Serials The Caprellidae (Crustacea, Amphipoda) of Atlantic and Arctic Canada The Caprellidae (Crustacea, Amphipoda) of Atlantic and Arctic Canada National Museum of Natural Sciences Publications in Biological Oceanography, No. 4 Published by the National Museums of Canada Staff editor Rigmore Adamson Musée national des Sciences naturelles Publications d’océanographie biologique, n° 4 Publié par les Musées nationaux du Canada The Caprellidae (Crustacea, Amphipoda) of Atlantic and Arctic Canada Diana R. Laubitz © Crown copyrights reserved Available by mail from the National Museums of Canada Marketing Services Ottawa, Ontario K1A OM8 Catalogue No. NM95-7/5 National Museum of Natural Sciences National Museums of Canada Ottawa, Canada 1972 PO0987654321 Y798765432 Litho in Canada Contents List of Plates, 6 List of Maps, 7 Résumé, 8 Summary, 9 Biographical Note, 10 Preface, 11 Acknowledgements, 12 Notes on the Family Caprellidae, 13 Methodology, 13 Key to the Caprellidae of Atlantic and Arctic Canada, 17 Aeginella, 19 C. septentrionalis, 48 A. spinosa, 19 C. trispinis, 53 Aeginina, 21 C. unica, 53 A. aenigmatica, 21 Cercops, 57 A. longicornis, 23 C. holbolli, 57 Caprella, 26 Hemiaegina, 59 C. andreae, 26 H. minuta, 59 C. carina, 26 Luconacia, 61 C. ciliata, 31 L. incerta, 61 C. dubia, 31 Mayerella, 61 C. equilibra, 34 M. limicola, 61 C. linearis, 35 Metacaprella, 63 C. microtuberculata, 39 M. horrida, 63 C. penantis, 41 Paracaprella, 67 C. natalensis, 47 P. tenuis, 67 C. neglecta, 47 Proaeginina, 68 C. rinki, 47 P. norvegica, 69 C. sarsi, 48 Zoogeography, 71 Table 1: Known Distribution of Canadian Arctic and Atlantic Caprellidae, 70 Canadian Atlantic Region, 71 Canadian Arctic Region, 72 Discussion, 75 Literature Cited, 77 List of Plates OMAN OORWNH — 10 Aeginella spinosa, 20 Aeginina aenigmatica, 22 Aeginina longicornis, 24 Caprella carina, 28 Caprella ciliata, 30 Caprella dubia, 32 Caprella linearis, 36 Caprella microtuberculata, 38 Caprella penantis, Caprella natalensis, Caprella neglecta, 42 Caprella penantis, Caprella natalensis, Caprella neglecta, 44 Caprella rinki, 46 Caprella septentrionalis, 50 Caprella unica, 52 Cercops holbolli, 56 Hemiaegina minuta, 58 Mayerella limicola, Metacaprella horrida, 62 Metacaprella horrida, 64 Paracaprella tenuis, 66 List of Maps 10 Known distribution of Aeginella spinosa and Aeginina aenigmatica north of 45°N, 16 Distribution of Paracaprella tenuis and Aeginina longi- cornis within the Canadian Atlantic region; based on material examined, 26 Known distribution of Aeginina longicornis north of 45°N, 27 Known distribution of Capre/la carina, Caprella dubia, and Metacaprella horrida, 34 Known distribution of Caprel/la ciliata, Caprella rinki, and Proaeginina norvegica, 35 Distribution of Caprella linearis and Caprella penantis within the Canadian Atlantic region; based on material examined, 40 Known distribution of Capre/la linearis within the North Atlantic and northern Pacific boreal and_ subarctic regions, 41 Known distribution of Capre/lla microtuberculata and Cercops holbolli within the North Atlantic and arctic regions, 49 Distribution of Capre/la septentrionalis and Caprella unica within the Canadian Atlantic region; based on material examined, 54 Known distribution of Capre/lla septentrionalis within the North Atlantic boreal and arctic regions, 55 Résumé L’auteur fait état de 21 especes de caprelles provenant des eaux arctiques et atlantiques du Canada. Les observations couvrent la partie septentrionale de l’Amérique du Nord depuis la Nouvelle-Ecosse jusqu’en Alaska, ainsi que l’est et l'ouest du Groenland. On trouve dans l’ouvrage la des- cription détaillée de 17 especes et une clé servant a leur identification. L’une d’elles, Aeginina aenigmatica, semble étre nouvelle. On a placé dans le genre Metacaprella, Caprella horrida G.O. Sars. On considére Caprella sedovi Gurjanova synonyme de C. carina Mayer. Quant a Caprella hystrix Bate et Westwood et a C. ultima Bate, on les a respectivement assimilées a C. linearis (Linné) et a C. septentrionalis Kr@yer. On considére non identifiables Caprella sarsi Honeyman et C. trispinis Honeyman. Caprella natalensis Mayer et C. neglecta Mayer acquiérent une identite specifique. On observe que les mers boréales se divisent en deux grandes régions zoogéographiques, |’une arctique-atlantique et l'autre subarctique-pacifique. La faune arctique est en grande partie d’origine atlantique alors que la plupart des especes venant du Pacifique ont des répartitions tres locales. Summary Twenty-one species of caprellids are reported from the Canadian Arctic and Atlantic. The area investigated includes northern North America from Nova Scotia to Point Barrow, Alaska, and eastern and western Greenland. Seventeen species are described in detail and a key to their identification is included. One species, Aeginina aenigmatica, is described as new. Caprella horrida G.O. Sars has been placed in genus Metacaprella. Caprella sedovi Gurjanova has been synon- ymized with C. carina Mayer. Caprella hystrix Bate and Westwood and C. ultima Bate have been assigned to C. linearis (Linnaeus) and C. septentrionalis Krgéyer respec- tively. Two species, Capre/la sarsi Honeyman and C. trispinis Honeyman, are reported as_ unrecognizable. Caprella natalensis Mayer and C. neglecta Mayer have been given specific status. The north-polar seas are found to comprise two major zoogeographical regions, Atlantic-arctic and _ Pacific- subarctic. The arctic fauna is predominantly Atlantic in Origin, and most Pacific species have very localized distributions. Biographical Note Born in London, England, Diana Laubitz graduated from Cambridge University with a B.A., specializing in Zoology. She came to Canada in 1956, and has been associated with the National Museum of Natural Sciences since 1964. Her research has been concerned primarily with the taxonomy and systematics of the Caprellidea, and in 1970 she published Studies on the Caprellidae (Crustacea, Amphipoda) of the American North Pacific, in the Museum's Biological Oceano- graphy series. Recently she has started investigating some of the Gammaridea closely related to this group. Preface The Caprellidae of the western North Atlantic have recently been treated by McCain (1968). He discussed the complicated taxonomy of this family, resolved certain systematic problems and listed 29 species, describing 24 in detail. The area under consideration in his paper was the western North Atlantic from the equator to Nova Scotia. Since Mayer’s three monographic papers (1882, 1890, 1903), caprellids have been reported only incidentally from North America north of Nova Scotia. Faunal checklists and ecological surveys (e.g. Bousfield 1952-62; Bousfield and Leim 1960; Brunel 19615, 1970; Dunbar 1942, 1954) have included caprellids found in certain areas, but no work has dealt exclusively with these animals. In contrast, the arctic region around Greenland and the northeastern Atlantic have been investigated fairly thoroughly. Most of the recent Arctic reports and new species are found in Stephensen (1912-44a). The present paper deals with the Caprellidae of the boreal and arctic regions of Atlantic North America. The geographical area investigated includes North America, from Nova Scotia northward and westward to the Canadian arctic islands and northern Alaska, and eastern and western Greenland. Seven- teen species are described, one of which is new to science. Four other species are listed but not described, since they were not present in the collections investigated. One of these is a cold-water species (Proaeginina norvegica). Three have . not yet been reported from the area investigated, but have been found as far north as Connecticut (Capre/l/a andreae, C. equilibra, and Luconacia incerta). Acknowledgements The major collections studied have been those of the National Museum of Natural Sciences, Ottawa. Other unidentified material was obtained from _ Universitetets Zoologiske Museum, Copenhagen; Station de Biologie marine, Grande- Riviere, Quebec; United States National Museum, Washington, D.C.; Fisheries Research Board of Canada, Arctic Unit, Ste- Anne-de-Bellevue, Quebec. The large amount of borrowed material has considerably increased the scope and value of this paper. | must offer my sincere thanks for their contributions to: Dr. Torben Wolff (Universitetets Zoologiske Museum, Copenhagen); Dr. A. Marcotte and Dr. M. Ledoyer (Station de Biologie marine, Grande-Riviere); Dr. Thomas Bowman (U.S.National Museum, Washington, D.C.); Dr. J.W. Wacasey (Fisheries Research Board of Canada). | am very grateful to Dr. Charlotte Holmquist (Naturhistoriska Riksmuseet, Stockholm) and Dr. Marit Christiansen (Zoologisk Museum, Oslo) for the loan of type material; and to Dr. J. Forest (Muséum National d'Histoire Naturelle, Paris) and Mr. R. Ingle (British Museum of Natural History) for some of C. Spence Bate’s original material. | also thank the National Museum of Natural Sciences for supporting this study, and particularly Dr. E.L. Bousfield for his advice and critical help, and for the use of his unpublished collecting station data. | am especially grateful to Dr. J.C. McCain (Hazleton Laboratories, Falls Church, Virginia) for his advice and comments, and for the use of his manuscript catalogue and bibliography of the Caprellidae. 12 Notes on the Family Caprellidae McCain (1968) and Laubitz (1970) have both commented on the various characters used in diagnosing caprellid genera. They have emphasized that certain characters need further investigation, and have pointed out characters on which the taxonomic division of the Caprellidae should eventually be based. Vassilenko (1968) divided the family Caprellidae into four subfamilies. These subfamilies, based extensively on super- ficial characters (e.g. pereopods 3 and 4, number of gills), are here considered to be unacceptable. McCain (1970) revised Vassilenko’s subfamilies on the basis of mandibular type, determined by the presence or absence of the molar process and the palp. The presence or absence of the mandibular molar is certainly more basic than the characters used by Vassilenko. However, other mouth- parts exhibit characters that may be of a more fundamental nature than the presence or absence of the mandibular palp. Thus, the number of spines on maxilla 1 outer plate is six in some genera, seven in others; also, the outer plate of the maxilliped sometimes bears a row of spines along its inner edge. These characters are here considered of higher systema- tic importance than the mandibular palp. No further decision on the classification of the Caprellidae can be reached until the nature of the mouthparts of as many genera as possible can be determined. Because of the un- satisfactory nature of the systems so far presented, the caprellids under review here have not been placed in sub- familes. If, however, one uses McCain's familial taxa, the genera in this paper can be assigned to families and sub- families as follows: Caprogammaridae Kudrjaschov and Vassilenko 1966, emend McCain 1970: Cercops. Aeginellidae Vassilenko 1968, emend McCain 1970: Aeginellinae Vassilenko 1968, emend McCain 1970: Aeginel/a, Aeginina, Proaeginina; Protellinae McCain 1970: Luconacia, Mayerella. Caprellidae White 1847, emend McCain 1970: Caprell/a, Hemiaegina, Paracaprella. Methodology A special terminology has evolved around the Caprellidae, and note must be made of certain usages. The swimming setae on antenna 2 are the many long, paired hairs set along the lower edge of the peduncle and, frequently, the flagellum of this antenna; these hairs are actually used in feeding rather than in swimming. Notes on the Family Caprellidae The inner and outer plates of the maxillae and the maxilliped have frequently been referred to as lobes. | have used the term plate, which is the standard amphipod terminology (e.g. J.L. Barnard 1969: 522). The pereopods and gnathopods are numbered according to the pereonite from which they arise. Because it was originally believed that the coxal segments were absent, the appendages have long been treated as six-segmented, the basis being segment one and the dactylus being segment six. This numbering is used here, despite the fact that coxae are found in caprellids, particularly on the gnathopods and pereopods 5 to 7. A decision concerning a reversion to normal amphipod terminology will be made when all the caprellid genera have been reviewed. All figures are drawn to scale, using a microprojector; unless otherwise stated, all scales shown equal 1 mm, and refer to the whole-mount only. The male and female whole- mount are to the same scale unless otherwise indicated. Measurements were made laterally along the midline of each pereonite, from the head, between the insertions of antennae 1 and 2, to the tip of the abdomen. All characters used in the Key refer to both males and females, unless specifically stated otherwise. Those species that were described in detail by McCain (1968) have been treated only generally; for details refer to the above paper. The present location of the specimens examined has been indicated by the following abbreviations: BM British Museum of Natural History, London FRBC Fisheries Research Board of Canada, Arctic Unit, Ste-Anne-de-Bellevue, Quebec GR Station de Biologie marine, Grande-Riviere, Quebec MN Muséum National d‘Histoire Naturelle, Paris NMC National Museum of Natural Sciences, National Museums of Canada, Ottawa NR Naturhistoriska Riksmuseet, Stockholm USNM United States National Museum, Washington, D.C. UZM __— Universitetets Zoologiske Museum, Copenhagen ZM Zoologisk Museum, Oslo Detailed collection data have been given only for the less common species. The predominantly intertidal and shallow- water collections from eastern Canada have been summarized by region, and by province within each region. Where used, station numbers refer to published station data (Bousfield 19566-1962). Other major National Museums of Canada collections, for which data is unpublished, came from the St. Lawrence estuary, 1953 (NMC 53-52); the Gulf of St. Lawrence, 1960 (NMC 60-180); eastern Nova Scotia, 1962 (NMC 62-116); western Nova Scotia, 1963 (NMC 63-293); 14 Methodology the St. Lawrence estuary, north shore, 1969 (NMC 69-211). Also deposited in NMC is a collection, gathered mainly in the Bay of Fundy region, 1911-1916, from the Atlantic Biological Station, St. Andrews, New Brunswick (NMC 51-15). Information for Maps 1, 3-5, 7, 8, and 10 is based on literature records, and on the material newly examined here. Information for Maps 2, 6, and 9 is based mainly on the published and unpublished data of Dr. E.L. Bousfield. 15 Map 1 - f\ @ Aeginella spinosa v Aeginina aenigmatica Known distribution of Aeginel/la spinosa and Aeginina aenigmatica north of 45°N. 16 Key to the Caprellidae of Atlantic and Arctic Canada Abdomen __ five-segmented; mandibular molar eM ae oe cin aia pu ecidpraen nsesle days Wane Wests COMUNRT Abdomen unsegmented; mandibular molar WN PRIIN 622i arcide cs semaine Warde cent tye dticaradD> 22 Fe aes AA Lower lip Mandibular palp Male, abdomen Female, abdomen Right mandible Left mandible Aeginina longicornis Gills elliptical. Pereopods 5 to 7 increasing in length; segments 1 and 3 com- paratively long, segment 4 short, propodus with proximal pair of grasping spines. Abdomen with one pair of biartic- ulate appendages bearing many minute knobs on their medial sur- face; one pair of unsegmented ap- pendages with an apical row of denticulations and an apical seta; and one pair of lobes. Penes lateral and short. Female: Essentially similar to male; abdomen as in male, but lobes setose. Distribution Type locality: Henry Land, eastern Greenland, depth 40 m, 21 July 1900. One male holotype, 1 female allotype, 2 mate and 3 female para- types (UZM). Remarks The largest male found was 10 mm, the largest female 9.4 mm. This species was based on seven speci- mens, all of which were so immature that there was no sign of develop- ment of the secondary sexual char- acters. As there was some doubt that they might be immature speci- mens of A. /ongicornis, they were compared with specimens of this species at an approximately similar developmental stage, and_ their distinctiveness was apparent. Apart from differences in antenna 1, gnathopod 2, and the pereopods, the abdomen shows slight differences; the most obvious is that in A. aenig- matica the penes are half as long as the unsegmented appendages in the immature specimens available, where- as in A. /ongicornis they are of the same length as the appendages in both immature and adult specimens. Aeginina longicornis (Kr¢yer 1842-43) Plate 3; Maps 2, 3 Synonymy Refer to McCain 1968. Also: [?] Caprella scolopendroides (non Lamarck) — Ross 1826 Aegina spinifera — Rodger 1894 Aegina spinosissima — G.O. Sars 1909 Aeginina longicornis — Vibe 1950 — Steele 1961 — Brunel 1970 Material examined Approximately 3,000 specimens. Hudson River estuary, off New Jersey, 12 specimens (NMC). Atlantic Canada: Bay of Fundy, 600 specimens (NMC); Nova Scotia, 100 specimens (NMC); off Newfoundland, 18 specimens (NMC). Gulf of St. Lawrence: Nova Scotia, 3 specimens (NMC); Prince Edward Island, 7O specimens (NMC); New Brunswick, 12 specimens (NMC); Gaspé, 2,100 specimens (GR, NMC). St. Lawrence River estuary: north shore, 8 specimens (NMC); Saguenay fjord, 27 specimens (GR). Arctic Canada, west to 106°20’, north to 76°46’, 65 specimens (FRBC). Greenland, western and southern, 30 specimens (UZM). Description Body smooth to very spinose, one pair of dorsal head spines and lateral spines over gnathopod 2 insertions, usually present. Length of largest male 50 mm, of largest female 30 mm, of smallest ovigerous female 7.7mm. 23 Plate 3 Aeginina longicornis Antenna 1 in adult males usually as long as body. Antenna 2 shorter than peduncle of antenna 1. Mouthparts typical of genus; setation of terminal article of man- dibular palp shows some variation. Gnathopod 1 propodus triangular, grasping margin of propodus and dactylus slightly serrate. Gnathopod 2 basis with strong anterolateral spine; propodus slender, and with triangular projection anterodistally; palm with proximal — projection bearing single spine, and with two distal projections separated by a cleft. In large adult males, both the distal cleft, and the notch distal to the grasping spine become much exag- gerated (Plate 3, fig. E). Gills elliptical. Pereopods 5 to 7 of increasing length, propodus being the longest segment and bearing proximal grasping spines. Abdomen typical of genus. Penes lateral, medially directed, and long. Distribution Type locality: Near Greenland. Frederikshaab, Other localities: Arctic Ocean from 140°E west to Baffin Bay; southern Norway; Denmark; Faeroe Islands; Shetland Isles; Iceland; southern Plate 3 Aeginina longicornis A Female B Mandibular palp C Male D- Right lacinia mobilis E Male, gnathopod 2 propodus F Pereopod 7 propodus G Male, abdomen H Female, abdomen Greenland; Newfoundland south to North Carolina. New records: Saguenay fjord and north shore of St. Lawrence estuary; various localities among the Cana- dian arctic islands from Baffin Bay west to 106°20'W. Remarks This species shows. considerable variation in both size and spination. It has not been possible to ascribe reasons for these variations, although McCain found that in general speci- mens from northern waters were more spinose than those from south- ern waters. Two female specimens were found to be _ parasitized by nematode worms; in one the worm was situated within pereonites 3 and 4, and in the other the worm was inside one of the brood plates. One_ intersexual specimen was found with developing brood plates, female genital opening, and one penis. This is a common North Atlantic and Arctic species. Although it is fre- quently found in deeper water (to 2258 m), nearly half of the Canadian Atlantic collections were from shal- low (less than 10 m) water. J-N Immature: J Male K Mandibular palp L Male, abdomen M Female, abdomen N Male, gnathopod 2 propodus 25 Caprella andreae Map 2 Distribution of Paracaprella tenuis and Aeginina longicornis within the Canadian Atlantic region; based on materia! examined. Genus CAPRELLA Lamarck 1801 Antenna 2 usually with swimming setae, flagellum with two articles; man- dibular palp absent, molar present; outer plate of maxilliped larger than, or equal to, inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 absent, pereopod 5 six-segmented; abdomen of male with one pair of appendages and one pair of lobes, abdomen of female with one pair of lobes. Type species Cancer linearis Linnaeus 1767 (designated by Dougherty and Steinberg 1953). Caprella andreae Mayer 1890 Synonymy Refer to McCain 1968. Remarks This species has not been reported north of Cape Cod. It appears to be pelagic in habit, being attached to floating objects, and it is therefore possible that it will be found in the region under consideration. C. andreae is generally similar to 26 C. penantis, differing most obviously in having a convex palm, with medial grasping spines, on the propodus of pereopods 5 to 7. For detailed description, McCain 1968. consult Caprella carina Mayer 1903 Plate 4; Map 4 Synonymy Caprella carina — dA. Thompson 1901 Caprella carina Map 3 Caprella carina Mayer 1903 — Stephensen 1933 Caprella sedovi Gurjanova 1933, 1935 Material examined Cape Parry, N.W.T., 7O°O7'N, 124° 39°;W, 23 Aug. 1962, depth 5.5m, 3 males; 7 Sept. 1962, depth 7 m, 4 males (FRBC). Point Barrow, Alaska: Eluitkak Passage, Elson Lagoon, 8 June 1948, 3 females; beach at Point Barrow Base, 28 Sept. 1950, 11 males, 10 females (USNM). Known distribution of Aeginina longicornis north of 45°N. Kara Sea, 70°40'N, 64°17°E, depth 20 m, female lectotype (NR). Slide mounts deposited in NMC. Description Head and body without dorsal spines, but with many large and small tuber- cles, particularly on dorsal surface; lateral spines present over gnathopod 2 insertions, over gills, and anteriorly on pereonite 3. Length of largest male 34.4 mm, of largest female 19.8 mm. Antenna 1 equal in length to cephalon plus pereonite 2 in male, 27 Plate 4 28 Caprella carina longer in female; flagellum of up to 19 articles, shorter than peduncle. Antenna 2 longer than peduncle of antenna 1, with strong swimming setae. Mouthparts typical of Caprel/l/a. Left mandible with five-toothed in- cisor, five-toothed lacinia mobilis, setal row of three (occasionally more) setae, molar with single plumose seta. Right mandible with five- toothed incisor, lacinia mobilis toothed but not five-toothed, setal row of two setae, molar with single plumose seta. Maxilla 1 outer plate with seven complex apical spines; palp with apical spines and many setae. Maxilla 2 plates both setose apically. Maxilliped inner plate con- cave apically, with four spines and many plumose and simple setae; outer plate with two large apical setae, row of spines along medial margin, and numerous’ setae on surface; articles of palp strongly setose, dactylus slender and bearing numerous comb setae. Gnathopod 1 propodus triangular, with one pair of proximal grasping spines; grasping margin of propodus and dactylus_ finely denticulate. Gnathopod 2 basis short, with lateral and medial anterior spines; ischium with lateral and medial anterior spines; propodus with slight antero- Plate 4 Caprella carina Female (posterior carinations not shown) Gnathopod 1 propodus Female lectotype, gnathopod 2 Male, gnathopod 2 Male Maxilla 1 Maxilla 2 QOummMoOdOWD>yYD distal projection, tuberculate anterior- ly, palm laterally setose in male, bearing proximal projection with single grasping spine, accessory spine present in female, and two triangular distal projections; dactylus strong, evenly tapered, in male denticulate along grasping margin. Gills elliptical. Pereopods 5 to 7 of increasing length, propodus' with proximal grasping spines. Abdomen of male with one pair of unsegmented appendages, and one pair of setose lobes separated by a median chitinous double hump; penes medial. Abdomen of female with one pair of setose lobes sep- arated by chitinous hump. Distribution Type locality: Kara Sea (70°40’'N, 64°17’E), depth 20 m. Other localities: Kara Sea; Disko, western Greenland. New records: Cape Parry, N.W.T.; Point Barrow, Alaska (71°22’N, 156°30'W). Remarks Mayer based this species on two female specimens. The specimen deposited in the Stockholm Museum Maxilliped Pereopod 6 Left mandible Male, abdomen Female, abdomen Lower lip Right mandible On Se rixtt 29 Plate 5 Caprella ciliata Female, gnathopod 2 Male lectotype, gnathopod 2 propodus Female Gnathopod 1 propodus Male, abdomen Pereopod 7 ™MONQWDYD 30 ACID Male lectotype Female, abdomen Right mandible, incisor region Maxilliped palp, terminal articles (setation not shown) Caprella ciliata (No. 2336) has been designated as lectotype. Laubitz (1970) noted that in C. a/as- kana Mayer, the mandibular setal row usually contained extra setae. Extra setae were observed in C. carina, but it was not possible to determine whether this condition is normal for this species. Caprella ciliata G.O. Sars 1883 Plate 5; Map 5 Synonymy Caprella ciliata G.O. Sars 1883, 1894 — Norman 1886, 1905 — Mayer 1890, 1903 — Holmes 1904 — Reibisch 1906 — Stephensen 1927, 1928, 1929a, 1940, 1942, 1944a — Oldevig 1933 — Enequist 1949. Material examined Thor Station 166, 62°57'N, 19°58'W, 14 July 1903, depth 957 m, 1 male, 5 (2 ovigerous) females (UZM). Ingolf Station 92, 64°44'N, 32°52’W 25 June 1896, depth 1838 m, 1 fe- male (UZM). Western Norway, 17 specimens, including Sars’s type material and slide mount (ZM). Description Body normally smooth; when dorsal spines present, there may be paired or single head spine, single posterior spine on pereonite 1, paired spines in the middle and a single spine posteriorly on pereonite 2. Pereonite 5 comparatively long, particularly in female. Length of largest male 13 mm, of largest female 9.1 mm, of smallest ovigerous female 8.8 mm. Mouthparts apparently typical of genus; maxilliped palp with dactylus longer than article proximal to it; right mandible lacinia mobilis denti- culate. not five-toothed. Gnathopod 1 propodus slender, triangular, with one pair of proximal grasping spines; not very setose; grasping margin of propodus and dactylus serrate. Gnathopod 2 pro- podus_ slightly setose in female, medial surface strongly setose in male; palm with proximal projection bearing grasping spine, and median and distal projections separated by a more or less deep cleft; dactylus of male setose, heavy. Gills small, elliptical. Pereopods 5 to 7 of increasing size, propodus with proximal pair of long slender grasping spines. Distribution Type locality: Western Norway. Other localities: Denmark; Sweden; Iceland; Alaska. Remarks Our present knowledge of the distri- bution of this species suggests that it may be present in the region covered by this paper; but since it has not yet been reported from this area, it has been treated only generally. For further details of this species, refer to Sars (1883, 1894) and Mayer (1903). A lectotype has been designated from Sars’s material deposited in the Oslo Museum (No. F 12537a). Caprella dubia Hansen 1888 Plate 6; Map 4 Synonymy Caprella microtuberculata f. spinosa — Norman 1886 Caprella microtuberculata f. spinigera Hansen 1888 — M.J. Grieg 1907 Caprella dubia Hansen 1888 — Vanhoffen 1897 — Scott 1899 — d’‘A. Thompson 1901 — Mayer 1903 — Stephensen 1933, 1942, 19445 — 31 32 Plate 6 Re RD e) XO ia. Caprella dubia Gurjanova 1935, 1936, 1964 Caprella septentrionalis f. spinigera — Mayer 1890 — Hansen 1895 Material examined Dana Station 2361, 68°O8'N, 57°30’ W, 26 June 1925, depth 398 m, 1 male, 1 female (UZM); 65°34'N, 54°31 W, 5 July 1895, depth 128 m, 1 male, 3 (1 ovigerous) females (UZM). Frobisher Bay, Baffin Island, 8 Oct. 1968, depth 62 m, more than 100 specimens, all immature (FRBC). 44°30'N, 49°OO’W, 25 June 1961, depth 120 m, 3 males, 1 female (NMC). Hansen’s cotypes: 66°32'N, 55°54’ W, 1884, depth 200 m, 2 males; 65°35'N, 54°50'W, 1884, depth 150 m, 1 male, 1 ovigerous female; 65°35'N, 54°50'W, 1884, depth 150 m, 2 males, 2 females (UZM). Slide mount deposited in UZM. Description Body spination varies slightly, as shown in male and female (Plate 6). In male, pereonite 1 and anterior portion of pereonite 2 elongate with maturity. Length of largest male 26.1 mm, of largest female 22 mm, of smallest ovigerous female 16.5 mm. Antenna 1 longer than head to pereonite 3 inclusive, flagellum with more than 26 articles and longer than peduncle; peduncle articles 2 and 3 setose in male. Antenna 2 with many Plate 6 Caprella dubia A Female B Female, gnathopod 2 propodus C Gnathopod 1 propodus D Male, gnathopod 2 propodus E Male lectotype F Maxilla 2 G Maxilliped strong swimming setae, article 2 minute. Mouthparts typical of genus; right mandible lacinia mobilis strongly toothed, but not five-toothed. Gnathopod 1 propodus triangular, with one pair of proximal grasping spines; dactylus slightly longer than propodus palm; grasping margin of dactylus and_ propodus _ serrate. Gnathopod 2 basis and ischium with distal anterolateral spine; propodus slender, palm setose, with proximal projection with spine, and two distal projections; dactylus thickened prox- imally, strongly curved, with setae along the inner margin in adult and subadult males. Gills long, oval. Pereopods 5 to 7 with proximal pair of grasping spines on the propo- dus, and moderately short dactylus. Abdomen of male and _ female typical of genus. flagellum Distribution Type locality: 65°35'N, depth 150 m. 54°50'W, Other localities: Eastern and western Greenland; Franz Joseph Land; Kara Sea. New records: Off Newfoundland; Baffin Island. Remarks A lectotype has been designated Maxilla 1 Lower lip Pereopod 7 Left mandible Male, abdomen Female, abdomen Right mandible S22 wor 33 Caprella equilibra 5 L : sf Vi ©? ° x > (ae A\ M@ Caprella carina @ C. dubia Vv M. “iA 130° N i Map 4 Known distribution of Caprel/la carina, Caprella dubia, and Metacap- rella horrida. from Hansen’‘s cotypes deposited in the Copenhagen Museum. C. dubia is very similar to C. micro- tuberculata G.O. Sars; for further discussion, see under the latter species, p. 39. Caprella equilibra Say 1818 Synonymy Refer to McCain 1968 Remarks This species is widely distributed in 34 the warm temperate waters of the world. Although it has not been re- corded from the area covered by this paper, it is common along the eastern coast of North America from Florida to Connecticut. It has also been re- corded from the Mount Desert region of Maine (Procter 1933), and in Europe is found at least as far north as Trondheimsfjord, Norway. C. equilibra is easily distinguished from the other Capre//a species of the northwestern Atlantic by the pres- ence of a single ventral spine be- Caprella linearis 1 @ Capreila ciliata MSOC. «*srinki VW Proaeginina norvegica Ve ip. \ Map 5 Known distribution of Caprel/la ciliata, Caprella rinki, and Proaeginina norvegica. tween the gnathopods 2. In the adult male the anterior pereonites are so elongated that the gnathopods 2 arise at mid-length of the body. For detailed description and figures, refer to McCain 1968. Caprella linearis (Linnaeus 1767) Plate 7; Maps 6, 7 Synonymy Refer to McCain 1968. Also: Caprella acuminifera (non Latreille) Bate 1862 — Mayer 1882 Caprella hystrix (non Krgéyer) Bate and Westwood 1868 Caprella septentrionalis (non Kr@yer) — Bousfield 19566 (in part) Caprella acutifrons (non Latreille) — Bousfield 1958 (in part) Caprella linearis — Steele 1961 — Brunel 1970 Material examined Approximately 1300 specimens. Hudson River estuary, off New Jersey, 11 specimens (NMC). Atlantic Canada: Bay of Fundy, 360 35 36 Plate 7 Caprella linearis A Male B Maxilliped palp, terminal articles (setation not shown) C Female Caprella linearis specimens (NMC); Nova Scotia, 60 specimens (NMC); off Nova Scotia, 44°02'N, 59°12'W, depth 180-360 m, 1 female (NMC). Gulf of St. Lawrence: Prince Edward Island, 25 specimens (NMC); New Brunswick, 8 specimens (NMC); Gaspé, 800 specimens (GR, NMC). St. Lawrence estuary, south shore, 30 specimens (NMC). ; Alaska: Ivik, 2 Aug. 1951, 7 juveniles (USNM); Elson Lagoon, Point Barrow, 6-20 Oct. 1948, depth 1.5-2 m, 16 juveniles (USNM); off Point Barrow, Aug. and Sept. 1948, depth 25-45 m, 125 specimens (USNM). BM specimens (1952.5.7.163) from Bate’s collection: These were labelled “Caprella acuminifera”, Bate’s original designation of C. hystrix Bate and Westwood. Hoeck (1879) suggested that C. Aystrix was a variety of C. /inearis. Mayer reverted to the use of C. acuminifera, and assigned this species to C. septen- trionalis. The specimens agree well with Bate’s original descriptions and figures, and are here assigned to C. linearis. Description Body usually smooth; tuberculations always paired when present. In adult males pereonites 1 and 2 considerably elongated. Length of largest male 22.2 mm, of largest female 13.1 mm, of smallest ovigerous female 6.2 mm. Male antenna 1 with’ setose peduncle and short flagellum. Anten- na 2 shorter than antenna 1 pedun- cular articles 1 and 2. Mouthparts typical of genus. Gnathopod 2 of male with basis longer than propodus; propodus palm with proximal projection bearing a spine and with one accessory spine, and two distal projections separated by a deep cleft; dactylus strongly curved and setose. In female, distal projections are smaller and closer together; dactylus not setose. Gills elliptical. Propodus of pereopods 5 to 7 with proximal grasping spines. Variation: Typically this species has a smooth body with usually one pair of dorsal spines on each of pereonites 5 to 7. A small number of markedly tuberculate specimens were found. These cannot be considered as a separate species, since intergrades between the two forms exist. The specimens figured (Plate 7) were of the intermediate type. Distribution Type locality: “‘Habitat in Oceano Europaeo”’ (Linnaeus 1767). Other localities: Siberian Polar Sea; Murmansk coast; Spitsbergen; Eu- ropean coast from Norway to France; British Isles; Faeroe Islands; Iceland; east coast of North America from Labrador to Connecticut; southwest Alaska; ? Japan, Kamchatka, and New Zealand. New records: Hudson River estuary; Bay of Fundy; Gulf of St. Lawrence; Point Barrow and lvik, Alaska. Remarks No explanation can be offered for the variation in body spination; such variation, shown by a number of caprellids, is possibly of ecological significance. There does not appear to be any geographical isolation of the variants, although it was noted that all the Alaskan specimens were strongly tuberculate. The C. Aystrix of Bate and Westwood, from the British Museum, were examined, and it was apparent that these are actually 37 Plate 8 38 Caprella microtuberculata strongly tuberculate C. /inearis. This common boreal species is mainly intertidal, and only a few collections of it were taken below 25 m. For further discussion of C. /inearis, see C. septentrionalis, p. 48. Caprella microtuberculata G.O. Sars 1879 Plate 8; Map 8 Synonymy Caprella microtuberculata G.O. Sars 1879, 1885, 1886, 1894 — Mayer 1882, 1890, 1903 — Vanhoffen 1897 — Scott 1899 — Stebbing 1900 — Bruggen 1909 — Stephensen 1918, 1929b, 1942, 1944a, 19445 — Derjugin 1915 — Gurjanova 1931, 1964 — McCain 1966 Material examined More than 75 specimens from east- ern Greenland eastward. Ingolf Station 2, depth 493 m, 3 males, 1 female; Ingolf Station 3, depth 512 m, 14 males, 20 females, many immatures and juveniles; In- golf Station 44, depth 1026 m, 1 male; Ingolf Station 93, depth 1444 m, 1 female; Ingolf Station 144, depth 520 m, 1 ovigerous female. Eastern Greenland, 65°54'N, 1902, 1 female; Henry Land, 1900, depth 40 m, 3 males, 14 females, 25 imma- tures; 72°35 N,. 19°35'W, 1891, depth 200 m, 1 male, 1 female; Plate 8 Caprella microtuberculata Female Gnathopod 1 propodus Male, gnathopod 2 propodus Female, gnathopod 2 propodus Maxilla 2 Maxilla 1 Maxilliped QOm7mMoOODD Finnmark, Norway, 1898, 2 males, 1 female. All specimens in UZM. Between Bear Island and Spits- bergen, 6 specimens, Sars’s type material (ZM). Slide mounts deposited in UZM. Description Body tuberculation variable; large adult males have paired head tuber- cles and dorsal tubercles on pere- onites 5 to 7 only; the most tuber- culate specimen observed’ was similar to the female in Plate 8, with the addition of a posterior pair of tubercles on pereonite 1. Length of largest male 30.1 mm, of largest female 21 mm, of smallest ovigerous female 20.5 mm. Antenna 1 flagellum longer than peduncle and with more than 28 articles; peduncle articles 2 and 3 setose in male. Antenna 2 shorter than peduncle of antenna 1, long swimming setae, flagellum article 2 minute. Mouthparts typical of genus, right mandible lacinia mobilis toothed, but not five-toothed. Gnathopod 1 propodus with one pair of proximal grasping spines; grasping margin of dactylus and propodus serrate. Gnathopod 2 pro- podus slender, palm with proximal projection bearing a spine, and with one or two accessory spines, and two distal projections; in the male, the more distal of these is much the Pereopod 7 Male Male, abdomen Female, abdomen Right mandible Left mandible 22S nA Cr 39 Caprella microtuberculata C. linearis e@ C. penantis a Map 6 Distribution of Caprella linearis and Caprella penantis within the Canadian Atlantic region; based on material examined. larger, but in the female they are the same size, and appear to have a com- mon point of origin; dactylus evenly tapered, may be setose in large adult males. Gills elliptical. Pereopods 5 to 7 of increasing length; propodus with proximal pair of grasping spines; dactylus long and slender. Abdomen of male with one pair of unsegmented appendages, and one pair of setose lobes separated by a median chitinous hump, penes medial. Abdomen of female with one pair of lobes separated by a median chitinous hump. Distribution Type locality: Northwest of Bear Island, and southern tip of Spits- bergen, depth 70-180 fm (128-329 m). Other localities: Eastern Greenland; northern Norway; Spitsbergen; Franz Joseph Land; Kara Sea; north 40 of New Siberian Islands. New record: Faeroe Islands (62°49'N, 7°12 Wi). Remarks A lectotype has been designated from Sars’s cotypes deposited in the Oslo Museum. The female lectotype (No. F 12317a) was from a vial labelled with both type localities; the para- lectotypes are numbered F 1231 7b-e, and F 1941. C. microtuberculata and C. dubia are very similar in both general body shape and the location of the body spines. However, the differences between them are_ sufficient to separate them as full species. The abdomen of C. microtuberculata is very much more obvious than that of C. dubia. Also, in C. microtuberculata the spine row on the maxilliped outer lobe consists of about five spines, the gnathopod 1 dactylus just reaches the end of the propodus palm, the Caprella penantis Map 7 Known distribution of Capre//a linearis within the North Atlantic and northern Pacific boreal and subarctic regions. pereopod dactylus is long and slender. In C. dubia the maxilliped outer lobe spine row consists of about eight spines, the gnathopod 1 dactylus reaches beyond the end of the pro- podus palm, the pereopod dactylus is fairly short. Hansen (1888) originally assigned C. dubia to C. microtuberculata, giving it the varietal name sp/nigera, but further on in the same paper (p. 217), he gave it specific status as C. dubia on the basis of its greater size and slight differences in the gnathopods 2. It should be em- phasized that both species may have setae on the dactylus of gnathopod 2. Caprella penantis Leach 1814 Plate 9, figs. A-E; Plate 10, figs. A-E; Map 6 Synonymy Refer to McCain 1968 Material examined About 400 specimens. Atlantic Canada: northeastern Nova 41 Plate 9 A-E Caprella penantis A Male, from Florida B Male, from Cape Cod C Male, from Prince Edward Island D Male, abdomen E Female, abdomen 42 F,G Caprella natalensis F Male, abdomen G Female, abdomen H-K Caprella neglecta H Female J Male K Female, gnathopod 2 propodus Caprella penantis Scotia, 25 specimens (NMC). Gulf of St. Lawrence: files de la Madeleine, 5 specimens (GR, NMC); Prince Edward Island, 300 specimens (NMC); Nova Scotia, 10 specimens (NMC); New Brunswick, 4 speci- mens (NMC). New England south to Florida, many specimens. The greatest depth recorded in these collections was 11 m. Description Head with large, triangular, anteriorly directed projection anterior to eye. Pleura present on pereonites 3 and 4 in mature specimens. Pereonite 1 normally shorter than head, pereonite 5 normally shorter than pereonites 6 plus 7. Length of largest male 9.5 mm, of largest female 5.9 mm, of smallest ovigerous female 3.5 mm. Antenna 2 longer than peduncle of antenna 1. Mouthparts typical of genus. Gnathopod 2 propodus palm setose, with large rectangular projection distally, and large proximal pro- jection. Gills circular, oval, or elliptical. Pereopods 5 to 7 of increasing length, propodus with concave palm and proximal grasping spines. Abdomen typical of genus. Variation: It was found that there is considerable variation within this species, particularly in the robustness of the body. Specimens from Florida were larger, stouter, with stronger pleural development and more setose antenna 2, than specimens from Prince Edward Island; intergradations between these two forms were found (Plate 9, figs. A-C). Most Crustacea spanning this geographical range grow to larger size in the north than in the south. No satis- factory explanation has been found for the peculiar size range of C. penantis. Distribution Type locality: Devonshire, England. Other localities: Atlantic coasts of Europe, British Isles, North and South America; Gulf of Mexico; Caribbean region; California; Hawaii; eastern Asia; New Zealand; Australia; South Africa. Remarks The Caprella acutifrons group, en- compassing the twenty varieties named by Mayer (1890, 1903), has been analysed by McCain (1968). He tentatively assigned eight of the varieties, plus C. angusta (Mayer 1903) to the species C. penantis. Of the eight varieties, five (C. acutifrons f. carolinensis, virginia, testudo, gibbosa, and Jusitanica) have the setose appendages (particularly the gnathopod 2 propodus) characteristic of C. penantis. These five are con- sidered to have been rightly assigned to this species. C. angusta (which has been considered the Pacific coast equivalent of C. penantis) and C. acutifrons f. natalensis, porcellio, and neglecta are not setose. As no other difference was observed be- tween them and C. penantis, it was suggested that they might, as a group, be given subspecific status. Vassilenko (1967) gave C. neglecta specific status, but since her specimens ap- pear to have been typical setose C. penantis, this species is considered invalid. Laubitz (1970) concluded that C. angusta was sufficiently different from C. penantis to be con- sidered a separate species, and re- instated it. Collections of C. penantis from the 43 HOAR HOOD D DONA Male, gnathopod 2 propodus A-E Caprella penantis, total length H 8mm (scale =0.5 mm) ae ae : J 11.1 mm (scale =0.5 mm) of individual: K 12.6mm (scale =0.5 mm) a ae ii i Ae: ee L-P Caprella neglecta, total length 6.1 mm (scale =0.5 mm) of individual: 8 mm (scale = 0.5 mm) 5 mm (scale = 0.5 mm) 9.5 mm (scale = 0.5 mm) 7.5 mm (scale = 0.5 mm) F-K Caprella natalensis, total length ieee aT Rea uh ree eee 12.2 mm (scale = 1mm of individual: { 14.7 mm (scale = 1 mm) F 4.9mm (scale =0.2 mm) G 6.2mm (scale =0.5 mm) mOOWD Uw @O.2—, 44 Caprella penantis Gulf of St. Lawrence south to Florida, and of C. angusta from the Canadian Pacific, have been investigated in great detail. The results confirm that these are indeed two separate spe- cies. The correct determination of these species was complicated by the discovery of the true C. penantis along the Pacific coast of California, which made it extremely difficult to determine the true identity of pre- viously published records. It must now-be pointed out that the specimens referred by Mayer (1903) to C. acutifrons f. angusta were from California, and were essentially simi- lar to the C. penantis from this same area. Therefore C. angusta must fall as a synonym of C. penantis, and the Canadian Pacific “acutifrons” species misidentified as C. angusta by Laubitz (1970) will have to be renamed. As specimens of C. acutifrons f. nata- lensis were also examined by me and found to be essentially similar to the false C. angusta,* it has been decided that the varietal name should be used, and C. natalensis (Mayer 1903) is herewith given specific status. Our present knowledge of the distribution of C. nata/ensis is limited to the south Atlantic and north Pacific. It is therefore highly probable that this name will eventually prove to be a junior synonym for either C. spinifrons Nicolet 1849, reported from Chile, or C. novae-zealandiae Kirk 1878, from New Zealand. Until specimens from either of these areas can be examined, no further decision can be made as to the correct name for this species. As has already been mentioned, * Slight differences between these two ‘‘varieties” were found. It is felt that, in the light of the known variability of the “acutifrons” group of species, these small differences should not be considered of a specific nature. C. penantis and C. nata/ensis are very similar in general appearance. Other- wise, C. penantis tends to be stouter than C. nata/ensis and, particularly in mature specimens, to have very obvious pleural development not present in C. natalensis. The most useful characters in distinguishing these two species are the gnathopod 2 propodus palm, and the ratio of the length of pereonite 5 to pereonites 6 plus 7. It was found that the gnathopod 2 differs in all stages of male develop- ment (Plate 10), and that differences are also apparent in the female gnathopods. In C. penantis, pereonite 5 is usually shorter than pereonites 6 plus 7, whereas in C. natalensis pereonite 5 is usually longer than pereonites 6 plus 7. Small differences are also present in the abdomen of both males and females (Plate 9, figs. D-G). Specimens of C. penantis from Monterey Bay, California, are less setose and smaller than the largest specimens found in the Atlantic (8 mm as opposed to 14 mm), but they show the typical stout body and strong pleural development. They are ob- viously more setose than C. nata- /ensis, particularly the adult males. Specimens of C. acutifrons f. neglecta, as identified by Mayer, were also examined, to try to determine their specific status. Differences be- tween these specimens and both C. penantis and C. natalensis, particu- larly in the head spine and the gnathopod 2 (Plates 9, 10), indicate that this variety must be considered a valid full species. Its specific status as C. neglecta (Mayer 1890, non Vassilenko 1967) is herewith pro- posed. This species has been re- ported from Hong Kong, Japan, and Australia. Although not examined, C. acutif- 45 Plate 11 Caprella rinki, female A Lateral view B Gnathopod 2 propodus 46 Caprella natalensis rons f. porcellio appears to be simi- lar to C. natalensis, and is recorded from the same area of the south Atlantic. It is therefore tentatively assigned to this species. Caprella natalensis Mayer 1903 Plate 9, figs. F, G; Plate 10, figs. F-K Synonymy | Caprella acutifrons var. natalensis Mayer 1903 — Stephensen 1949 Caprella acutifrons var. porcellio Mayer 1903 Caprella penantis (non Leach 1814) — Stebbing 1910 — [?] Penrith and Kensley 1970a, 19706 Caprella penantis var. natalensis — K.H. Barnard 1916 Caprella angusta (non Mayer 1903) — [?] Dougherty and Steinberg 1953 — Laubitz 1970 Material exaniined Specimens from Tristan da Cunha, 23 Dec. 1937, depth 40-45 m (USNM), and from Pacific North America (USNM, NMC). This spe- cies has been described and figured by Laubitz (1970). It is recorded from South Africa (Durban, Cape Town, Cape Peninsula), South West Africa, Tristan da Cunha, and Pacific North America (Queen Charlotte Islands south to Oregon; Pacific Grove and Santa Cruz, California). It is probable that further investiga- tion will show this species to be the same as C. spinifrons Nicolet, from Chile. Caprella neglecta Mayer 1890 Plate 9, figs. H-K; Plate 10, figs. L-P Synonymy Caprella acutifrons — Mayer 1882 (in part) — Schellenberg 1938 — Edmondson and Mansfield 1948 Caprella acutifrons var. neglecta Mayer 1890, 1903 — Utinomi 1943a, 19436, 1947, [?] 1968, [?] 1969 Caprella acutifrons var. natalensis — Arimoto 1930 — Hiro 1937 Material examined Specimens from Hong Kong, 9 March 1872, Mayer’s type specimens (USNM). This species is closely similar to C. penantis, and it appears from a study of the literature that both species are present in the seas around Japan. C. neglecta of Vassilenko, and C. acutif- rons f. neglecta of Utinomi (1943c) appear to refer to C. penantis. Uti- nomi (1968, 1969) mentions other Japanese “C. acutifrons” records; it has not been possible to determine to which species these records refer. The probable distribution of C. neg- lecta includes Hong Kong, Japan, Taiwan, and Hawaii. Remarks It is apparent that the “C. acutifrons” group is still in need of further clarifi- cation. Study of specimens from all parts of its geographical range will be needed before it can be decided which species are truly valid and which are subspecies or variants. Caprella rinki Stephensen 1917 Plate 11; Map 5 Synonymy Caprella Rinki Stephensen 1917 Caprella rinki — Stephensen 1933, 1944a — McCain 1966 — Brunel 1970 — Laubitz and Mills 1972 Material examined Gulf of St. Lawrence: Laurentian channel, 10 Sept. 1959, depth 377 m, 1 female (GR); 17 August 1957, 1 female (GR). 47 Caprella sarsi Description Body with small tuberculations dorsally on pereonites 1 to 5, ven- trally on pereonites 2, 3, and 4. Un- paired spines present dorsally at mid- length of pereonites 2 to 7, and laterally at the anterior of pereonites 3 and 5. Length of largest female 10.1 mm. Antenna 1 peduncle articles 2 and 3 slightly setose. In female, antenna 2 longer than peduncle antenna 1. Mouthparts not examined. Eyes very large. Gnathopod 2 of female with large anterolateral spine on basis; propodus with proximal projection with grasp- ing spine and accessory spine, and medial and distal low projections, palm slightly setose; dactylus quite heavy. Gills slender, elliptical. Pereopods 5 to 7 with proximal pair of grasping spines on propodus. Distribution Type locality: Southern Greenland, depth 460-700 m. Other localities: 61°42'N, 9°36'W, depth 1026 m; 65°14'N, 30°39'W, depth 1416 m; 65°30'N, 55°26'W, depth 550 m; Gulf of St. Lawrence, depth 180-380 m. Remarks The two specimens of this species available to me were females in rather poor condition. They are described only generally here, as this species has been treated in detail by Laubitz and Mills (1972). Caprella sarsi Honeyman 1889 Remarks Honeyman described two species of Caprella, from a glacial boulder and 48 from two cable hauls. C. sars/ was described as follows: We have already noticed this fantastic little Crustacean on Lawson Boulder (A)... On Cable Il they abound. We have found them among the Hydroida, where they had been evidently feasting. They are of various sizes, some very small. We regard all as one species. All that we now note regarding them is: Under the microscope they are plain, without spines, and have small red eyes. They are male and female. We would name it Caprella Sarsii. Lawson Boulder (A) is described as: ‘“‘a ‘glacial boulder’ of the Nova Scotia Fishing Banks”, from a depth of 65 fathoms (119 m). C. sarsi was attached to a sponge. Cable Il was brought up on 26 Oct. 1888, after 19 years. Its location was 43°4.38'N, 66°14.3'W, at 48 fathoms (88 m), in the Bay of Fundy. The description is insufficient to identify this species, but it is Suggested that it may be C. linearis. Caprella septentrionalis Kr¢yer 1838 Plate 12; Maps 9, 10 Synonymy Refer to McCain 1968. Also: Caprella ultima Bate 1862 Caprella_ septentrionalis — Smith 1879 — G.O. Sars 1909 — Vibe 1950 — Steele 1961 — Brunel 1970 Material examined Total of 4,500 specimens. Atlantic Canada: Bay of Fundy, 2,100 specimens (NMC); Nova Scotia, 640 specimens (NMC); Newfoundland, 1 specimen (NMC); Labrador, 30 specimens (NMC). Gulf of St. Lawrence: Prince Edward Island, 33 specimens (NMC); Gaspé, 300 specimens (GR, NMC); Anticosti Island, 20 specimens (NMC); Newfoundland, 1 specimen (NMC). Caprella septentrionalis “A\ W Caprella microtuberculata HM Cercops holbolli v4 ws Map 8 Known distribution of Capre/la microtuberculata and Cercops holbolli within the North Atlantic and arctic regions. St. Lawrence estuary: north shore, 100 specimens (NMC); south shore, 330 specimens (GR, NMC). Arctic Canada: Hudson Bay, 30 specimens (NMC); Hudson _ Strait, 36 specimens (NMC); Baffin Island, 27 specimens (NMC). Greenland: western, N of 70°, 84 specimens (UZM); western, S of 70°, 560 specimens (UZM); southern, 14 specimens (UZM); eastern, 14 specimens (UZM). Jan Mayen Island, 13 specimens (UZM); Iceland, 2 specimens (UZM); Spitsbergen, 1 specimen (UZM). Bate’s specimens of C. u/tima (MN). The collections were intertidal, to a depth of 15 m. Description Body spination extremely variable, single spine or tubercle usually pres- ent on head. Length of largest male 31.4 mm, of largest female 23.6 mm, of smallest ovigerous female 6.4 mm. Antenna 1 peduncle articles 2 and 3 normally setose in male. Antenna 2 normally at least as long as pe- 49 Plate 12 (\ Tad v ( is E NS Ape Caprella septentrionalis A-C_ Males, lateral view, showing differences of body spination D Pereopod 7 E Large female 50 Caprella septentrionalis duncle of antenna 1. Mouthparts typical of genus. Gnathopod 1 propodus with one pair of proximal grasping spines; grasping margin of dactylus and pro- podus serrate. Gnathopod 2 basis short; propodus palm with proximal projection bearing a spine and with accessory spine, with distal small projection plus a large triangular projection in the angle of the dacty- lus; dactylus becomes more strongly curved with maturity. Gills elliptical, oval, or round, oc- casionally grossly inflated. Pereopods 5 to 7 with proximal pair of grasping spines on the pro- podus. Abdomen typical of genus. Variation: This species shows great variation in both body spination and size. Adult males can be recognized by the elongation of pereonites 1 and 2, by the situaiion of the gnathopod 2 posterior to mid-pereonite 2, and by the setose antenna 1. They have been found as small as 11 mm. Females and immature specimens are usually spinier than the adult males. Three variations of body spination are shown in Plate 12, and intergrades between these variations exist. There is also variation in the form of the body spines, particularly on the anterior pereonites. The spines may be slender with rounded apex, blunt with flat apex, or bifid. The _ bifid spines frequently give the impression of paired spines, particularly when on the head. An interesting condition was noted in some of the female specimens. The largest females found (Plate 12) had small, immature brood plates, while in the same lot were found ovigerous females of half the size. No explanation of these ‘super fe- males” can be given until more is known about the growth and life cycle of these animals. Distribution Type locality: Greenland. Other localities: Murmansk coast, Novaya Zemlya, White Sea, and northern Russia; Europe from Nor- way, south to France and Britain; Spitsbergen; Faeroe Islands; Jan Mayen Island; Iceland; eastern and western Greenland; Baffin Bay, Davis Strait, and the eastern coast of North America from Cumberland Sound south to Georges Bank. Remarks As has already been stated, C. septen- trionalis exhibits a variety of body spination. Many of the varieties have at one time been treated as distinct species, particularly in the immature stages. G.O. Sars (1894) treated the main variations (C. punctata, C. lovéni, and C. monocera) as species, while Mayer (1903) appeared to consider them as variants. Stephen- sen (1928-19446) treated them sometimes as species and sometimes as subspecies. McCain (1968) con- cluded that, because of the existence of intergradations between the var- ious forms, they should be con- sidered infra-subspecific variants of C. septentrionalis. During the pres- ent investigation of this species, a large number of specimens were examined. It became apparent that McCain’s so-called ‘‘monocera-like variant’ is actually a typical adult male, and that all the so-called varieties can indeed by assigned to C. septentrionalis, with the exception of C. monocera. No specimens of this variety were found. Examination of type material showed that, while 51 Plate 13 Caprella unica A Male B Maxilliped palp, terminal articles (setation not shown) C Pereopod 5 propodus D Female 52 Caprella trispinis this variety is extremely similar to C. septentrionalis, the antenna 1 is obviously different, in that it has a flagellum with at least 30 articles, which is subequal to the peduncle. Until specimens of C. monocera can be examined in greater detail and numbers, it should probably be con- sidered a valid species. C. septentrionalis and C. linearis, particularly the females and im- mature stages, are very similar in form. McCain has shown that they can readily be distinguished by the ratio of total length to length of basis of gnathopod 2 (greater than 13.0 for C. septentrionalis, less than 13.0 for C. linearis). Another distinction is that in C. /inearis the head and anterior body spines are always paired, in C. septentrionalis they are always unpaired. The literature on these two species is very confused, and McCain (1968) has sorted out their synonymies most helpfully. However, as_ has already been mentioned, C. /Aystrix Bate and Westwood 1868 (non Krgyer) is to be assigned to C. /inearis. The original specimens of C. ultima Bate 1862 were also examined and as Mayer stated (1903), these are synonymous with C. septentrionalis, not with C. eqgui/ibra as suggested by McCain. Caprella trispinis Honeyman 1889 Remarks This is the second species from cable hauls in Nova Scotia, as described by Honeyman: On Cable |, we found among the Hydroida three complete specimens of another species. These have a long spine on the back of the head curved backward, and two on the back, opposite the Che/ae, curved forward; also granulation. These are blind. They are, one male and two females. We would give this the provisional name - Caprella trispinis. Cable | was brought up on 11 July 1888, after 19 years. Its location was 44°38'N, 54°6'W, at a depth of 570 fathoms (1042.5 m), “‘on the side of the Grand Bank near the Gulf Stream”. It has not been possible to identify this species. There are a number of deep-water caprellids, such as Pro- tellina ingolfi and Caprella rinki, that have both spines and ‘‘granulations”, but they also have eyes. The identity of Caprella trispinis must therefore remain in doubt. Caprella unica Mayer 1903 Plate 13; Map 9 Synonymy Caprella linearis (non Linnaeus) — [?] Ohlin 1895 — Bousfield 1956a (in part), 19566 Caprella unica Mayer 1903 — Rath- bun 1905 — McCain 1968 Caprella grahami Wigley and Shave 1966 — Patton 1968 Material examined Hudson River estuary, off Jersey, 2 specimens (NMC). Atlantic Canada: Newfoundland, 1954, A 14, Ferryland, 2 specimens; B 8, Lord’s Cove, 4 specimens (NMC). St. Lawrence estuary: north shore, 4 August 1969, Ile Grande Basque, 1 specimen (NMC); south’ shore, 23 August 1953, La Lorraine Cove, 30 specimens, Pte. Ste-Anne-des- Monts, 20 specimens (NMC). All collections are intertidal, tem- perature range 8.8°-14.5°C. New Description Body smooth to quite spiny. Length of largest male 9.2 mm, of largest 53 Caprella unica / 60° C.unica | C. septentrionalis ® 45°n _ | 55°W Map 9 _ Distribution of Capre//a septentrionalis and Caprella unica within the Canadian Atlantic region; based on material examined. female 7.2 mm, of smallest ovigerous female 4.9 mm. Antenna 1 of male with short flagellum and setose peduncle. An- tenna 2 shorter than, or subequal to, peduncle of antenna 1. Mouthparts typical of genus; dactylus of maxillipedal palp heavy. Gnathopod 1 propodus with one pair of proximal grasping spines; grasping margin of dactylus and pro- podus serrate. Gnathopod 2 propodus similar to C. /inearis, but basis shorter and heavier. Gills elliptical. Pereopods 5 to 7 with propodus palm not delineated, grasping spines absent, few palmar spines. Abdomen typical of genus. Distribution Type 45"’W, depth 59 m. 54 locality: 40°34'°30"N, 69°50’ Other localities: Long Island Sound; southern Cape Cod and off Cape Cod; Casco Bay, Maine; St. John’s, New- foundland. New records: Hudson River estuary; north and south shores, St. Lawrence estuary; Burin Peninsula, New- foundland. Remarks Patton (1968) has shown. that C. unica appears to be an obligate commensal with starfish (Asterias spp.), and that in New England waters it shows a definite, strong prefer- ence for Asterias forbesi (Desor) over A. vulgaris Verrill. A. forbesi has not been recorded north of Maine while A. vulgaris is common in the south- ern Gulf of St. Lawrence, but is re- placed in the northern Gulf and in the estuary by Leptasterias polaris Caprella unica Map 10 Known distribution of Capre//a septentrionalis within the North Atlantic boreal and arctic regions. (Muller and Troschel). The collection data on the specimens examined is not specific, but it seems probable that in this region it is A. vulgaris and probably also L. polaris that act as host to C. unica. C. unica is very similar to C. /inearis, differing in being more robust in both body and appendages. The most distinctive features are the absence of grasping spines on the pereopods, and the heavy dactylus to the maxil- liped palp in C. unica. In C. linearis the dactylus is quite slender, and grasping spines are present on the pereopods. 55 Plate 14 Cercops holbdlli Genus CERCOPS Kroyer 1842-43 Antenna 2 without swimming setae, flagellum with two or three articles; mandible with triarticulate palp, setal formula for terminal article 3, molar absent; outer and inner plates of maxilliped very small, subequal; gills on pereonites 2, 3, and 4; pereopods 3 and 4 one-segmented, pereopod 5 six- segmented; abdomen five-segmented, with two pairs of biarticulate uropods and in male, two pairs of very rudimentary pleopods. Type species Cercops holbolli Kréyer 1842-43 (by monotypy). Cercops holbolli Kr¢éyer 1842-43 Plate 14; Map 8 Synonymy Cercops Holbolli Kréyer 1842-43, 1846 — Bate 1862 — Boeck 1871, 1872-76 — Mayer 1882, 1890, 1903 — Stuxberg 1882 — Hansen 1888 — Vanhoffen 1897 — Utinomi 1947 Cercops holboelli — Lutken 1875 — Stephensen 1918 Cercops holbolli — d'A. Thompson 1901 Cercops holbolli — Stephensen 1933, 1944b Material examined Alaska: Ivik, 2 Aug. 1951, 1 male (USNM); off Point Barrow’ Base, 9 Sept. 1948, 15 Sept. 1948, depth 33-38 m, 4 males, 1 female (USNM), Plate 14 Cercops holbolli, immature Male, gnathopod 2 propodus Gnathopod 1 Male Female Maxilliped Maxilla 1 Maxilla 2 QOmmooooapyp depth 37-38 m, 1 female (USNM). Description Head with anteriorly directed median spine and paired spines anterior to eyes. Position and size of body spines variable, dorsal unpaired spines or humps may be present on pereonites 1 to 6. Length of largest male 5.7 mm, of largest female 7.5 mm. Antenna 1 setose, just longer than head to pereonite 2; flagellum shorter than peduncle, having about 12 articles. Antenna 2 setose and shorter than peduncle of antenna 1, flagellum with two articles. Mandible with triarticulate palp having three setae on the terminal article; left mandible with six-toothed incisor and lacinia mobilis, setal row of seven setae; right mandible with Pereopod 6 Lower lip Pereopod 3 Male, abdomen Left mandible Right mandible me rR Ge 57 Plate 15 Hemiaegina minuta, male Pereonites 1 and 2, lateral view B Gnathopod 1 propodus C Pereopod 7 propodus D Abdomen > 58 Hemiaegina minuta six-toothed incisor, finely denticulate lacinia mobilis, setal row of six setae. Maxilla 1 palp with few spines and setae; outer plate with six complex spines. Maxilla 2 very small, with few apical setae on each plate. Maxilliped inner plate small, with three apical setae; outer plate small, with two apical setae; palp slender. Gnathopod 1 propodus with prox- imal, single grasping spine, and strongly serrate grasping margin; dactylus grasping margin serrate. Gnathopod 2 basis and ischium with anterolateral spine, propodus palm with proximal projection bearing a spine, and small distal projection. In adult male, palm is setose and bears two proximal and two distal pro- jections. Gills small, elliptical. Pereopods 3 and 4 with one apical and one subapical seta. Pereopods 5 to 7 normal, propodus’ grasping spines proximal. Abdomen __five-segmented; seg- ments 4 and 5 each bear one pair of biarticulate appendages with a row of fine denticulations along their length; in the male, abdominal seg- ments 1 and 2 each have a pair of small humps bearing single setae. Penes lateral. Distribution Type locality: Greenland. Other localities: Near Greenland; off Cape Raper, Baffin Island; ? Tsugaru Strait, Japan. New records: Off Point Barrow, Alaska, depth 33-38 m; Ivik, Alaska. Remarks The seven specimens examined were all immature. They extend our know- ledge of the distribution of this species to the western Arctic. The Japanese locality record for C. holbolli is questionable; it is probable that either C. compactus Laubitz 1970, or per- haps an undescribed third species of this genus, is responsible for this record. Genus HEMIAEGINA Mayer 1890 Antenna 2 without swimming setae, flagellum with two articles; mandible without palp, molar present; outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 one-segmented, pereopod 5 six-segmented; abdomen with one pair of biarticulate appendages. Type species Hemiaegina minuta Mayer 1890 (by monotypy, subsequently designated by McCain 1968). Hemiaegina minuta Mayer 1890 Plate 15 Synonymy Refer to McCain 1968. Also: Hemiaegina minuta — Utinomi 1964, 1969 — Fine 1970 Material examined One specimen from mid-Atlantic, 45°40.3'N, 36°45.5'W, 24 July 1966, depth 180 m, midwater trawl! (NMC). Description Body smooth except for one pair of 59 Hemiaegina minuta ventral spines between gnathopods 2; pereonites 2 to 6 have a hexagonal outline when viewed dorsally. Length of male, 4.7 mm. Antenna 1 subequal in length to pereonites 1 to 4. Antenna 2 just longer than peduncle of antenna 1. Left mandible with five-toothed incisor, five-toothed lacinia mobilis; right mandible with five-toothed incisor, serrate lacinia mobilis. Maxilla 1 with six spines on outer plate. Maxilliped inner and _ outer plates quite small, palp slender with tapering dactylus having’ serrate grasping margin. Gnathopod 1 propodus with large proximal knob bearing many minute projections and one small grasping spine, tip of dactylus complex;* grasping margin of propodus and dactylus not serrate. Gnathopod 2 propodus large, palm with proximal grasping spine and medial projection separated from distal bilobed pro- jection by a notch. Gills elongate. Pereopods 3 and 4 one-segmented, minute. Pereopods 5 to 7 with pro- podus bearing several proximal knobs, each with a spine; dactylus with a few strong serrations just past the centre of its grasping edge. Abdomen with one pair of biarti- culate appendages bearing rows of minute knobs along medial edge. Penes lateral. Distribution Type locality: Off Amoy, China, depth 15-46 m. Other localities: Off ©Bermuda; Virgina; North Carolina; Florida; Tortuga Island; 29°44'N, 88°23'W; * There appears to be an anterior claw and a posterior marginal lobe (Plate 15, fig. B). 60 Texas; Virgin Islands; South Africa; Hawaii; Bora Bora Island; Tateyama, Nomasaki, and Sunohana, Japan; 1°42.5'S, 130°47.5’E; Fremantle, Australia; Krudadai Island, India; coast of South Arabia. New record: Mid-Atlantic, 45°40.3'N, 36°45.5'W, depth 180 m. Remarks | The single specimen examined was a very large male in rather poor con- dition. Because the new locality was much farther north than previously recorded for this species, the speci- men was compared with H. minuta from Bermuda. Apart from being very much larger and more robust, it agrees in all essential characters with this species. It was also noted that the ventral spines were supported by a large ventral protrusion in the large male. H. minuta has not been re- corded from the region under con- sideration, but has been recorded in plankton tows and from. floating Sargassum sp. It is therefore sug- gested that the specimen recorded here was carried north within the Gulf Stream. Luconacia incerta Genus LUCONACIA Mayer 1903 Antenna 2 without swimming setae, flagellum with two articles; mandible with triarticulate palp, setal formula for terminal article 1+x+1, molar present; outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 two-segmented, pereopod 5 six-segmented; abdomen of male with one pair of appendages and one pair of lobes; abdomen of female without appendages or lobes; pereopod 5 inserted near mid-length of pereonite 5. Type species Luconacia incerta Mayer 1903 (by monotypy, subsequently designated by McCain 1968). Luconacia incerta Mayer 1903 Synonymy Refer to McCain 1968. Remarks Although this species has not been recorded north of Cape Cod, it has been taken in plankton tows and from that specimens may be found in the region covered by this paper. L. incerta can most readily be distinguished from the other caprellids of this re- gion by the fact that it has the pereo- pod 5 insertion near the mid-length of pereonite 5. For further details consult McCain 1968. Sargassum sp. It is therefore possible Genus MAYERELLA Huntsman 1915 Antenna 2 without swimming setae, flagellum with two articles; mandible with triarticulate palp, setal formula for terminal article 1, molar present; outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 two-segmented, pereopod 5 two- or three-segmented; abdomen of male with one pair of appendages and one pair of lobes, of female with one pair of lobes. Type species Mayerella limicola Huntsman 1915 (by monotypy, subsequently designated by McCain 1968). Material examined Five females from Bay of Fundy, Har- bor de Loutre, 1913, 9-18 m, (NMC). Mayerella limicola Huntsman 1915 Plate 16, figs. A-F Synonymy Mayerella limicola Huntsman 1915 — Procter 1933 — McCain 1968 — Laubitz and Mills 1972 Description Body smooth. female 4.9 mm. Length of largest 61 Plate 16 A-F Mayerella limicola, female G,H Metacaprella horrida A Abdomen G Male lectotype, lateral view B Pereopod 7 H Female, semidiagrammatic dorsal view, C_ Lateral view showing body spination D Pereopod 3 E Pereopod 5 F Gnathopod 2 62 Metacaprella horrida Antenna 1 longer than head to pereonite 2. Antenna 2 longer than peduncle of antenna 1. Mouthparts typical of genus. Gnathopod 17 propodus_ slender, with single proximal grasping spine, and serrate palm; dactylus toothed, grasping margin serrate. Gnathopod 2 basis long and slender; propodus with proximal grasping spine, smooth palm with setae; dactylus slender. Gills elliptical, anterior pair being segmented. Pereopods 6 and 7 slender, with long, slender propodus and dactylus, propodus’ without obvious palm or grasping spines. Abdomen of female with one pair of setose lobes. Distribution Type locality: St. Croix estuary, New Brunswick. Other localities: Bay of Fundy, to the larger. depth of 91 m; Mount Desert region, Pereopods 3 and 4 small, two- Maine; 44°N, 68°15’'W, depth 101 m; segmented. Pereopod 5 two- 39°54'N, 70°20'W, depth 713 m. Genus METACAPRELLA Mayer 1903 Antenna 2 with swimming setae, flagellum with two articles; mandibular palp absent, molar present; outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 absent, pereopod 5 six- segmented; abdomen of male and female with one pair of appendages and one pair of lobes. Type species Caprella kennerlyi Stimpson 1864 (designated by Dougherty and Steinberg 1953). Remarks This genus is now composed of three species, VW. anomala (Mayer), M. ken- nerlyi, and M. horrida (G.O. Sars). The genus was previously thought to be endemic to the Pacific Ocean. The addition of MW. horrida to the genus extends its range to the Arctic and North Atlantic. The only character that separates Metacaprella from Caprella is the female abdomen, which bears minute appendages in Metacaprella. The abdomina of all three species were investigated, and it was found that lobes are present in the females, in addition to the appendages. The generic definition has therefore been altered to include this character. Metacaprella horrida(G.O. Sars 1877) G.O. Sars 1885, 1886 — Hansen comb. n. 1887 — Bruggen 1909 — J.A. Grieg Plate 16, figs. G, H; Plate 17; Map 4. 1925 Caprella horrida G.O. Sars 1877 — Mayer 1882, 1890, 1903 — Hansen 1895 — Vanhoffen 1897 — dA. Synonymy Caprella_ spinosissima (non Bate 1862) Norman 1874, 1886, 1905 — Miers 1877 — Mayer 1882, 1890 — Thompson 1901 — Stephensen 1918, 1933, 1940, 1942, 19444, 19445 — 63 Plate 17 64 Metacaprella horrida Steele 1961 — Gurjanova 1964 — McCain 1966 Material examined 44°30'N, 49°00'W, 25 June 1961, depth 120 m, 3 males, 4 females (NMC); Frobisher Bay, 8 Aug. 1970, depth 70 m, 1 male (NMC). Sars’s type material: Norwegian North-Atlantic Expedition: Station 124, 66°41'N, 6°59’E, 19 June 1877, depth 640 m, more than 60 spec- imens; Station. 200, 71°25’N, 15° 41'E, 17 July 1877, depth 1134 m, more than 50 specimens; Stations 18, 48, 137, 164, 200, 343, depth 547- 1359 m, more than 60 specimens (ZM). Description Head and body strongly spined, size and number of spines variable. Large paired dorsal spines always present on head, just behind eye, and on pereonite 2 over gnathopods. Spines usually also present on head just over the eye, and at the base of an- tenna 2. Spination for least spiny and most spiny specimens shown in Plates 16 and 17. The adult male of the less spiny form shows con- siderable reduction in the number of spines on pereonites 1 and 2 (Mayer 1903, Pl. 4, fig. 20). In all adult males, pereonites 1 and 2 are elongated to nearly half of the body length. Length Plate 17 Metacaprella horrida Gnathopod 1 propodus Female Male, gnathopod 2 propodus Female, gnathopod 2 propodus Maxilla 2 Maxilla 1 Lower lip QOm7moaowvp of largest male 25 mm, of largest female 14.3 mm. Antenna 1 longer than head to pereonite 3, flagellum very much shorter than peduncle; in adult male, peduncle articles 2 and 3 very long, so that the antenna is subequal to the body length. Antenna 2 shorter than antenna 1 peduncle, and with long swimming setae. Mouthparts similar to Caprella. Right mandible lacinia mobilis strongly toothed, but not five-toothed. Gnathopod 1 propodus heavy, very setose, with one pair of proximal grasping spines; grasping margin of propodus serrate, of dactylus denti- culate. Gnathopod 2 basis with lateral distal spine; propodus of male setose, with small lateral spines and a single anterodistal projection; palm with proximal projection bearing small spine, and with median projection separated by a cleft from distal, com- plex projection in angle of dactylus; dactylus heavy, setose. Propodus of female similar to male, but not parti- cularly setose and with smaller palmar projections. In adult male, the propodus is slender and longer than the basis; the dactylus is thickened proximally, but quite slender distally. Gills elliptical. Pereopods 5 to 7 of increasing length, propodus with proximal pair of grasping spines. Maxilliped Left mandible Right mandible Pereopod 7 Subadult male Male, abdomen Female, abdomen Sse ae 65 Plate 18 Paracaprella tenuis 66 mooawp Male Right mandible Male, gnathopod 2 Female, gnathopod 2 propodus Male, abdomen Cae G)e Female, abdomen Pereopod 4 Pereopod 5 Pereopod 7 propodus Paracaprella tenuis Abdomen of male with one pair of appendages and one pair of setose lobes; abdomen of female with one pair of very small appendages and one pair of lobes. Distribution Type locality: depth 753.5 m. 62°44.5'N, 1°48'E, Other localities: Siberia; Kara Sea; Spitsbergen; Norwegian Sea; _ Ice- land; Faeroe--\Islands; Greenland; Baffin Island; Hudson Bay. New record: Off Newfoundland. Remarks This is a high arctic species, and has been reported to depths of 1359 m. The record for Newfoundland is a considerable southward extension of the known range of this species, but within the same zoogeographic region. In his description of this species, Sars (1885) noted the female ab- dominal appendages, but when Mayer proposed genus Metacaprella (1903), he made no mention that C. horrida should be included in that genus. M. horrida is readily distinguished from M. anomala and M. kennerlyi by its very strongly spined head and body, and by the enormously elongated peduncle of antenna 1 and setose gnathopod 2 of the male. A lectotype has been designated from Sars’s specimens deposited in the Zoologisk Museum, Oslo (No. F 1944a), a subadult male of the more spiny form, as described and figured by Sars (1885). Genus PARACAPRELLA Mayer 1890 Antenna 2 without swimming setae, flagellum with two articles; mandibular palp much reduced with up to three articles, setal formula for terminal article 1, molar present; outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 two-segmented, pereopod 5 six- segmented; abdomen of male with one pair of appendages and one pair of lobes; abdomen of female with one pair of lobes. Type species Paracaprella pusilla Mayer 1890 (by monotypy, subsequently designated by McCain 1968). Paracaprella tenuis Mayer 1903 Plate 18; Map 2 Synonymy Refer to McCain 1968. Material examined Hudson River estuary, off Jersey, 1 male (NMC). Gulf of St. Lawrence: Nova Scotia, 9 specimens (NMC); Prince Edward New Island, 130 specimens (NMC); New Brunswick, 13 specimens (NMC). The greatest depth recorded in these collections was 11 m. Description Body dorsally smooth, head of characteristic high-domed — shape. Laterally there is some _ pleural development on pereonites 2, 3, and 4. Length of largest male 6.8 mm, of 67 Paracaprella tenuis largest female 5.1 mm, of smallest ovigerous female 3.6 mm. Antenna 1 as long as head to pereonite 2, peduncle setose. Antenna 2 setose, shorter than peduncle an- tenna 1. Mouthparts typical of genus; man- dibular palp present but much re- duced, composed of from one to three articles, and usually having single long apical seta. Gnathopod 1 propodus with single proximal grasping spine and strongly serrate margin; dactylus toothed and with serrate grasping margin. Gnathopod 2 with slender basis having distal anterolateral projection; propodus of female slightly setose, palm serrate, grasping spine proxi- mal. Propodus of male with small proximal spine separated by serrate portion of palm from very large pro- jection with deep triangular cleft immediately distal; remaining palm more or less smooth, with slight, serrate, triangular projection in the angle of the dactylus; dactylus with proximal thickening, setose and tapering more or less evenly distal to the thickening. Gills oval. Pereopods 3 and 4 biarticulate, the terminal article being minute, both articles being setose. Pereopods 5 to 7 of increasing length, propodus with proximal grasping spines and, in pereopods 6 and 7, having a series of palmar knobs, each bearing a small spine. Abdomen of male with one pair of small unsegmented appendages and one pair of setose lobes; penes arise laterally but meet distally. Abdomen of female with one pair of setose lobes. Distribution Type locality: Woods Hole, Mass. Other localities: Southwestern Gulf of St. Lawrence, around Prince Edward Island; Pennellville, Maine; coast of North America from Cape Cod to Florida; Gulf of Mexico. Remarks McCain (1968) stated that one of the differences between P. tenuis and the very similar P. pusilla Mayer, is that setae are ‘‘not generally’ pres- ent on the dactylus of the gnathopod 2 of male P. tenuis. In the collections examined by myself, however, all mature males had a few setae on the dactylus, but these were both shorter and sparser than those of P. pusi/la. Genus PROAEGININA Stephensen 1940 Antenna 2 without swimming setae, flagellum with two articles; mandible with triarticulate palp, setal formula for terminal article 1+x+1, molar present; outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 absent, pereopod 5 six-segmented; abdomen of male and female with two pairs of biarticulate appendages and one pair of lobes. Type species Parvipalpus norvegicus Stephensen designated by Stephensen 1940). 68 1931 (by monotypy, subsequently Proaeginina norvegica Proaeginina norvegica (Stephensen 1931) Map 5 Synonymy Parvipalpus norvegicus Stephensen 1931 Proaeginina norvegica — Stephensen 1940, 1942, 1944a, 19446 — McCain 1966, 1968 — Laubitz and Mills 1972 Remarks This species has been recorded from Davis Strait and off Cape Cod, and it is therefore to be expected that it will eventually be recorded from. the Canadian Atlantic. As it is primarily a deep-water species (174-2702 m), it is not surprising that it was not found in the predominantly intertidal collections examined here. P. norve- gica can be recognized by the distinctive abdomen of this genus, and by its long and slender body with elongated pereonites 4 and 5. For detailed description, consult Laubitz and Mills (1972). 69 Table 1. Known Distributions of Canadian Arctic and Atlantic Caprellidae to Long Island Sound W Arctic (140°E to 125°W) NE Spitsbergen (S) Novaya Zemlya (N) Franz Joseph Land (F) D4 < < & & | KaraSea Siberia to 140°E Labrador to Gulf of St. Lawrence Faeroe Islands (F) SW Spitsbergen Pacific Alaska and Siberia Gulf of St. Lawrence < | W Greenland x< < x << XX x< SE Greenland . x< x x D4 NW Greenland NE Greenland (G) Iceland (I) N Norway and Barents Sea S of Cape Cod Species () Caprella dubia Metacaprella horrida Caprella carina C. microtuberculata Cercops holbolli Aeginina longicornis Aeginella spinosa Aeginina aenigmatica Caprella rinki Proaeginina norvegica Caprella septentrionalis C. ciliata C. linearis C. unica Mayerella limicola Caprella andreae C. equilibra C. penantis Paracaprella tenuis Hemiaegina minuta Luconacia incerta >< << | N American Arctic to 125°W Xx X X X X X X X X X x KK 70 Zoogeography The known distributions of the Cana- dian Atlantic and Arctic Caprellidae, illustrated by Maps 1 to 10, are sum- marized in Table 1. The’ three zoogeographical regions considered here, arctic, subarctic, and boreal, conform to Grainger’s (1966) modi- fication of Dunbar’s (1951, 1953) definitions, with the exception of the position of the boundary between subarctic and boreal regions. Dun- bar’s definition of this boundary was based on the southward penetration of arctic water. The caprellid fauna shows a marked change in the Gulf of St. Lawrence region. Such a faunal change has been noted for other groups (Hall 1964; Powell 1968). With division based on climate and faunal distribution, the subarctic province probably includes, besides southern Greenland, most of Hudson Bay and Labrador, northern and eastern Newfoundland only, and the northern Gulf of St. Lawrence, in- cluding the St. Lawrence estuary (Ganong 1890; Stephenson = and Stephenson 1954; Bousfield 1956a, 19566; Brunel 1961a). It is here considered that the boundary between subarctic and boreal faunal regions lies in the northern half of the Gulf of St. Lawrence. Although detailed ecolog- ical knowledge of the region is lacking, it appears probable that this boundary runs from northern New- foundiand (Strait of Belle Isle) near the north shore of the Gulf, north and west of Anticosti Island, and across the St. Lawrence estuary to northern Gaspé. Canadian Atlantic Region The Canadian Atlantic region lies within the subarctic and_ boreal zoogeographical provinces. These provincial divisions are based on coastal conditions, and pertain pri- marily to shallow-water regions. In deeper waters within each province, conditions are more like those of more northerly regions, with corres- ponding faunal differences. Brunel (1961a) and _ Drainville, Tiphane and Brunel (1963) have observed that the depths of the Saguenay fjord, which are colder and have a different fauna from both the St. Lawrence estuary and the Gulf, contain an “arctic enclave” isolated from the arctic province. Similarly, subarctic conditions exist within the boreal province, particularly in the Lauren- tian channel (Gulf of St. Lawrence), at the mouth of the Bay of Fundy and in its cold-water estuaries, and in the Gulf of Maine. Within the boreal province there are also ‘‘pockets” of southern, or warm-temperate, forms. The warm- temperate Virginian province’ is generally defined as lying between Cape Cod and Cape Hatteras. Ganong (1890) pointed out that typically Virginian faunas were to be found in variouS unconnected areas of the boreal region. These findings have been corroborated and expanded by later workers. The presence of warm marine pockets has been established in the following areas: warm-water bays and estuaries of the Bay of Fundy, particularly the St. John estuary, Chignecto Bay, Minas Basin, St. Marys Bay; sheltered coves and warm-water estuaries of Atlantic Nova Scotia; Newfoundland, in sheltered bays around the coast; the Magdalen shallows of the south- western Gulf of St. Lawrence (Ganong 1890; Huntsman 1924; Stephenson and Stephenson 1954; Bousfield 1956a-62). These warm pockets support relict populations of species widespread in this region 7 / Zoogeography during the postglacial period of minimum sea _ levels and warm temperate climate (Bousfield 1962, 1967). Seven caprellid species are here recorded from the Canadian Atlantic. Caprella rinki, a sublittoral species, was found only in the Laurentian channel, below 300 m. The distribu- tions of the other species reflect the zoogeographical conditions of this region. Caprella penantis and Paracap- rella tenuis, both warm-temperate species, were found almost ex- clusively within the warm Magdalen shallows (Maps 2 and 6). Except for a few isolated locations, these two species are otherwise recorded from south of Cape Cod. Caprella linearis and C. unica, and more particularly C. septentrionalis and Aeginina_ longicornis, were found mostly within the cooler northern part of the Gulf, in the St. Lawrence estuary, along the Atlantic shores of Nova Scotia, and in the Bay of Fundy (Maps 2, 6, and 9). Canadian Arctic Region Knowledge of the distribution of caprellids within the Canadian Arctic, and indeed within the Arctic as a whole, is limited and discontinuous. While certain areas of the Arctic have been studied quite extensively (for example, Greenland and northern Norway), we have very little know- ledge of the fauna of other regions such as the western Canadian Arctic. Therefore, any conclusions reached on the distribution of caprellids within the arctic and subarctic regions must be of a tentative nature. More ex- tensive collections are needed, parti- cularly from the far eastern and far western portions of these regions. A number of caprellid species have been reported from the Pacific (Mayer 72 1903), but they appear to be con- fined to south of the Bering Strait. As has been pointed out by Hedgpeth (1963) and Coachman (1963), the shallow waters of the Beaufort and Siberian seas have reduced salinity. Also, because of the effects of ice during the winter, they offer a very hostile type of environment (Mohr and Tibbs 1963). However, caprellid species recorded from other parts of the north-polar seas may well be present in the deeper, more saline waters to the north of the Pacific subarctic. Within the arctic region, Caprella dubia and Metacaprella horrida are probably endemic species. The ma- jority of records for these two species within the subarctic region are at depths below 100 m. Their dis- tribution is therefore of the high arctic type, as defined by Ekman (1953). In the subarctic region, two distinct patterns of distribution are found. The panarctic species, Caprella carina, C. microtuberculata, and pos- sibly Cercops holbolli, are found in arctic and subarctic regions, but do not extend southward into the boreal region. The _ arctic-boreal species, Aeginina longicornis and Aeginella spinosa, though primarily subarctic in distribution, are found also in the arctic region and as far south as Cape Cod, and for A. /ongicornis, Cape Hatteras. Caprella unica and Mayerella limicola are possibly endemic to the boreal region. The other species characteristic of this region show a variety of distribution patterns. Caprella linearis and C. ciliata have discontinuous circumpolar distribu- tion, being recorded from the north- ern Atlantic and Pacific oceans, with some occurrences in the subarctic. C. rinki and Proaeginina norvegica Canadian Arctic Region are sublittoral species, found usually below 200 m. Caprella septentriona- lis, a boreo-arctic species, is found mainly in boreal and_ subarctic regions, with few records for the Arctic. The single record for Aeginina aenigmatica is insufficient for con- clusions to be reached on its distri- bution, but it will probably prove to be a subarctic or arctic species. There appear to be two main zoo- geographical regions inhabited by north-polar caprellids: Atlantic- arctic, reaching from approximately 125°W eastward to at least 140°E, probably to 170°E; and Pacific sub- arctic, which includes the Chukchi and Beaufort seas. The only caprellid species recorded east of 140°E is Caprella micro- tuberculata. \t is possibly Siberian in origin, and has not yet been recorded west of Greenland. Four species have been recorded west of 125°W. Two of these, C. ci/- jata and C. /inearis, are circum- polar. C. carina may prove to be a circumpolar species, but its present distribution record indicates a Siberian or Pacific origin. Cercops holbolli appears to be confined to the North American Arctic. It may be a relict, one of a group of invertebrates that, according to Nesis (1962), crossed the Arctic during the Upper Pliocene. Certainly, the primitive nature of this species, and its possible presence in Japan, indicates a Pacific origin. The remaining species have not been recorded either east of 140°E or west of 125°W. Although sub- sequent collecting may show some of them to be circumpolar in distri- bution, it is apparent that the majority of north-polar caprellids are Atlantic in origin. Study of the literature indicates that similar distribution patterns exist in other faunal groups. Both asteroids (Grainger 1966) and pycnogonids (Hedgpeth 1963) have an arctic fauna that is predominantly Atlantic, with a Pacific subarctic fauna limited to the Chukchi Sea region. Filatova’s (1957) investigation of arctic bivalve mollusks of the U.S.S.R. showed that, while a number of Atlantic species are found as far east as the northern Chukchi Sea, _ relatively few north Pacific species were found to have penetrated into the East Siberian Sea. The arctic Bryozoa show a high degree of circumpo- larity, and Osburn (1955) and Powell (1968) have concluded that “there is no. significant difference between the bryozoan faunas of the Pacific-Arctic and Atlantic-Arctic areas’. However, the majority of the arctic species are Atlantic in origin, the few Pacific-arctic species having localized distribution. It appears that a similar situation exists in the gam- maridean amphipods, with a fairly homogeneous fauna in most of the cold-water region. There is, how- ever, a definite Pacific element in this fauna. There is also a smaller ele- ment radiating from the central Siberian Seas (Gurjanova 1951; Steele 1961). Two caprellids that may have originated in the latter region are Caprella carina’ and C. microtuberculata. Other caprellids have penetrated it from the west. Gurjanova’s other cold-water regions (1951) do not appear to be sup- ported either by amphipod or by other faunal distributions. However, the Chukchi-American transitional, as modified by Steele (1961), is ap- proximately equivalent to what has here been called the Pacific-sub- arctic. Lemche that (1941) recognized 73 Zoogeography zoogeographical classification within the arctic-subarctic-boreal region cannot. satisfactorily be achieved using only Ekman’s system. Geo- graphical location is less important than climatological and _ hydro- graphical conditions. Thus, cold- stenothermal species may be re- corded from all three regions, though from different depths. Lemche there- fore based his faunal classification on both propagative and vegetative requirements of the species. Such a system may be usefully applied to all arctic faunas. Study of the compo- sition and collecting data of the collections of most museums would produce much information on breed- ing biology and_ distribution of caprellids. Unfortunately, no analyses have been made of the available facts, and Lemche’s system cannot as yet be applied to the caprellids. In summary, we may say that there are two major faunal regions within the north-polar seas. The Atlantic-arctic (including the sub- arctic) extends from at least 125°W, eastward to at least 140°E. The Pacific-subarctic is confined to 140°E to 125°W at most. The ma- jority of arctic caprellids are of Atlantic origin, while the Pacific- arctic species have very localized distributions, being found for the most part south of the Bering Strait. 74 Discussion Many of the problems encountered when dealing with the Caprellidae will be solved only when detailed ecological and _ genetical studies have been carried out, and when we have detailed knowledge of the life history of these animals. Some of these problems will be discussed briefly. In certain caprellid species, for example, Aeginella spinosa, body Spination appears to be quite con- stant; more frequently, however, the body spination is highly variable. In some species, such as Aeginina longicornis or Caprella unica, both smooth and spiny forms are found. In others, the size of the spines varies, with only certain spines always being present, as in the case of Cercops holbolli. \t has been observed that spination is often related to growth, so that in many species, immature and female specimens are spinier than adult males. However, this relationship does not hold for all species, nor for all specimens of species where it is Common. It Is therefore probable that other factors are involved in determining the ex- tent of spine development. Ecological, and perhaps geographical, conditions may prove to influence the develop- ment of spines in certain caprellids. Experiments need to be carried out to ascertain what external factors are here involved. A second problem is lack of know- ledge of the growth, development, and life cycles of caprellids. Collec- tions of caprellids in many institutions could yield much information on these subjects. The data are avail- able, but need to be analysed. For example, until more is known we Cannot know why Caprella penantis is larger and more robust in the south, i.e. Florida, than in the north, i.e. Gulf of St. Lawrence. It may be that the north, being colder, is less congenial to this species, affording a shorter season for growth. How- ever, the explanation may not be as simple as this. In both Aeginina longicornis) and Caprella_ septen- trionalis, large individuals were found at both northern and southern extremes of the range, and the local- ity records for small individuals (one- third to one-half of the usual adult size) offered no reason for their size. Similarly, no explanation can be given for the large female C. sep- tentrionalis, with undeveloped brood plates, in a collection where the ovigerous females were half the size. A third problem lies in the literature of the group. It is apparent that many species, particularly of Capre//a, are very similar to one another. It is not always possible to determine to which species of a similar pair a record refers, and many records may be suspected of referring to both. It should therefore be emphasized that, for such species as C. /inearis and C. septentrionalis, C. microtuberculata and C. dubia, some of the references are questionable, and synonymy lists should be regarded only as guides to the literature. 75 : . i i Y ry ary t : +> | ; aj vi a D Ne a") » 7 DY, i ’ aa _ A Pease. Wa ot Oe all i ue "3 Spisrtlwvts te DA SHAE EE es weet! * Leese “— rey ey al | oa uae ae reat ae nts ees ref ' ey vrei nt ¢ Jae and 7 ra? i v ; evita: re 4 ¢ .. ne es ce v me epee ye he) errant art ; 7 ot Fad oe i ae vier it fis ty} taht: bli =e Gert os a aa) OEY 1 ea “ij vi raed yh : ’ ; we Hele 84 Ase ( i 4 is) (ae oor oehmee int soem he UR? NPA ite Vee the ; 4 : ’ . RR. Ride ery Ghia pe SF to eae i) daiadliaets ates, «akc aancanat vo dtnpira i israel Sier Coe en ej et Ce f Gyo - t mye syeeig ih Are Hada hd? ha 4 i pie be te MS ls a ee ne ) . oh ems We ea PEvid of we Ge ¢ te} nl |= a Hae Seven re fe Tiah ? oe shal Ae Ooh ) | ag VENTE So wes y eal ie ag! Me | ay MRM Gn PEN eG Bie gate’ y = Say ange CHINO AiR OP be Retr BOP BAN IG cton 1KWOt toi aes ‘Crehet wipe bar ta’ WLS kee’ FT a o ee bal ea 1 GAG eet Kase ; Prac Ve f > = v% Lal, Ss WHE ‘ 7 (Oda a5 OF anergy =~ Toy fut cert reuth | , i i 7 i: i eS i+ eRe Ae | ; Ir : b a. eae et a oe ‘ " . Wee lie id Veiled ee hte. , 7: V9 / peerey: i ; ‘ : % ‘ ¥ Lom “4 hips Ey) ae nae ae o ne 4 ri ea | 4 ; d's he ve % ' e i. ie 7 os a) { ‘ : oe ar im ae! hitelfuooahe ‘wth pies oti iat soos gh Th paca ati drive hi eat ny / ele 7 tov iae wine bevdou ‘ a "eee bi i oe Fin , . : oP oj Q N — = , i 2 ie vel’ ermeaihgg. mt Us) ria * 5 ‘ : WW a ‘i Qf tary , - - = i = i ene bh F8 “i civ be Re :7 » . e,' weed iva ’ ‘i ying ‘ ‘ S } Ff ~ ny - é ® Pes Tl = whcpe . ¢ ve A ’ ri 6 i 4 "4 eure be) act eM TUM Myo ° - —_ \ ae | — ' > F ve 1 mdeoe ow vr f¢ van { ' r| (6 aie 7 r ‘ U » if 7. : C1) ay 2 ; af _ b 4 4 5 MAA igo nYo{ af gS ‘ } ons , eNOS stale wDys Wd hi tar ay gt saa fa. -evgana nn), dau: ei liven ocyg Atel ott tenis t Gayyliete ad ag Dawe gud rN TUVEnY i O9iet) MAA Apieit yp yorves) rire ait rin. tude were bee Atom FT ub ost sleet ely a. A Literature Cited Arimoto, I. (1930). Studies on the Caprellidae from Tateyama Il. J. Tokyo Natur. Hist. Soc. 28 (39): 45-56. Barnard, J.L. (1969). 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Al ia » Copratiden./ foc Se) Mothada Univ, sor © NERY URE T89-KH _ - 1947) Canrelidiae @ depen wed atidoon waters io Soper) FePvid Svaot 1 60-02 | 19964) Cepcolides, p> 11-15 jn T Kaech Le ‘Paune and Aart of tha tee ground tiie Amenosa : Moria Litlagwol Lihoratar, o! 8 Amphipes ry Crustacee 21988), Eptenti« or planktonic capresiids / ‘Ria eritiora of Tanabe Gey (40s ymnie | | Pubi Sete Mar, Boi Lad, Kyore i. 7 281.89 trom Kemet Cay, sorth- | tAmohinoda® Cepretnine! ry os ae aos awe a La a = a a at oo Bik Leb. 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