van Beneden, 1851)

Nemesis robusta (van Beneden, 1851)

Ergasilina robusta van Beneden, 1851, p. 97, pl. 3, figs. 1-2; van Beneden, 1870, p. 15; Valle, 1880, p. 67.

Pagodina robusta van Beneden, 1853, p. 246; van Beneden, 1870, pp. 4-5.

Nemesis robusta (van Beneden), Heller, 1868, p. 221; Valle, 1884, p. 1; Brian, 1906, p. 74; Brian, 1924, p. 400; Leigh-Sharpe, 1936, p. 410; Delamare Deboutteville and Nunes-Ruivo, 1953, p. 211; Nunes-Ruivo, 1956.

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Nemesis robusta (van Beneden), 1851) from Alopias vulpinus. Female—fig. 51: dorsal view; fig. 52: anterior, ventral view; fig. 53: first antenna; fig. 54: second antenna; fig. 55: mouth tube and mandibular palp; fig. 56: tip of maxilla; fig. 57: maxilla; fig. 58: maxilliped; fig. 59: first pereiopod; fig. 60: second pereiopod; fig. 61: third pereiopod; fig. 62: fourth pereiopod; fig. 63: fifth pereiopod; fig. 64: caudal lamina.

Nemesis robusta (van Beneden), 1851) from Alopias vulpinus. Female—fig. 51: dorsal view; fig. 52: anterior, ventral view; fig. 53: first antenna; fig. 54: second antenna; fig. 55: mouth tube and mandibular palp; fig. 56: tip of maxilla; fig. 57: maxilla; fig. 58: maxilliped; fig. 59: first pereiopod; fig. 60: second pereiopod; fig. 61: third pereiopod; fig. 62: fourth pereiopod; fig. 63: fifth pereiopod; fig. 64: caudal lamina.

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N. pallida Wilson, 1932, p. 464, pl. 33, figs, b-p; Wilson, 1935, p. 340; Bere, 1936, p. 604; Wilson, 1937, p. 29; Pearse, 1947, p. 9; Barnard, 1948, p. 250; Causey, 1953, p. 14.

N. aggregatus Cressey, 1967, p. 6.

Material

On Alopias vulpinus: 21 females and 5 males probably from Cooks Strait or near this area, collected by J. A. F. Garrick, 11 March 1960(Dominion Museum Collection).

Description

Female (figs. 51-64).

Overall length 3.0-5.2 mm, composed of two types of individual, contracted forms (3.0 mm-3.9 mm) in which the second, third and fourth thoracic segments overlap each other slightly (see Plate A figs 1-2) and extended forms (4.3 mm-5.2 mm) in which the second, third and fourth thoracic segments are not overlapped to any extent and may be separated by a short neck (see Plate A figs. 3-5). Of 19 individuals, 10 were contracted, the other nine extended, see Graph 1 below.

Graph 1

Graph 1

Cephalothorax subovate, width four-fifths length (1.3 mm-1.45 mm × 1.1 mm-1.2 mm), separated from the second thoracic segment in extended specimens by a short neck (ca 0.2 mm × 0.7 mm).

Second thoracic segment, length half width (0.75 mm-1.05 mm × 1.6 mm-1.85 mm), anterior angled broadly rounded, posterior angles less broadly rounded, segment widest posteriorly, narrowing by almost one-third anteriorly, separated from third thoracic segment in extended specimens by a short neck (ca 0.07 mm × 1.2 mm).

Third thoracic segment subrectangular, length two-fifths width (0.7 mm-0.9 mm × 1.6 mm-1.9 mm), angles rounded, narrowing very slightly anteriorly, separated from fourth thoracic segment in extended specimens by a short neck (ca 0.06 mm × 1.0 mm).

Fourth thoracic segment subrectangular, length half width (0.65 mm-0.8 mm × 1.4 mm-1.7 mm), angles rounded, anterior angles slightly more broadly rounded than posterior.

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Fifth thoracic segment, length three-quarters width (0.45 mm-0.75 mm × 0.75 mm-0.9 mm), anterior margins sublinear, lateral and posterior margins united in a somewhat irregular entire curve.

Genital segment, length half width (0.3 mm-0.35 mm × 0.6 mm-0.7 mm), lateral margins rounded.

First abdominal segment subrectangular, length one-third width (0.07 mm-0.13 mm × 0.33 mm-0.40 mm), narrowing slightly posteriorly.

Second abdominal segment disc-shaped, length one-third width (0.06 mm-0.12 mm × 0.29 mm-0.32 mm).

Third abdominal segment subrectangular, length two-thirds width (0.18 mm-0.21 mm × 0.27 mm-0.35 mm), posterior angles slightly rounded, the caudal laminae borne laterally on posterior margin.

Caudal laminae subovate, width one-third length (0.20 mm-0.25 mm × 0.07 mm-0.10 mm), with six spines on distal margin, the innermost and the two outermost being very small.

Egg strings missing or damaged in all specimens so that total length could not be reliably measured.

First antenna apparently of twelve segments, their proportionate lengths being given by the following ratio—16:8:2:4:2:4:2:3:3:2:5:7; all segments subrectangular, distal segment rounded distally; first segment length twice width; second segment as long as wide, with two setae on distal margin and six setae on outer margin; third segment length half width, with one seta on distal margin; fourth segment length two-thirds width, with four setae on or near outer distal angle; fifth segment, length two-fifths width, with one seta on outer distal angle; sixth segment, length three-quarters width, with one seta on outer distal angle; seventh segment, length half width; eighth segment length two-thirds width, with one seta near outer distal angle; ninth segment length three-quarters width, with one seta on outer distal angle; tenth segment as long as wide, with one seta on outer distal angle; eleventh segment width two-thirds length, with one seta on outer distal angle; twelfth segment width two-fifths length, with about eight setae distally and four setae placed more proximally.

Second antenna of four segments, first and third segments subequal in length, second and terminal segments two-thirds this length; first segment subrectangular, width half length; second segment subrectangular, width two-thirds length with a few to 39 very small spines on inner distal area; third segment subrectangular, width one-third length with two to 21 very small spines in rows and patches: terminal segment, basal width one-third length, narrowing rapidly from the base to terminate in a sharp point distally, sharply curved into a claw, with three small spines near the base, one on a raised boss.

Mouth tube 0.3 mm in length, basal width two-thirds length, narrowing gradually to two-fifths this width distally, distal margin irregularly rounded.

Mandibular palp biramous, rami placed on a subrectangular base; outer ramus subrectangular, width one-third length, with one long and two short setae distally; inner ramus subrectangular, width two-fifths length, with two long setae distally.

Maxilla of two segments, segments subequal in length, first segment subrectangular, width half length; second segment width one-third length, rounded distally, with two processes distally, each one-quarter length of segment, outer process width three-quarters length, rounded distally, distal half covered in stout spines, inner process width two-fifths length, narrowing distally, with rows of small sharp spines on inner and outer margins, segment also bears a tuft of long cilia just proximal to the bases page 18 of these processes, and a group of small spines on outer margin near base of outer process.

Maxilliped subchelate, of two segments; first segment basal width half length, narrowing to half this width distally, with a blunt spine or process on inner margin near base; second segment basal width one-quarter length, narrowing gradually to a sharp point distally, segment a smooth curve, with two small spines on inner margin, the first one-quarter distance from base, the second three-fifths distance from base, the more distal spine with a small hair arising at its midpoint.

First pereiopod biramous, the basipod of two subrectangular segments, the second narrower and shorter than the first; exopod of two segments, first segment width half length, a little narrower at the base, inner distal angle expanded to increase width by one half, expansion rounded and covered in short stout spines, distal half of outer margin covered in very short spines, one spine and one very short spine on distal margin beside outer distal angle, inner margin covered in very long cilia; second segment one-third length of first, width three-quarters length, distal angles rounded, inner margin with about seven very small spines, distal margin with three long spines; basal segment of endopod two-thirds length of second segment of exopod, length two-thirds width, with a long plumose seta at outer proximal angle, and with two large processes on distal margin, each twice length of segment, the innermost, width two-fifths length, has very short spines on the rounded distal and inner distal region of margin, the outer process, which may be second segment of endopod, is rounded distally, basal width half length, slightly narrower distally, with a small spine on outer margin one-third distance from base, and a group of very short spines on and near distal margin.

Second pereiopod biramous. each ramus of two segments; basipod of two subrectangular segments, the second shorter and narrower than the first, larger than those of the first pereiopod; first segment of exopod as long as wide, inner margin two-sevenths length of curved outer margin, with a large spine on outer distal angle, a spine on inner distal angle, and a row of very short spines on distal margin near outer distal angle; second segment two-fifths length of first, as long as wide, rounded distally, with six to eight spines on distal margin, one of which may be significantly smaller than the rest, most spines about half length of segment; endopod subequal in length to exopod, first segment subrectangular, length half width, with very large spines on outer and inner distal angles, and a row of very small spines on distal margin near outer distal angle; second segment two-thirds length of first, as long as wide, rounded distally, with five or six spines on distal margin, one of which may be significantly smaller than the rest.

Third pereiopod biramous, each ramus of two segments, the basipod and segments similar in shape and size to those of second pereiopod, the first segment of exopod with a large spine on outer and inner distal angles, and a row of small spines on distal half of outer margin, distal margin of exopod second segment with six or seven spines, of which none to three may be significantly smaller than the rest, first segment of endopod with a large spine on inner distal angle and a slightly shorter spine on outer distal angle, second endopod segment with four spines on distal margin of which one may be significantly shorter than the rest.

Fourth pereiopod similar in form to second, but a little smaller; endopod first segment armed as in second pereiopod, second segment with six to eight small spines on distal margin; first endopod segment armed as in second pereiopod except that spines on distal angles are a little smaller, second endopod segment with three to six spines on distal margin.

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Nemesis robusta (van Beneden, 1851) from Alopias vulpinus. Male—fig. 65: dorsal view; fig. 66: anterior, lateral view; fig. 67: first antenna; fig. 68: second antenna; fig. 69: mandible and mandibular palp; fig 70: mouth tube; fig. 71: maxilla; fig. 72: maxilliped; fig. 73: first pereiopod; fig. 74: second pereiopod; fig. 75: third pereiopod; fig. 76: fourth pereiopod; fig. 77: caudal lamina.

Nemesis robusta (van Beneden, 1851) from Alopias vulpinus. Male—fig. 65: dorsal view; fig. 66: anterior, lateral view; fig. 67: first antenna; fig. 68: second antenna; fig. 69: mandible and mandibular palp; fig 70: mouth tube; fig. 71: maxilla; fig. 72: maxilliped; fig. 73: first pereiopod; fig. 74: second pereiopod; fig. 75: third pereiopod; fig. 76: fourth pereiopod; fig. 77: caudal lamina.

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Fifth pereiopod of one segment, small, subrectangular, width two-thirds length, with three setae on distal margin.

Male (figs. 65-77).

Overall length 3.3 mm-3.7 mm, with no evidence in this sample of the contractile powers possessed by the female.

Cephalothorax as in female, width three-quarters length (1.25 mm-1.32 mm × 1.00 mm-1.04 mm).

Second thoracic segment length three-fifths width (0.61 mm-0.66 mm × 1.07 mm-1.18 mm), narrowing slightly posteriorly and anteriorly, lateral margins rounded.

Third thoracic segment similar in shape to second but smaller, length two-fifths width (0.41 mm-0.47 mm × 1.06 mm-1.08 mm).

Fourth thoracic segment similar in shape to third, length two-fifths width (0.39 mm-0.47 mm × 0.94 mm-1.0 mm).

Genital segment subovate, anterior and posterior margins sublinear, a little wider than long (0.76 mm-0.80 mm × 0.81 mm-0.91 mm).

First abdominal segment, length three-quarters width (0.27 mm-0.34 mm × 0.43 mm-0.45 mm), lateral margins rounded.

Second abdominal segment subrectangular, angles rounded, length one-third to two-thirds width (0.11 mm-0.24 mm × 0.31 mm-0.33 mm).

Third abdominal segment subrectangular, angles rounded, length two-fifths width (0.10 mm-0.12 mm × 0.26 mm-0.27 mm).

Fourth abdominal segment subrectangular, angles rounded, length three-fifths width (0.14 mm-0.17 mm × 0.24 mm-0.26 mm).

Caudal laminae subovate, anterior margin sublinear, width two-fifths length (0.25 mm-0.28 mm × 0.10 mm-0.11 mm), with three long plumose setae on posterior margin, two smaller setae just medial to these, three small setae just lateral to these, and a row of short cilia near base of outer margin.

First antenna of twelve segments, the proportionate lengths of these segments being given by the following ratio—14:9:4:4:2:3:3:4:4:4:6:8; all segments subrectangular, distal margin of terminal segment rounded; first segment width two-thirds length; second segment width three-quarters length, with three setae placed widely apart on distal margin, and a further six setae proximal to these; third segment length half width, with two setae on distal margin; fourth segment length two-thirds width, with four setae on or near distal margin; fifth segment length half width, with one seta on distal margin; sixth segment length two-thirds width, with one seta on distal margin; seventh to tenth segments as long as wide, the seventh with one seta on distal margin, the others with two setae on distal margin; eleventh segment width half length, with one seta on distal margin; twelfth segment width two-sevenths length, with seven setae on or near distal margin, and one seta proximal to these.

Second antenna, Mouth tube, Mandibular palp, Maxilla and maxilliped as in female.

First pereiopod as in female except that second segment of exopod has small spines on outer margin instead of inner, and first segment of endopod bears three processes, all rounded distally, two of which are twice length of first segment, the other two-thirds their length, all have groups of small spines on and near their distal margins, and in addition the two longer processes have one and two larger spines respectively; the larger process with two spines is a little wider distally than the process with one spine and probably represents the second segment of the endopod.

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Table 3
Previous records (females)
Species Author Year Host (if only one specified) Seg. 2 Seg. 3 Seg. 4 Seg. 5 Gen. Seg. Abd. 1 Total Length (mm)
(width divided by cephalothorax width)
N. atlantica Wilson 1922 Scoliodon terrae-novae 1.22 1.33 1.22 0.77 0.68 0.33 5
N. macrocephalus Shiino 1957 Carcharinus melanopterus 1.28 1.22 0.97 0.71 0.39 0.29 3.14
N. pilosus Pearse 1951 Carcharias littoralis 1.47 1.47 1.14 0.80 0.50 0.31 3.0
N. versicolor Wilson 1913 Sphyrna zygaena 1.54 1.49 1.33 0.72 0.53 0.30 3.1
N. pallida Wilson 1932 Several species (type from Vulpecula marina) 1.49 1.52 1.33 0.95 0.65 0.30 4.5-5
N. pallida Wilson Barnard 1955 Alopias sp. 1.56 1.61 1.33 0.94 0.62 0.44 4.5
N. aggregatus Cressey 1967 Alopias vulpinus 1.56 1.65 1.63 0.85 0.67 0.33 4.5
N. robusta (van Beneden) Delamare Deboutteville and Nunes-Ruivo 1953 Carcharias glaucus 1.59 1.64 1.56 0.72 0.54 0.28 3.8
N. tiburo Pearse 1952 Sphyrna tiburo 1.79 7.19 1.56 0.79 0.61 0.31 2.7
N. robusta (van Beneden) Brian 1906 2.17 2.17 1.96 1.28 1.0 0.55 5
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Second pereiopod biramous, each ramus of two segments; basipod two segmented, second segment narrower and shorter than first; first segment of exopod width three-quarters length, inner margin one-third length of outer, outer margin curved, with a long plumose seta on inner distal angle, a well developed spine on outer distal angle, and a row of very short spines just proximal to the outer distal angle; second segment subovate, proximal margin sublinear, width three-quarters length, with four long setae on inner part of distal margin and four spines on outer part of distal margin; endopod a little shorter than exopod, first segment length four-fifths width, subrectangular, outer margin curved, with a well developed spine on outer distal angle, a long seta on inner distal angle, and a row of short spines along distal margin; second segment half as long again as first, width two-thirds length, distal margin rounded, with six long setae on distal margin.

Third pereiopod similar to second except as follows: rami are a little shorter, first segment of basipod bears a plumose seta on inner distal angle, distal margin of second segment of exopod bears five setae and three spines and distal margin of second segment of endopod bears five normal setae and one very stout seta.

Fourth periopod very similar to third except as follows: rami are a little larger, endopod has rows of cilia along outer margins of both segments, and lacks the row of spines along the distal margin of the first segment, and distal margin of second endopod segment has only five setae which are all normal.

Discussion

These specimens are members of the first broad group defined in the discussion of females of Nemesis lamna, i.e. small specimens (3.0 mm-5.0 mm) with a three-segmented abdomen and the fifth thoracic segment significantly smaller (25%-50%) than the fourth.

A number of described species come within this group.

In an attempt to separate these species I used the method described above to produce a ratio of widths, the cephalothorax being counted as 1, and the measurements placed in order of increasing relative width (see tables 3 and 4).

Table 4
Present material (females)
Host Area Seg. 2 Seg. 3 Seg. 4 Seg. 5 Gen. Seg. Abd. 1 Total Length (mm)
Alopias Probably 1.39 1.35 1.22 0.74 0.55 0.31 3.0
vulpinus Cooks 1.47 1.56 1.39 0.69 0.54 0.29 4.6
Strait 1.52 1.56 1.47 0.65 0.61 0.31 5.0
1.54 1.59 1.37 0.71 0.58 0.27 3.6
1.56 1.56 1.35 0.78 0.54 0.35 3.8
1.56 1.61 1.39 0.69 0.59 0.31 3.85
1.59 1.45 1.27 0.72 0.57 3.0
1.64 1.69 1.54 0.72 0.59 0.30 5.0
1.64 1.69 1.54 0.72 0.64 0.32 4.9
1.64 1.69 1.54 0.86 0.59 0.33 4.3

The following characters distinguish the described females in this group of the genus Nemesis:

N. atlantica Wilson (1922, p. 60) has a comparatively broader cephalothorax than other described species and is also unique in having notches on the lateral margins of its second thoracic segment.

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N. macrocephalus Shiino (1957, p. 392) and N. pilosus Pearse (1951, p. 362) are similar in having their fourth thoracic segment subequal in width to the cephalothorax (in other species, with the exception of N. atlantica, this segment is 33%, or more, wider than the cephalothorax) and also in having the spines on the second segment of the exopod of the second pereiopod subequal in length to this segment, in contrast to the much smaller spines described for other species. I am unable to separate these two species on the basis of descriptions in the literature.

N. versicolor Wilson (1913, p. 236) is unique in possessing a knob of each lateral margin of the cephalothorax.

N. pallida Wilson (1932, p. 464) and N. robusta (van Beneden) as described by Delamare Deboutteville and Nunes-Ruivo (1953, p. 211) differ mainly in the comparative width of their fifth thoracic segments. Considering the range of variation of this segment in the present material this character does not seem sufficient to regard these as separate species.

Cressey (1967, p. 6) described N. aggregatus which he separated from most other species on the basis of 10-12 small spines on the second segment of the second antenna, stating that these other species had 25-40 even smaller spines, and from N. tiburo and N. versicolor since in his species the row of very small spines, on the third segment of the second antenna of these species, was missing. In my material the second segment of this appendage has from a few to 39 very small spines, which vary considerably in size, although none are as large as those shown in his figure (p. 7, fig. 9). Also in my specimens the spination of the third segment varies from two to 21 very small spines, some of which may be arranged in one or two longitudinal rows. Under these circumstances the use of the spination of the second antenna as the sole criteria of taximonic distinctness must be considered doubtful until it has been investigated in further collection.

N. robusta (van Beneden) as described by Brian (1906, p. 72) is quite distinct from any described species of Nemesis in the narrowness of its cephalothorax which is about half the width of the second thoracic segment, while in other species it is two-thirds the width of this segment or wider. However, it seems likely, considering the usual accuracy of Brian's identifications and that similar specimens have not been recorded since, that the specimen figured was simply an abberrant individual.

N. tiburo Pearse (1952, p. 217) is described from a single, small (2.7 mm total length) specimen, without egg strings. It could easily be a juvenile form of one of the above species.

The present material clearly belongs to N. robusta as described by Wilson (1932) and Delamare Deboutteville and Nunes-Ruivo (1953).

Unfortunately van Beneden's original description and figures appear to refer to the male, and his figures (1851, pl. 3, figs. 1-2) show his specimen only in side view. However his specimens seem to correspond quite closely with the males in the present collection.

Previous records which appear to be of this species include:

Northeast atlantic—on Prionace glauca, Mustelus mustelus, Dasyatis pastinaca: Belgium (van Beneden, 1851, p. 97, 1870, pp. 4, 5 and 15).

on Alopias vulpinus: Brighton, England (Leigh-Sharpe, 1936, p. 410).

on Sphyrna sp.: Mauritania (Brian, 1924, p. 400).

on Cetorhinus maximus:* Ayrshire, England (Jenkins, 1936, p. 316).

* Jenkins' record is the only one from this host fish and unless confirmed by later collections it must be regarded as doubtful. Nemesis lamna is a common parasite on this species of shark.

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Southeast atlantic—on Sphyrna zygaena and Mustelus sp.: Angola (Nunes-Ruivo, 1956 *).

on Alopias vulpinus: Table Bay, South Africa (Barnard, 1948, p. 250).

Mediterranean—on Prionace glauca: Sète (Delamare Deboutteville and Nunes-Ruivo, 1953, p. 211).

on Mustelus mustelus, Dasyatis aspersa, Raja oxyrincha, and R. macrorhynchus: Adriatic (Valle, 1880, p. 67).

on Hexanchus griseus: Adriatic (Valle, 1884, p. 1).

Northwest atlantic—on Alopias vulpinus, Carcharhinus milberti, C. obscurus, Carcharodon carcharias, Odontaspis taurus, Galeocerdo cuvieri: Marthas Vineyard, Mass. (Wilson, 1932, p. 464).

on Carcharhinus limbatus: Beaufort, Carolina (Pearse, 1947, p. 9).

on C. obscurus, Rizoprionodon terraenovae, Sphyrna sp., S. tiburo, smooth-toothed shark (Aprionodon isodon (?)), spot tipped ground shark: Lemon Bay, Florida (Bere, 1936, p. 604).

on C. limbatus: Port Aransas, Texas (Causey, 1953, p. 14).

on Negaprion brevirostris: Dry Tortugas (Wilson, 1935, p. 340).

So far as I am aware, this is the first record of this species outside the Atlantic and Mediterranean.

* From a translation, original page numbers not retained.