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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 275

Revision of the Pelagic Amphipod Genus Primno (Hyperiidea: Phrosinidae)

Thomas E. Bowman

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978

ABSTRACT

Bowman, Thomas E. Revision of the Pelagic Amphipod Genus Primno (Hyperiidea: Phrosinidae). Smithsonian Contributions to Zoology, number 275, 23 pages, 13 figures, 1978.—Four species are recognized in the genus Primno, previously considered to be monotypic. These species, which are all described and illustrated in detail, are Primno macropa Guerin-Meneville, 1836 (the type-species), P. latreillei Stebbing, 1888, and 2 new species, P. brevidens and P. johnsoni. A key to the species is given. The 2 new species are confined to tropical and subtropical waters; P. brevidens inhabits cooler water, and the large P. macropa is found only at still higher latitudes. Brief discussions are given of associations with siphonophores, food, parasites, and the conspicuous gland in the basipod of pereopod 2.

OFFICIAL, PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Bowman, Thomas E.

Revision of the pelagic amphipod genus Primno (Hyperiidea: Phrosinidae).

(Smithsonian contributions to zoology; no. 275)

Bibliography: p.

Supt. of Docs. no.: SI 1.27:275

1. Primno—Classification. 2. Crustacea—Classification. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology; no. 275.

QL1.S54 no. 275 [QL444.M315] 591′.08s [595′.371] 78–606011

Revision of the Pelagic Amphipod Genus Primno (Hyperiidea: Phrosinidae)

Thomas E. Bowman

____________

Thomas E. Bowman, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560

Introduction

The genus Primno was proposed by Guérin-Méneville* (1836) for a single new species, Primno macropa. A second nominal species, P. guerinii White (1847), was a nomen nudum, and P. macropa remained the only species in the genus until Stebbing (1888) described 3 new species: P. antarctica, P. latreillei, and P. menevillei. Stebbing's species were short-lived, being reduced to synonyms of P. macropa the following year by Bovallius (1889), and since that time Primno has been considered a monotypic genus, although Monod (1926) recognized P. macropa var. menevillei. Recently Thurston (1976) expressed doubts about the monotypy of Primno and suggested the possible existence of a series of physiological races or a group of sibling species. I believe that Thurston's doubts were well founded, and that for 88 years 4 species of Primno have been lumped as P. macropa.

The present study is based largely on the following collections: (1) From CalCOFI cruises 1, 5, 9, and 20, off the North American west coast from the Columbia River to the southern tip of Baja California (Figures 4, 8, 10). At the stations occupied oblique tows were made from about 70–0 m (Cruises 1, 5, 9) or about 140–0 m (Cruise 20). (2) Specimens from the Climax I Project in the North Pacific Central Gyre (28°N, 155°W), sent to me by Dr. Eric Shulenberger. (3) Collections from the SOND Expedition, near the Canary Islands, sent to me by Dr. Michael Thurston. (4) Collections from the Ombango cruises in the Gulf of Guinea, sent to me by Dr. S. Frontier. (5) Specimens from the Bermuda Oceanographic Expeditions, 1931, led by Dr. William Beebe. In addition to these 5 major collections, I have examined samples in collections of the Smithsonian Institution from various localities.

These collections do not give adequate worldwide coverage for a definitive revision of Primno. Only the North Pacific can be claimed to be reasonably well covered; the South Pacific and Indian Oceans are barely represented. Despite these shortcomings, the present revision is offered as a step toward better understanding of the taxonomy and distribution of the species of Primno, now obscured by the longstanding universal misconception that Primno is monotypic.

Of the 4 species of Primno that I have been able to distinguish, one, which agrees in body length and structure of pereopod 5 with Guérin-Méneville's account, appears to be P. macropa. I have not been able to confirm this by comparison with Guérin-Méneville's type-material, since the latter is not in the collections of the Muséum National d'Histoire Naturelle, Paris, and must be considered lost (J. Forest, in litt., 1 July 1976). Efforts to obtain specimens from the type-locality, “les mers du Chile,” have been unsuccessful. Thus the designation of Primno macropa given herein is not altogether certain.

Because Primno is so abundant and widespread, there are more than 50 references to it in the scientific literature. In the majority of these references it is impossible to determine which of the 4 species is referred to; hence accurate and full synonymies cannot be given here. The synonymies given for each species include only those references in which descriptions and/or illustrations contain sufficient information to identify with reasonable assurance the species referred to. The indeterminable references are given as the synonymy of Primno incertae sedis.

ACKNOWLEDGMENTS.—For making available collections that were essential to this study, I wish to thank Drs. S. Frontier, Eric Shulenberger, and Michael Thurston. Dr. Shulenberger and Dr. Frank Ferrari searched thru plankton samples from Pacific South American waters in unsuccessful attempts to obtain topotypes of Primno macropa; their efforts are much appreciated. The manuscript benefited from a careful review by Ms. Anne Cohen. My drawings were arranged and mounted by Ms. Martina Picciano.

PHROSINIDAE

Primno Guérin-Méneville

Primno Guérin-Méneville, 1836:2–3. [Type-species, by monotypy, Primno macropa Guérin.]

Euprimno Bovallius, 1889:397.

Euprimnus Fraser, 1961:43 [lapsus calami].

ETYMOLOGY.—“Nymphe, fille de l'Océan” (Guérin-Méneville, 1836). Gender feminine. Bovallius (1889:397) proposed changing the name to Euprimno, “because Primno was previously applied to a genus of Crustacea by Rafinesque-Schmaltz in 1814.” However, as Stebbing (1888) pointed out, Rafinesque's Primno was a nomen nudem, one of 11 names simply listed without further information in a paragraph that began, “Obs. Voici les noms des autres N. G. et N. Esp. des Familles Aselotia et Oniscia.” Thus Bovallius' action was invalid, and the correct name of the genus is Primno.

DIAGNOSIS.—Head produced into single very short rostrum. Pereonites 1 and 2 separate. Pereonite 7 and pleonites 1 and 2 produced posteriorly into middorsal spines. Posteroventral corner of pleonite 3 produced posteriorly into spine. Telson much narrower than urosome. Antenna 1 of art longer than head. Antenna 2 of art reduced to a tubercle. Pereopod 1 basis with large gland with duct running from center of gland to tip of dactyl. Pereopods 3, 4, and 6 simple with some teeth on margin of carpus. Pereopod 5 prehensile, entire anterior margin of carpus dentate, with several long teeth separated by groups of short teeth; propus shorter than carpus. Pereopod 7 with all segments present; dactyl digitiform, with ring of spinules at apex in art. Uropods pointed; uropods 2 and 3 with teeth on outer margins. Gills without folds. (Modified from Bowman and Gruner, 1973:40.)

Key to the Species of Primno

Primno macropa Guérin-Méneville

FIGURES 1–2, 3a–c, 4

Primno macropa Guérin-Méneville, 1836:4, pl. 17: fig. la–f.—Bovallius, 1887:28.—?Spandl, 1927:168–169.—Barnard, 1930: 424–425 [at least larger specimens—up to 12 mm]; 1932: 287–288 [except 6 and 7 mm ovig. art], pl. 1: fig. 8.—Mackintosh, 1934, passim.—Hardy and Gunther, 1935, passim.—Thorsteinson, 1941:93–94, figs. 98–102.—Hurley, 1955:172–174, figs. 219–235; 1969:33, map 7.—Vinogradov, 1956:209; 1962:22.—Yoo, 1971a:59 [part]; 1971b: passim; 1972a: passim [part]; 1972b:174–175 [part.].—Sanger, 1973:20; 1974:7.—Lorz and Pearcy, 1975: 1445–1446.

Primno menevillei Stebbing, 1888:1447–1448, pl. 179, B.

Primno antarctica Stebbing, 1888:1448–1451, pl. 209, B.

Euprimno macropus (Guérin-Méneville).—Bovallius, 1889:400–407 [part].—Walker, 1907:9.—Wailes, 1929:161; 1931:41; 1933:9.—Behning, 1939:363.

Euprimno abyssalis Bowman.—Fulton, 1968:104, 109.

Euprimno macropa Guérin-Méneville var. menevillei Stebbing.—Monod, 1926:50–51, fig. 49.

ETYMOLOGY.—From Guérin-Méneville's center heading, “P. à grands pieds. P. Macropa. Guérin,” it is clear that the specific name “macropa” is composed of μαχρoσ (large) and Πoνσ (foot), and should have been transliterated as macropus. Guérin-Méneville apparently changed its ending from -us to -a to make it agree in gender with the feminine Primno. Bovallius (1889) was linguistically correct in changing macropa to macropus, but subsequent authors continued to use macropa, perhaps intimidated by Stebbing's (1904) admonition “that macropus is a more scholarly form than macropa must be admitted, but the practice of polishing and improving the names which our scientific ancestors invented has to my mind the same effect as daylight upon Melrose Abbey, where the rays of the sun ‘gild but to flout the ruins grey.’”

Stebbing's view is supported by Article 32(a)(ii) of the International Code of Zoological Nomenclature, which prohibits correction of an original spelling because of incorrect transliteration. Guérin-Méneville's spelling is therefore the “correct original spelling” as defined by Article 32(a).

DIAGNOSIS.—Body narrow, up to 21 mm in length. Rostrum truncate in dorsal view. Pereon about 3.3× as long as head, slightly longer than pleon. Middorsal spines and posteroventral spine of pleonite 3 sharper and more pronounced than in other species of Primno. Antenna 1 slightly longer than head. Buccal mass about 2/3 height of head. Pereopod 5 basis nearly 3× as long as broad; carpus slightly longer than basis, proximal 1–2 teeth short, long teeth slightly shorter than width of carpus; short teeth about as long as width of gently curved propus. Peropod 6 basis slightly more than 1/3 as wide as long, narrowed proximately, proximal part of anterior margin concave; ischium, merus and carpus narrower than in other species of Primno; propus armed with spinules on anterior and posterior margins. Pereopod 7 basis about as long as remaining segments combined. Uropod 3 with well developed medial shoulder.

REMARKS.—The large size, short proximal teeth on the carpus of pereopod 5, and the spinulose margins of the propus of pereopod 6 make identification of P. macropa relatively simple. None of the other species reaches more than 9 mm in length, and most are considerably shorter. The usual length of ovigerous females of P. macropa in the North Pacific is 12–15 mm. The greatest length reported is 21 mm (Vinogradov, 1956) for females from the western Bering Sea. Guérin-Méneville (1836) reported a length of 12–14 mm for his specimen(s).

The shape of its uropod 3 shows that Primno antarctica (“three twentieths of an inch” = 3.8 mm) is clearly juvenile. Uropod 3 of juvenile Primno are narrow with sharp apices, in contrast to the broad shape in mature individuals. This difference was pointed out by Bovallius (1889:400) and confirmed by Stebbing (1904). Stebbing illustrated a juvenile, only 1.4 mm in length, with an articulated process on the lateral margin of uropod 3 that he interpreted as an outer ramus since a muscle was inserted on it. A juvenile P. johnsoni from Bermuda, removed from its maternal marsupium, has uropods closely matching those of Stebbing's juvenile (Figure 13i). A later developmental stage is shown in a 2.3 mm P. brevidens (Figure 7e). It resembles Stebbing's P. antarctica in having a narrow pointed uropod 3 with a tooth on the lateral margin. As Stebbing (1904) suggested, this tooth may represent the outer ramus, now coalesced.

The short proximal teeth of the pereopod 5 carpus of P. antarctica (Figure 3k) places it in P. macropa. Whether the proximal teeth are long or short is apparent in very young stages, although marsupial young have a smooth carpus except for a pair of distal teeth (Figure 13e). Stebbing's (1904) 1.4 mm Primno had only 3 long teeth, and Bovallius (1899) recorded a 1.5 mm specimen in which the right pereopod 5 has 4 long teeth and the left pereopod 5 has a short proximal tooth followed by 3 long teeth. It appears that the long teeth are the first to be discernible as such, but the short teeth soon follow. Figure 5j shows the pereopod 5 of a 2.3 mm female P. brevidens; as in the adult the proximal spine is long.

FIGURE 1.—Primno macropa, art: a, habitus, lateral; b, posteroventral corner of pleonite 3; c, right mandible; d, incisor and lacinia of left mandible; e, pereopod 2; f, gland in basis of pereopod 2; g, pereopod 5; h, pereopod 6.

FIGURE 2.—Primno macropa, art: a, maxilla 1, outer lobe, anterior; b, same, posterior; c, maxilla 1, palp; d, maxilla 2; e, maxilliped (inner lobe omitted); f, pereopod 3; g, pereopod 4; h, pereopod 7; i, j, same, apex of dactyl; k, pleopod 3, anterior.

FIGURE 3.—Primno macropa, art: a, head, dorsal; b, pereopod 1; c, telson and uropod 3. Primno brevidens: d, art habitus, lateral; e, art head, lateral; f, same, dorsal; g, posteroventral corner of pleonite 3; h, art antenna 1, dorsal; i, art antenna 1, proximal segments, lateral; j, art antenna 2, proximal segments, lateral. Primno antarctica, art syntype: k, pereopod 5.

FIGURE 4.—Distribution of Primno macropa on 3 CalCOFI cruises, showing numbers of specimens per 1000 m³ of water filtered.

In P. menevillei the short proximal tooth of the pereopod 5 carpus and the shape of the uropod 3 apex reinforces Bovallius' opinion that this species is identical with P. macropa.

At the time I was preparing my doctoral thesis (Bowman, 1953), I had not seen Guérin-Méneville's original account of P. macropa and based my concept of this species on Stebbing's (1886) description and illustrations of the species herein referred to P. brevidens. I considered the large Primno to be a new species, and gave it the name Euprimno abyssalis. Plans to publish the present review were delayed, and the inevitable happened; Fulton (1968) made the name abyssalis available by quoting from my thesis to the effect that Thorsteinson's (1941) figures of P. macropa were referable to E. abyssalis. This action makes E. abyssalis a junior synonym of P. macropa.

DISTRIBUTION.—South Atlantic, S of 44°32′S (Barnard, 1932); Antarctic (Barnard, 1930); S of Australia, 48°18′S (Stebbing, 1888); New Zealand (Hurley, 1955); Chile (Guérin-Méneville, 1836); Japan, Okhotsk and Bering Seas (Behning, 1939); western Bering Sea (Vinogradov, 1956); SE Bering Sea (Sanger, 1974); Gulf of Alaska and off Nanaimo, B.C. (Thorsteinson, 1941); Strait of Georgia, British Columbia (Fulton, 1968); British Columbia (Wailes, 1929, 1931, 1933); British Columbia and Washington (Sanger, 1973); Oregon (Lorz and Pearcy, 1975).

In the CalCOFI area P. macropa was not taken on Cruise 20, which did not extend north of Point Conception. In the other cruises it occurred almost entirely north of Point Conception and was most abundant at the northern stations. Only 3 adults, all females, were taken on these CalCOFI cruises; the remaining specimens were juveniles, 3–7 mm long. The adult population presumably lives at greater depths than the 70–0 m layer sampled.

The collections of the Smithsonian Institution contain a art from Eltanin station 138, N of the South Shetland Islands (ca. 62°S, 61°W). The head is missing, and the body length without the head is about 16 mm.

Primno brevidens, new species

FIGURES 3d–j, 5–8

Primno macropa Guérin-Méneville.—Stebbing, 1888:1441–1445, pI. 178.

ETYMOLOGY.—From the Latin “brevis” (short) plus “dens” (tooth), alluding to the relative shortness of the carpal teeth on pereopod 5.

DIAGNOSIS.—Body rather stout, up to 9 mm in length. Rostrum very short, rounded in dorsal view. Pereon 2.5–3 × as long as head, subequal in length to pleon. Antenna 1 about as long as head. Buccal mass about half height of head. Pereopod 5 basis slightly more than twice as long as broad; carpus subequal in length to basis, proximal tooth long, long teeth ¹/₃ or less as long as width of carpus, short teeth about half as long as width of curved propus. Pereopod 6 basis about twice as long as wide, anterior margin convex throughout; posterior margin of propus apparently bare, but with minute spinules visible only under high magnification. Pereopod 7 basis about as long as remaining segments combined. Uropod 3 apex less abruptly pointed than in P. macropa.

TYPES.—All from CalCOFI Cruise 5, Station 901, off San Diego, California, 32°39′N, 118°09′W, 1 Jul 1949, 0035 hrs PST, oblique tow 68.3–0 m. Female holotype USNM 170203; 12 art, 6 art paratypes USNM 170204.

DISTRIBUTION.—Challenger station 287, South Pacific, 36°32′S, 132°52′W. In addition to its occurrence in the CalCOFI area (discussed below) I have identified it in collections of the Ombango in the SE Gulf of Guinea.

P. brevidens is one of the most abundant amphipods in the CalCOFI area, with the largest numbers tending to occur at the inner stations. In the 1949 cruises the southern boundaries roughly paralleled 30 m isotherms: 15°C in March, 18°C in July, and 20°C in November (Figure 8). In November 1950, however, it extended into much warmer water, more than 26°C at 2 stations. The greater southern penetration might be explained by the greater depth sampled in 1950, about 140–0 m, in contrast to about 70–0 m in 1949. It is likely that P. brevidens frequents deeper and colder water in the southern part of its range.

P. brevidens was much more abundant in July (61.2 per positive station) than in March (18.0) or November (18.5). A possible explanation is afforded by the change in vertical distribution. In both March and November the average number per positive station was about twice as great for the night stations as for the day stations. Hence at least during the day much of the population was below the depth sampled. In July the average number per positive station was about the same for day and night stations. Either the population had increased in numbers, was distributed over the same depth range, and had ceased vertical movements; or it had migrated closer to the surface so that most of it was being sampled in both day and night net tows. The latter interpretation seems more reasonable.

FIGURE 5.—Primno brevidens, art: a, palp of mandible. art: b, maxilla 1; c, maxilla 2; d, maxilliped, anterior; e, same, lateral; f–g, anterior and lateral aspects of apex of maxilliped outer lobe; h, pereopod 1; i, pereopod 2; j, pereopod 5 of juvenile, 2.3 mm; k, pereopod 7 of same juvenile.

FIGURE 6.—Primno brevidens: a, art pereopod 5; b, art pereopod 6; c, art pereopod 6; d, same enlarged section of propus; e, art pereopod 7; f, subadult art pereopod 7; g, adult art pereopod 7; h, juvenile art pereopod 7; i–k, art pereopod 7, spinulose ring of dactyl of 3 different specimens.

Primno latreillei Stebbing

FIGURES 9–10

Primno latreillei Stebbing, 1888:1445–1447, pI. 179A.

ETYMOLOGY.—“The specific name is given in honour of the celebrated French naturalist, Latreille” (Stebbing, 1888:1447).

DIAGNOSIS.—Body slender, length of ovigerous art 3.7–6.5 mm. Rostrum short, pointed. Pereon about 3.5 × as long as head, about as long as pleon. Female antenna 1 about ¹/₃ longer than head. Buccal mass about half height of head. Pereopod 5 basis slightly less than 3× as long as broad; carpus subequal in length to basis, proximal tooth long, long teeth nearly as long as width of carpus, short teeth subequal in length to width of nearly straight propus. Pereopod 6 like that of P. brevidens. Female pereopod 7 basis more than 1.5× length of remaining segments combined; art basis about equal to remaining segments combined. Uropod 3 similar to that of P. brevidens.

FIGURE 7.—Primno brevidens, art: a, telson and uropods; b, uropod 3; c–d, apex of right and left uropod 3 (from different individuals); e, apex of left uropod 3, juvenile, 2.3 mm; f, pleopod 3 of same juvenile.

TYPES.—Syntypes, British Museum (Natural History) 89–5–15–243, mounted on 4 slides as follows: art head; art pleon; art gnathopods 1–2 and pereopods 1–5; art. These are the remains of Stebbing's 2 art and 1 art from Challenger station 164D, off Sidney, Australia, 34°3′S, 152°20′E, surface temperature 67.5°F.

REMARKS.—Identification of my material with Stebbing's P. latreillei is not altogether satisfactory. Stebbing's description and illustrations are based almost entirely on the 2 immature art syntypes, and most of the information given is of little help at the specific level. Stebbing seemed to attach most importance to the structure of pereopods 5 and 7. Pereopod 5 of P. latreillei agrees closely with those of my specimens, but in pereopod 7 illustrated by Stebbing the basis is only about ¼ longer than the remaining segments combined, whereas in my specimens it is more than ½ longer. This discrepancy might favor identifying P. latreillei with the larger species described herein as P. johnsoni, but the shape of the pereopod 7 basis of P. latreillei is quite unlike the pyriform basis in P. johnsoni. In my opinion the shape of the basis is less likely to vary than the combined length of the remaining segments, hence I have assigned the smaller species to P. latreillei.

FIGURE 8.—Distribution of Primno brevidens on 4 CalCOFI cruises, showing numbers of specimens per 1000 m³ of water filtered. Some isotherms at depth of 30 m are shown.

FIGURE 9.—Primno latreillei, art: a–b, head, lateral and dorsal; c, posteroventral corner of pleonite 3; d, telson and uropods; e, maxilliped, outer lobe; f, pereopod 1; g, pereopod 2; h, pereopod 3; i, pereopod 6; j, pereopod 5; k, pereopod 7 (California); l–m, pereopod 7 (Gulf of Guinea). art: n, pereopod 7, juvenile (3.3 mm) (Gulf of Guinea). art: o, pereopod 7, basipod, syntype, Challenger station 164D (E of Australia).

FIGURE 10.—Distribution of Primno latreillei on 4 CalCOFI cruises, showing numbers of specimens per 1000 m³ of water filtered. Some isotherms at depth of 30 m are shown for Cruise 9. Star on Cruise 1 map shows Cruise 5 (July) station where P. latreillei was collected.

I borrowed the type-material of P. latreillei from the British Museum (Natural History) through the kindness of Dr. Roger J. Lincoln. The mountant had deteriorated so badly that I could not make out clearly the details of the mounted appendages, but I was able to confirm the shape of the pereopod 7 basis (Figure 9o).

DISTRIBUTION.—Off Sydney, Australia. In addition to its occurrence in the CalCOFI area (discussed below) I have identified it in collections of the Ombango in the southeast Gulf of Guinea, and in collections made by the Hebrew University of Jerusalem in the eastern Mediterranean and the Red Sea.

In the CalCOFI area P. latreillei occurred in limited numbers on all 4 cruises studied. It was never abundant, and generally occurred well offshore. In Cruise 5 (July) it occurred at only 1 station, but in Cruise 9 (November) it was taken at 20 stations, and its northern boundary roughly paralleled the 17°C isotherm at 30 m (Figure 10).

Only a small part of the P. latreillei population extends into the CalCOFI area, but its distribution there is consistent with that of a “Central” species (Brinton, 1962; McGowan, 1971, 1974). Additional evidence that P. latreillei is a Central species is given by its abundance in the North Pacific Central Gyre, where it is one of the most numerous hyperiids (Shulenberger, 1977).

Primno johnsoni, new species

FIGURES 11–13

Primno macropa Guérin-Méneville.—Dakin and Colefax, 1940: 123, fig. 210.—Shoemaker, 1945:234, 236.—Thurston, 1976: 427–431.

Euprimno macropus (Guérin-Méneville).—Chevreux and Fage, 1925:416–417, fig. 411.

ETYMOLOGY.—Named for Dr. Martin W. Johnson, in recognition of his many significant contributions to our knowledge of zooplankton.

DIAGNOSIS.—Closely resembling P. latreillei in almost all respects, but larger, up to 9 mm. Female pereopod 7 basis broader proximally, hence more pyriform in shape; about 1.2× length of remaining segments combined. Uropod 3 distal end with well developed medial shoulder.

TYPES.—All from Beebe's “cylinder,” an area about 8 mi in diameter located about 5 mi SE of Bermuda (Beebe, 1931); collections made during Bermuda Oceanographic Expeditions, 1931, led by William Beebe. Holotype: USNM 170234 ovigerous art, 7.8 mm, net 1249, 1 Sep 1931, towed at 1280 m. Paratypes: Net 1249, 2 art (USNM 170235); Net 1072, 10 Jul 1931, 549 m, 6 art (USNM 170236); Net 1088, 15 Jul 1931, 549 m, 17 art (USNM 170237); Net 1218, 24 Aug 1931, 1280 m, 2 art (USNM 170238); Net 1247, 1 Sep 1931, 914 m, 17 art, 2 art (USNM 170238); Net 1248, 1 Sep 1931, 1097 m, 2 art, 1 art (USNM 170240); Net 1312, 16 Sep 1931, 732 m, 19 art, 2 art (USNM 170241) (data from Beebe, 1932).

REMARKS.—P. johnsoni is very close to P. latreillei. If adequate material eventually shows a continuum in the characters used to separate them, it will be necessary to combine them under P. latreillei. At present it seems best to keep them separate to prevent a possible loss of zoogeographic information such as has resulted in the past from lumping all forms of Primno under P. macropa. I have had available only the Beebe Bermuda material (Shoemaker, 1945) in the collections of the Smithsonian Institution, and a large series from the SOND Cruise, 1965, kindly loaned to me by Dr. Michael H. Thurston.

DISTRIBUTION.—Off New South Wales, Australia (Dakin and Colefax, 1940); Bermuda (Shoemaker, 1945); Canary Islands (Thurston, 1976).

Primno incertae sedis

Primno macropa Guérin-Méneville—Stebbing, 1904:38–40, figs. 1–2.—Tattersall, 1906:25.—Stewart, 1913:258.—Spandl, 1924a:25; 1924b:266–267; 1927:168–169.—Barnard, 1925:375; 1931:128; 1937a:187; 1937b:4.—Pirlot, 1939:40.—Hurley, 1956:17–18; 1960a:113; 1960b:280.—Bogorov, 1958:156.—Bary, 1959: passim.—Irie, 1959:32.—Ruffo, 1959:34.—Kane, 1962:307–309, fig. 5.—Siegfried, 1963:9.—Pillai, 1966:216–218, fig. 9.—Brusca, 1967a:386–387; 1967b:450–451; 1973:17.—Dick, 1970:63, fig. 9.—Bradford, 1972:68.—Repelin, 1972: passim.—Tashiro and Jossi, 1972:9, 19, 31.

Euprimno macropus (Guérin-Méneville).—Chevreux, 1900:148.—Lo Bianco, 1902:419, 423, 425, 426, 447; 1903:198.—Vosseler, 1901:87–88, pl. 8, fig. 21.—Chilton, 1921:234.—Stephensen, 1924:143–146, chart 22; 1947:77.—Schellenberg, 1927:643–644, fig. 46.—Pirlot, 1929:130–131; 1930:22.—Bulycheva, 1955:1048.—Hure, 1955:50–51; 1961:34.—Hure, Scotto di Carlo, and Basile, 1971: passim.—Reid, 1955:23.—Vives, 1966:96; 1968:460.—Vives, Santamaria, and Trepat, 1975: 58–59.—Jossi, 1972:120, fig. 12.—Daniel, 1973: passim.

Euprimno macropa (Guérin-Méneville).—Walker, 1909:52.

Primno macropus Guérin-Méneville.—Chevreux, 1935:181–182.—Gamulin, 1948:179.

Euprimnus macropus (Guérin-Méneville).—Fraser, 1961:43 [lapsus calami].

FIGURE 11.—Primno johnsoni, art: a, pereopod 1 (Bermuda); b, pereopod 2 (Bermuda); c, pereopod 1 (Canary Is.); d, apex of same, slightly flattened under cover glass; e, pereopod 3 (Canary Is.); f, pereopod 6 (Bermuda); g–h, pereopod 7 (Canary Is.), lateral and medial; i, pereopod 7 (Bermuda).

REMARKS.—The widespread distribution of Primno in the world oceans, except the Arctic and the Atlantic north of about 60°N, is well documented, but the majority of published records of Primno do not give information that will permit specific identification. Consequently, distributions of the 4 species of Primno are poorly known. P. macropa lives at higher latitudes and in cooler waters than the other species, being found in subarctic and subantarctic parts of the Pacific and in the Antarctic. It is absent from the Indian Ocean, and has not been found north of 44°32′S in the Atlantic. P. brevidens inhabits warmer water than P. macropa, but, unlike P. latreillei and P. johnsoni, is not limited to tropical and subtropical regions. P. latreillei is probably circumtropical-subtropical, but P. johnsoni is thus far known with certainty only from the Atlantic.

FIGURE 12.—Primno johnsoni, art: a–b, posteroventral corner of pleonite 3; c, maxilliped; d, pereopod 4; e, pereopod 5; f, pleopod 1; g, telson and uropods.

FIGURE 13.—Primno johnsoni, young from marsupium: a, antenna 1; b, pereopod 1; c, pereopod 3; d, pereopod 4; e, pereopod 5; f, pereopod 6; g, pereopod 7; h, pleopod 1; i, telson and uropods.

I have given above, in the form of a synonymy, references to Primno that I cannot assign to species. Doubtless some records in the voluminous literature on marine plankton have escaped my attention, but I believe the listing above is reasonably complete back to Bovallius (1889). References prior to 1889 are given in detail by Bovallius, and are not repeated here.

The Canadian literature contains a number of reports that may or may not be considered publications under the provisions of the International Code of Zoological Nomenclature. Since the provisions of the ICZN as to what constitutes publication are undergoing revision, I am not including these “gray area” reports in my synonymies or bibliography. Most of them can be found in Shift, Figueira, and Grainger (1971).

Natural History of Primno

ASSOCIATIONS.—Recent observations by Madin and Harbison (1977) and Harbison, Biggs, and Madin (1977) support the view that most hyperiids spend at least part of their lives in association with gelatinous zooplankton. The only report known to me of such an association for Primno is that of Daniel (1973), who found “Euprimno macropus” within the posterior nectophores of the siphonophores Abylopsis tetragona (Otto) and Sulculeolaria chuni (Lens and Van Riemdijk).

FOOD.—I have examined the contents of the midguts of 8 specimens of P. brevidens. Most of the contents consisted of unrecognizable granules, but there were some remains of crustaceans: parts of copepods, crystalline cones from compound eyes, pereopods of a small hyperiid (Parathemisto pacifica), and pereopods of an unidentified platysceloid amphipod.

PARASITES.—In 4 of the 8 midguts examined, there were from one to many unidentified gregarines. No gregarines have been reported from Primno, but from Phrosina semilunata (a member of the same family, Phrosinidae) the gregarine Cephaloidophora phrosinae was recently described by Théodoridès and Desportes (1975).

GLANDS.—In all species of Primno a conspicuous gland occupies most of the basis of pereopod 2. The gland is composed of larger proximal and smaller distal parts. At the juncture of these parts is a vesicle from which collecting ducts radiate throughout the gland. An efferent duct runs from the central vesicle thru the distal segments and opens at the tip of the dactyl.

The function of the gland is unknown. Similar glands were found in pereopods 5–7 of Phronimella by Mayer (1878), who suggested they might be poison glands, but Laval (1968), in observations on living Phronima curvipes, found no evidence to support Mayer's hypothesis.

Glands have been reported from the pereopod 1 basis of several members of the gammaridean family Lysianassidae, e.g., Eucallisoma glandulosa Barnard, 1961. Their function is unknown; possible uses for the glandular secretions suggested by Barnard—paralysis, digestion, excretion of toxic wastes—remain to be tested.

PEREOPOD 7.—The differences in structure between the male and female pereopod 7 appear to have been overlooked in the past. In the female the dactyl is slightly expanded at its apex into an oblique oval surface that is ringed by spinules. A few additional spinules are situated in the center of the oval surface. In the adult male the segments distal to the basis are longer than in the female, and the digitiform dactyl lacks the spinulose ring. Young males, however, may have the spinulose ring.

The function of the specialized pereopod 7 is unknown, but it seems likely that it may play a role similar to that in Vibilia, which also has a digitiform pereopod 7 dactyl with a ring of spinules. In Vibilia, Laval (1963) observed that toward the end of the incubation period of about 20 days, the larvae begin to move about in the marsupium. The mother then reaches into the marsupium with the very flexible 7th pereopods, grasps a larva between the dactyls, and deposits it on the surface of a salp. The larva enters the salp's interior through a siphon and develops there, feeding on the salp's tissues.

VERTICAL DISTRIBUTION.—The most informative study of vertical distribution in Primno is that of Thurston (1976) on P. johnsoni (identified as P. macropa). Thurston found that the habits of vertical migration are complex, with differences in behavior associated with sex and maturity. During the day adult males occurred at depths of 450–940 m, with a mean depth of 647 m. At night they ranged between 100 and 830 m, with a mean depth of 616 m. Thus vertical migration was insignificant. Adult females and immature and juvenile males and females were found at lesser depths than adult males and moved upward at night for distances of 110 m (adult females), 350 and 222 m (immature males and females), and 114 and 204 m (juvenile males and females), based on mean depths. Juveniles not identified to sex were found almost exclusively in the shallowest samples and moved slightly downward at night.

Other studies of the vertical distribution of Primno are summarized by Thurston (1976) and need not be discussed here. Most of these studies are of limited usefulness, either because non-closing nets were used, or because the species involved is uncertain.

____________

* Guérin modified his name to Guérin-Méneville, and both versions are cited in the zoological literature without regard to the date of the change. Standard reference works usually cite only Guérin-Méneville (e.g., Larousse's Grand Dictionnaire Universal du XIXe Siècle, Catalog of the Library of the British Museum (Natural History), Library of Congress (catalog of printed cards), a practice that is followed herein.

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1971a. Pelagic Hyperiids (Amphipod-Hyperiidea) of the Western North Pacific Ocean. Journal of the National Academy of Sciences, Republic of Korea, Natural Sciences Series, 10:39–89.

1971b. The Biology of the Pelagic Amphipod, Primno macropa Guér., in the Western North Pacific, 1:Systematics. Korean Journal of Zoology, 14(3):132–138.

1972a. The Biology of the Pelagic Amphipod, Primno macropa Guér., in the Western North Pacific, 2: Geographical Distribution and Vertical Distributional Pattern. Korean Journal of Zoology, 15(2): 87–91.

1972b. Faunal Studies on the Amphipods in Korea. Floral Studies on Some Taxa of Plants and Faunal Studies on Some Taxa of Animals in Korea, R–72–82: 163–181. Ministry of Science and Technology.

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