4* 0f c s a o UNITED STATES / NATIONAL OCEANIC AND / National Marine DEPARTMENT OF COMMERCE / ATMOSPHERIC ADMINISTRATION / Fisheries Service Rogers C. B. Morton, Secretary / Robert M White Administrator / Robert w Schonmg Director a ^* For ■ nlr by the Superintendent of Document*. U.S. Government Printing Office. Washington. D.C. 20402 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. CONTENTS Page Introduction 1 Field methods 2 Laboratory methods 3 Material studied 6 The Arctic Spinocalanidae 9 Arctic hydrography 9 Vertical distribution of Arctic Spinocalanidae 9 Family Spinocalanidae Vervoort, 1951 13 Description L3 Female 13 Male 14 Remarks 14 Key to the Genera of Spinocalanidae 15 Genus Spinocalanus Giesbrecht, 1888 15 Description 15 Key to the species of Spinocalanus 16 1. Spinocalanus abyssalis Giesbrecht, 1888 17 Description 17 Remarks 19 Distribution 20 2. Spinocalanus longicornis Sars, 1900 20 Description 24 Remarks 25 Distribution 25 (3. Spinocalanus schaudinni Mrazek, 1902) 26 4. Spinocalanus magnus Wolfenden, 1904 26 Description 26 Remarks 28 Distribution 29 5. Spinocalanus antarcticus Wolfenden, 1906 30 Description 30 Remarks 35 Distribution 35 (6. Spinocalanus major Esterly, 1906) 35 (7. Spinocalanus latifrons Sars, 1907) 35 8. Spinocalanus hirtus Sars, 1907 35 Description 35 Remarks 36 9. Spinocalanus spinosus Farran, 1908 36 Description 36 Remarks 36 Distribution 37 10. Spinocalanus horridus Wolfenden, 1911 37 Description 37 Remarks 40 Distribution 41 (11. Spinocalanus giesbrechti Brady, 1918) 41 Remarks 41 12. Spinocalanus validus Sars, 1920 41 Description 42 Remarks 42 Distribution 42 13. Spinocalanus angusticeps Sars, 1920 42 Description 43 Remarks 43 Distribution 43 (14. Spinocalanus caudatus Sars, 1920) 43 iii (15. Spinocalanus abyssalis var. pygmaeus Farran, 1926) 43 (16. Spinocalanus heterocaudatus Rose, 1937) 44 (17. Spinocalanus pacificus Mori, 1942) 44 (18. Spinocalanus stellatus Brodsky, 1950) 44 (19. Spinocalanus spinipes Brodsky, 1950) 44 (20. Spinocalanus pseudospinipes Brodsky, 1950) 44 21. Spinocalanus similis Brodsky, 1950 44 Description 44 Remarks 46 Distribution 46 (22. Spinocalanus dorsispinosus Brodsky, 1950) 46 (23. Spinocalanus sp.? Brodsky, 1950) 46 (24. Spinocalanus longispinus Brodsky, 1950) 46 25. Spinocalanus elongatus Brodsky, 1950 46 Description 46 Remarks 49 Distribution 50 26. Spinocalanus polaris Brodsky, 1950 50 Description 50 Remarks 53 Distribution 53 27. Spinocalanus brevicaudatus Brodsky, 1950 53 Description 54 Remarks 55 Distribution 56 (28. Spinocalanus similis var. profundalis Brodsky, 1955) 57 (29. Spinocalanus longipes Tanaka, 1956) 57 (30. Spinocalanus? sp. Johnson, 1963a) 57 31. Spinocalanus abruptus Grice and Hulsemann, 1965 57 Description 57 Remarks 57 Distribution 57 (32. Spinocalanus oualis Grice and Hulsemann, 1965) 57 (33. Spinocalanus ventriosus Grice and Hulsemann, 1967) 57 (34. Spinocalanus sp. Grice and Hulsemann, 1967) 57 Remarks 57 (35. Spinocalanus parabyssalis Park, 1970) 57 (36. Spinocalanus pteronus Park, 1970) 57 37. Spinocalanus usitatus Park, 1970 58 Description 58 Remarks 58 Distribution 59 38. Spinocalanus hoplites Park, 1970 59 Description 59 Remarks 59 Distribution 59 39. Spinocalanus aspinosus Park, 1970 59 Description .59 Remarks 59 Distribution 59 40. Spinocalanus oligospinosus Park, 1970 59 Description 60 Remarks 60 Distribution 60 (41. Spinocalanus paraoligospinosus Hure and Scotto di Carlo, 1971, nomen nudum) .... 60 (42. Spinocalanus neospinosus Grice, 1971) 60 43. Spinocalanus terranovae, new species 60 Description 60 Remarks 62 Distribution 62 iv Genus Monacilla Sars, 1905 62 Description H2 Key to the species of Monacilla 64 1. Monacilla typica Sars, 1905 64 Description 64 Remarks 65 Distribution 65 2. Monacilla tenera Sars, 1907 65 Description 65 Remarks 66 Distribution 66 (3. Oxycalanus spinifer Farran, 1908) 66 (4. Oxycalanus semispinus Scott, 1909) 66 (5. Monacilla dubia Scott, 1909) 66 (6. Hypsicalanus gracilis Wolfenden, 1911) 66 7. Monacilla gracilis (Wolfenden, 1911) 66 Description 66 Remarks 66 Distribution 66 (8. Monacilla typica var. asymmetrica Tanaka, 1956) 66 9. Monacilla sp 66 Remarks 66 Distribution 67 Genus Mimocalanus Farran, 1908 67 Description 67 Key to the species of Mimocalanus 67 1. Mimocalanus cultrifer Farran, 1908 68 Description 68 Remarks 69 Distribution 69 2. Mimocalanus nudus Farran, 1908 69 Description 69 Remarks 72 Distribution 72 3. Mimocalanus major Sars, 1920 72 Description 73 Remarks 73 Distribution 73 4. Mimocalanus inflatus Davis, 1949 73 Description 73 Remarks 73 Distribution 73 (5. Mimocalanus distinctocephalus Brodsky, 1950) 73 6. Mimocalanus ovalis (Grice and Hulsemann, 1965), new combination 73 Description 73 Remarks 74 » Distribution 74 (7. Mimocalanus species Grice and Hulsemann, 1967) 74 8. Mimocalanus crassus Park, 1970 74 Description 74 Remarks 70 Distribution 76 9. Mimocalanus sulcifrons Wheeler, 1970 76 Description 79 Remarks BO Distribution SO 10. Mimocalanus heronae, new species 80 Description Remarks Distribution B2 Genus Teneriforma Grice and Hulsemann, 1967 82 Description 82 Teneriforma naso (Farran, 1936) 83 Description 83 Remarks 83 Distribution 83 Summary 83 Acknowledgments 84 Literature Cited 84 Tables 1. Source of Arctic specimens 3 2. Estimated numbers of Arctic Spinocalanidae from 24 depth intervals 4 3. Sources of borrowed specimens 6 4. Spinocalanus and Mimocalanus, Al armature 8 1. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. Figures Positions and track of Fletcher's Ice Island ("T-3") in the Canadian Basin, June-September 1967 Positions and track of Fletcher's Ice Island ("T-3") in the Canadian Basin, June-September 1968 Vertical distribution of temperature and salinity for four stations and the principal water masses of the central Arctic Ocean Spinocalanus abyssalis, female, habitus, dorsal view Spinocalanus abyssalis, female, habitus, lateral view Spinocalanus abyssalis, female, Mxp Spinocalanus abyssalis, female, PI Spinocalanus abyssalis, female, P2 Spinocalanus abyssalis, female, P3 Spinocalanus abyssalis, female, P4 Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis Spinocalanus longicornis female, habitus, dorsal view 21 female, habitus, lateral view 21 female, Al 21 female, Mn blade 21 female, Mxl 21 female, Mx2 21 female, Mxp 21 female, PI 22 female, P2 22 female, P3 22 female, P4 22 male, A2 22 male, Mn blade .22 male, Mn blade 22 male, Mxl gnathobase 22 male, Mx2 22 male, habitus, dorsal view 23 male, habitus, lateral view 23 male, Al 23 male, Mxp 23 male, P4 23 male, P5 23 male, P5 23 male stage V, P5 23 Spinocalanus magnus, female, habitus, lateral view 27 VI Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus Spinocalanus magnus, female, habitus, dorsal view 27 magnus, female, habitus, lateral view 27 magnus, female, Th5 and urosome, dorsal view 27 magnus, male, habitus, dorsal view 27 magnus, male, habitus, lateral view 27 magnus, male, P5 27 magnus, male stage V, P5 27 e, genital segment 30 e, A2 30 e, habitus, dorsal view 31 e, habitus, lateral view 31 e, Al 31 e, Mxl 31 e, Mxp 31 e, Mn 32 e, Mn blade 32 e, Mx2 32 e, PI 32 e, P2 32 e, P3 32 e, P4 32 habitus, dorsal view 34 habitus, lateral view 34 Al 34 Mn blade 34 Mxl gnathobase 34 Mxp 34 PI 34 P2 34 P3 34 P4 34 P5 34 male stage V, P5 34 horridus, female, habitus, dorsal view 38 horridus, female, habitus, lateral view 38 horridus, female, Mxl gnathobase 38 horridus, female, Mx2 38 horridus, female, Mxp 38 horridus, female, PI 38 horridus, female, P2 39 horridus, female, P3 39 horridus, female, P4 39 horridus, male, habitus, dorsal view 39 horridus, male, habitus, lateral view 39 horridus, male, Mn blade 39 horridus, male, Mxp 39 horridus, male, P5 39 horridus, male stage V, P5 39 spinosus, female, PI 40 angusticeps, female, P3 42 similis, female, habitus, dorsal view 45 similis, female, habitus, lateral view 45 similis, female, Mxp Bl-2 45 similis, female, Pi 45 similis, female, P2 4"-> similis. female, P3 45 similis, female, P4 similis, male, habitus, dorsal view similis, male, habitus, lateral view (5 similis, male, Mn blade 45 vii antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus femal antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male, antarcticus male 96. Spinocalanus similis, male, Mxp 45 97. Spinocalanus similis, male, P5 45 98. Spinocalanus elongatus, female, habitus, dorsal view 47 99. Spinocalanus elongatus, female, habitus, lateral view 47 100. Spinocalanus elongatus, female, Mn blade 47 101. Spinocalanus elongatus, female, PI 47 102. Spinocalanus elongatus, female, P2 47 103. Spinocalanus elongatus, female, P3 47 104. Spinocalanus elongatus, female, P4 47 105. Spinocalanus elongatus, female, Mxp 48 106. Spinocalanus elongatus, male, habitus, dorsal view 48 107. Spinocalanus elongatus, male, habitus, lateral view 48 108. Spinocalanus elongatus, male, Mxp 48 109. Spinocalanus elongatus, male, PI 48 110. Spinocalanus elongatus, male, P2 48 111. Spinocalanus elongatus, male, P3 48 112. Spinocalanus elongatus, male, P5 48 113. Spinocalanus elongatus, male stage V, P5 48 114. Spinocalanus polaris, female, habitus, dorsal view 51 115. Spinocalanus polaris, female, habitus, lateral view 51 116. Spinocalanus polaris, female, Mx2 lobe-1 51 117. Spinocalanus polaris, female, PI 51 118. Spinocalanus polaris, female, P2 51 119. Spinocalanus polaris, female, P3 51 120. Spinocalanus polaris, female, P4 51 121. Spinocalanus polaris, female, Mn blade 52 122. Spinocalanus polaris, female, Mxl gnathobase 52 123. Spinocalanus polaris, female, Mxp 52 124. Spinocalanus polaris, male, habitus, dorsal view 52 125. Spinocalanus polaris, male, habitus, lateral view 52 126. Spinocalanus polaris, male, Mn blade 52 127. Spinocalanus polaris, male, Mxp 52 128. Spinocalanus polaris, male, PI 52 129. Spinocalanus polaris, male, P5 52 130. Spinocalanus brevicaudatus, female, Mxp Bl-2 54 131. Spinocalanus brevicaudatus, female, P2 54 132. Spinocalanus brevicaudatus, male, PI 54 133. Spinocalanus brevicaudatus, male, P2 54 134. Spinocalanus brevicaudatus, male, P3 54 135. Spinocalanus brevicaudatus, male, P4 54 136. Spinocalanus brevicaudatus, male, P5 . 54 137. Spinocalanus usitatus, female, habitus, dorsal view 58 138. Spinocalanus usitatus, female, habitus, lateral view 58 139. Spinocalanus usitatus, female, Mx2 58 140. Spinocalanus usitatus, female, Mxp Bl-2 58 141. Spinocalanus terranovae, female, habitus, dorsal view 61 142. Spinocalanus terranovae, female, habitus, lateral view 61 143. Spinocalanus terranovae, female, Mxp Bl-2 61 144. Spinocalanus terranovae, female, PI 61 145. Spinocalanus terranovae, female, P2 61 146. Spinocalanus terranovae, female, P3 61 147. Spinocalanus terranovae, female, P4 61 148. Spinocalanus abyssalis, female, Al terminal segments 63 149. Spinocalanus longicornis, female, Al terminal segments 63 150. Spinocalanus magnus, female, Al terminal segments 63 151. Spinocalanus antarcticus, female, Al terminal segments 63 152. Spinocalanus antarcticus, female, Al terminal segments 63 153. Spinocalanus horridus, female, Al terminal segments 63 154. Spinocalanus similis, female, Al terminal segments 63 155. Spinocalanus elongatus, female, Al terminal segments 63 viii 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. 212. 213. 214. 215. Spinocalanus Spinocalanus Spinocalanus Spinocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus Mimocalanus polaris, female, Al terminal segments 63 breuicaudatus, female, Al terminal segments 63 oligospinosus, female, Al terminal segments 63 terranouae, female, Al terminal segments 63 ovalis, female, Al terminal segments 63 crassus, female, Al terminal segments 63 sulcifrons, female, Al terminal segments 63 heronae, female, Al terminal segments 63 cultrifer, female, habitus, dorsal view 68 cultrifer, female, habitus, lateral view 68 cultrifer, female, PI 68 cultrifer, female, P2 68 cultrifer, female, P4 68 nudus, female, habitus, dorsal view 70 nudus, female, habitus, lateral view 70 nudus, female, Mn blade 70 nudus, male, habitus, dorsal view 70 nudus, male, habitus, lateral view 70 nudus, male, Mn blade 70 nudus, male, P5 70 nudus, female, PI 71 nudus, female, PI Re2-3 71 nudus, female, P2 71 nudus, female, P3 71 nudus, female, P4 71 crassus, female, habitus, dorsal view 75 crassus, female, habitus, lateral view 75 crassus, female, Mn blade 75 crassus, female, PI 75 crassus, female, P2 75 crassus, female, P3 75 crassus, female, P4 75 crassus, male, habitus, dorsal view 75 crassus, male, habitus, lateral view 75 crassus, male, Mn blade 75 crassus, male, P5 75 sulcifrons, female, habitus, dorsal view 77 sulcifrons, female, habitus, lateral view 77 sulcifrons, female, A2 77 sulcifrons, female, Mn blade 77 sulcifrons, female, Mxl gnathobase 77 sulcifrons, female, Mx2 77 sulcifrons, female, PI 77 sulcifrons, female, P2 77 sulcifrons, female, P3 77 sulcifrons, female, P4 77 sulcifrons, female, Mxp 78 sulcifrons, male, Mn blade 78 sulcifrons, male, Mxp 78 sulcifrons, male, PI 78 sulcifrons, male, P2 78 sulcifrons, male, P2 Re3, abnormal 78 sulcifrons, male, P3 - sulcifrons, male, P4 78 sulcifrons, male, P5 3 sulcifrons, male, P5 B sulcifrons, male stage V, habitus, lateral view 78 sulcifrons, male stage V, P5 3 sulcifrons, male, habitus, lateral view 79 sulcifrons, male, habitus, dorsal view 79 IX 216. Mimocalanus heronae, female, habitus, lateral view 8 217. Mimocalanus heronae, female, habitus, dorsal view 8 218. Mimocalanus heronae, female, Mn blade 8 219. Mimocalanus heronae, female, Mxp 8 220. Mimocalanus heronae, female, PI 8 221. Mimocalanus heronae, female, P2 8 222. Mimocalanus heronae, female, P3 8 223. Mimocalanus heronae, female, P4 8 224. Teneriforma naso, male, P5 8 225. Scales A-H, 0.1 mm 8 CALANOID COPEPODS OF THE GENERA SPINOCALANUS AND MIMOCALANUS FROM THE CENTRAL ARCTIC OCEAN, WITH A REVIEW OF THE SPINOCALANIDAE DAVID M. DAMKAER 1 ABSTRACT The family Spinocalanidae includes small to medium-sized marine calanoid copepods belonging to the genera Spinocalanus, Monacilla, Mimocalanus, and Teneriforma. All species are deep-living and often comprise a large proportion, or even a majority, of the copepods in deep samples. In spite of their prevalence, definitive knowledge of the Spinocalanidae has lag- ged behind that of other copepod groups because adequate collections from deep water have been few, and specimens from widely separated localities have seldom been compared. Most im- portant, however, is the fact that the fragility of the specimens makes them very difficult to study; most investigators attempting to describe or identify Spinocalanidae have indicated that their specimens were damaged and incomplete. The present study is based on collections of zooplankton from Fletcher's Ice Island, T-3, in the Canadian Basin of the Arctic Ocean in 1967-68. The seven species of Spinocalanidae from these collections are redescribed, and their vertical distributions are discussed, based on series of samples from discrete depth intervals to 3,000 m. The systematics of the Spinocalanidae has been reconsidered, using characters in addition to those most commonly lost in sampling. All published descriptions and records are discussed. Several critical type specimens and specimens forming the bases of widespread records have been examined and are redescribed. Keys to the genera and all of the species have been prepared, with the goal of enabling an investigator to identify even damaged specimens. Many named species or forms have been placed in synonymy, and two new species (Spinocalanus terranovae and Mimocalanus heronae) are described. The family is now considered to comprise 32 species, distributed as follows: Spinocalanus (19), Monacilla (4), Mimocalanus (8), and Teneriforma (1). INTRODUCTION The family Spinocalanidae includes small to medium-sized marine calanoid copepods belonging to the genera Spinocalanus (42 named species or forms), Monacilla (8), Mimocalanus (8), and Teneriforma (1). The Spinocalanidae have been difficult to study because they are prone to damage in sampling, resulting typically in the loss of the first antennae and most swimming legs. Grice and Hulsemann (1965) said that "the identification of species of Spinocalanus was difficult as almost all the specimens examined were damaged .... This appears to be the case for other collections also because most previous descriptions are accompanied by few il- lustrations .... Most of the species in this genus need 'Systematica Laboratory, National Marine Fisheries Service, NOAA, Washington, D.C. 20560; present address: Pacific Marine Environmental Laboratory, NOAA, University of Washington WB-10. Seattle, WA 98195. to be restudied and redescribed, a task which will be difficult unless specimens in better condition than the present ones can be obtained." This fragility and rather generalized body form have given the Spinocalanidae a potential to be overlooked or ig- nored in surveys of deep-living copepods, even though they often comprise a large proportion, or even a ma- jority, of the copepods in deep samples. Also, as in many other calanoids, adult males are scarce. John- son (1963a), in discussing this group, said that "the clarification of the position of these males and the whole complex of Spinocalanus species requires ad- ditional material and much more thorough study." Definitive knowledge of the Spinocalanidae has also lagged behind that of other copepod groups because the species are mostly deep-living; adequate collec- tions have been few, and specimens from widely separated localities have seldom been compared. Most have been collected by towing plankton nets vertically from great depths to the surface, so that depth limits of the species are difficult to define. Their absence from surface-layer samples, however, indirectly fixes their upper depth limit, generally at about 100 m. Spinocalanids have been collected in a lesser number of deep closing-net samples taken at discrete depths. A lower depth limit, if there is such a limit, has not been fixed; some species have an ex- traordinary vertical distribution, from 50 m to at least 5,000 m. The great vertical range gives these species a potential for wide, in some cases worldwide, distribu- tion in the relatively uniform and interconnected deep-ocean water masses. The lack of records for some species from entire oceans cannot yet be accepted as an indication of real absence. In particular, the ap- parent bipolar distribution of Spinocalanus antarc- ticus should be reconsidered, to determine its dis- tribution into lower latitudes. In the decades since World War II interest in the Arctic Ocean has greatly increased, for commercial, strategic, and scientific reasons. The heroic days of the earliest Arctic explorations, when observations were gained under the harshest conditions, are past. In spite of the difficult and distant environment, in- formation now accumulates relatively rapidly. The development of air transportation and the use of floating ice platforms have made all Arctic areas accessible throughout the year. Still, most subjects of investigation remain in an exploratory stage; this is especially true of the oceanic plankton of the high polar basins. The oceanic copepods, the most numerous of the net-zooplankton, are the subject of less than a dozen published reports, and only three or four of these consider deepwater species. The present study is based on collections of zooplankton from Fletcher's Ice Island, in the Canadian Basin in 1967- 68. The spinocalanids from these collections are redescribed, and their vertical distributions are dis- cussed. The systematics of the Spinocalanidae has been reconsidered using characters in addition to those usually considered, the antennae and legs. Many named species are placed in synonymy, and two new species are described. The family is now considered to comprise 32 species, distributed as follows: Spinocalanus (19), Monacilla (4), Mimocalanus (8), and Teneriforma (1). Keys to all the species have been prepared, with the goal of enabling an investigator to identify even damaged specimens. Field Methods The sampling base for the Arctic collections was Fletcher's Ice Island, also called "T-3." T-3 is a tabular ice sheet ca. 8 X 13 km and 30 m thick, fragmented from northern Ellesmere Island. T-3 is caught up in the pack ice, with which it drifts at about 2-4 km/day. At the time the material to be described was collected, T-3 was in the Canadian Basin, ranging in latitude from 79°30'N to 85°N and in longitude from 142° W to 175° W (Figs. 1, 2). Positions were determined only every few days. Mohr (1959) and Schindler (1968) have summarized the history and use of the ice island. Zooplankton samples were collected through a hole in the sea ice (3-4 m thick) bordering the island. A small prefabricated plywood hut was assembled over the hole; the hut housed a winch, with 4,000 m of hydrographic wire, powered by a 5-hp gasoline engine. The wire ran through a meter wheel attached to a 2-m tripod placed over the hole; a 50-kg weight was secured to the end of the wire. TSE-(T-3)-4 JUNE-SEPTEMBER 1967 Figure 1. — Positions and track of Fletcher's Ice Island ("T-3") in the Canadian Basin, June-September 1967. TSE-(T-3)-7 JUNE -SEPTEMBER 1968 i T-3 (to scale) Table I uens. Figure 2.— Positions and track of Fletcher's Ice Island ("T-3") in the Canadian Basin, June-September 1968. Samples were collected in 1-m closing nets with mesh apertures of 0.215 mm (1967) and 0.110 mm (1968). A series of vertical samples was obtained in summer 1967 (July and September), and a second series in summer 1968 (June, July, and August), a total of 142 samples. Depth intervals sampled were: 10-100 m, 10-m intervals; 100-200 m, 20-m intervals; 200-400 m, 50-m intervals; 400-500 m, 100-m inter- vals; 500-3,000 m, 500-m intervals. Usually more than one sample was obtained from each interval each summer. Some specimens were obtained from samples not in the above routine series; these samples are included in Table 1, but the specimens are not enumerated in Table 2. Samples were fixed in 5% formaldehyde/seawater solution, buffered with sodium borate. Laboratory Methods Arctic specimens were preserved in 4'. form- aldehyde/3' ( sodium acetate solution; most borrowed specimens were preserved in ethyl alcohol. Gross ob- servations and drawings were made in water or water/ glycerine solution, to and from which the specimens Program ■ Depth Sample Date 573 12 VII 67 8 • - 847 8 848 - K67 850 9 10 5 60 10 VI 68 500 61 10 \1 68 62 10 \ 1 68 ! 63 10 300-350 64 10 VI 68 80 12 VI 68 500- l.i xrfi 81 12 V 1 68 400 120 14 VI 68 400 121 14 VI 68 1,000 122 24 VI 68 3.000 123 24 VI 68 2.00< 124 24 VI 68 125 24 VI 68 1,000-1.500 126 24 VI 68 500-1.000 127 24 VI 68 400-500 167 7 VII 68 500-1. UN. 168 7 VII 68 1.000-1. .500 169 7 VII 68 170 7 VII 68 212 19 VII 68 500 213 19 VII 68 1.000 214 19 VII 68 1,500 215 19 VII 68 216 26 VII 68 2,000 217 26 VII 68 !.."«' X 1-2.000 218 26 VII 68 1,000-1.500 219 26 VII 68 500-1,000 220 26 VII 68 500 282 7 VIII 68 500-1,000 283 i VIII 68 1.000-1.500 284 7 VHI68 1.500-2,000 285 * vm6a 2.00i • 286 13 VIII 68 1,501 287 13YIII6S 1.000-1. .Vm 288 L3Vm68 500-1,000 289 t3vra6a 500 309 20VTJI68 1.5IM-2.000 310 21 VIII tW 1,000 311 21 \! 1,000 2272 26 \ 3005 5 Mil us 3006 -Mil 68 21 Villus 3076 14 \ 111 68 3090 18 VTA 68 Sample Al A2 A:i A4 A6 A6 A7 AH A9 A10 All A12 A 13 A14 A15 A16 A17 A18 A19 A20 A21 A22 A23 A24 A25 A26 A27 A28 A29 A30 A31 A32 A3.') A34 A35 A36 A37 A38 A39 A40 A41 A42 A43 A44 A45 A46 A47 A48 A49 A50 A51 A.-.2 were moved slowly through series ol solutions. Dis- sected specimens were stained overnight in methyl blue/lactic acid solution; detached appendages were oriented, drawn (usually without cover slips), mounted in glycerine under 9-mm diameter cover slips and sealed with a heavy ring of nail-polish Lacquer I sually •9 a S w v I c + © 2 cu CO 3 e § a E o 03 c CD D. > j: ed tie CD £ 3 C CD .2/3 ■O 9) ■S 3 « .s -O est £ £ 3 «g 0! II Jo fa 13 • - * 2 01 03 £ 3 C •a CD CG £ fa CM 3 CO 3 so c ••2 CO CO u o o Or B c a o M "3 u 3 c u a H o oj o < CO t/3 a ~ oooooooooo oooooooooo o o o mm^ior-t-Nujujio lO lO m oooooooooo oooooooooo oooooooooo © © © © ©©©©©©©©oo iolo-^loc^ol-^lololo ©©©©©©© !D ■* iH CM © CO © CO i-B !-S .. lO i-H CM fa co LO a 1-3 © s ►-3 O © 3 CO T-H CM *-3 ""3 •"3 © CO LO © a >-3 lO co t— t lO 1-3 rH i-a *~3 © . © fi- LO fa fa CM fe CD io •>* © Ol • - o co r-i co »— I co *"» CM lo ... CM o © © © © © >-3 fa © CM © © © © 115 lO "* lO •-3 i-a >-3 © -rH lO CM ... CO "I .. fa r - "-3 r - >-3 5 i_h i-a co fa © fa co © 2 © h ►, n h o ..n >-3 ^ ■* ^,- N ,_l »ir ^ 5 ^- .[Ju[i 4 , -5©'-3'^>00 » - » »v _ >S -* fe - 1 ©©©ocTo^^w^co^ C— C- C- CO CM r- I © CO CM >-3 © © CM ° s rH CO .. © ._ © -3 - 1 ■ © r - LO fa © fa lO lOLO-rfLOt— C~C— lOLOLO © o ©0©©©©00©CM NCO^mCOC-COCDHH o o CM Mi i-H - - o "* CM r fa LO o o CO >-3 o CO !-3 1-1 !-3 CO © © fa >-3 LO fa © ■- 1 CO . . 1-3 !-3 o in CO !-S CO © © 3 LM fa t> fa o lO ^-1 S" CM o 1—1 fa 2 Pi- rn S fe CM s oo 05 fa CO CO fa CM !-3" lo •"3* fa CM CM co CO CO s lO 2 CO © o © CM fa lO fa o fa a CO £ fe m s Ix ^ fa CO Oi a co CO [I, CM © fa 00 fa fa © fa CO CM 3 Ian .—I 2 co o t> 2 co CM fa CM -5 © fa oo CN fa © r^ CO b Of) CO «l ro CO t— CM ■* t- lO :S fa fa © . fa fa ^ TO .fa fa M ^Ifa o CO a a © ^ §3 3 a" a" a" a CS| -^< i-H ^H - fa" fa" fa" fe CO ■"* 00 CO CO H .. >-3 '"S « o in »n co co >-3 CM rH r- ( , fc -IT fa C&" % .. fa © "^ lO m fa 2 CN fa © §'S CM CM CM fa" -3 LO LO rH a W LO fa fe CM fa 03 fa o CO 1-3 "- 1 LO ■ - •rj" fa" fa" fa >-3 fa is CO © CM 1-3 O Ol ^ C5 CM rH fa" CO fa CM ■>* § , - Oi fa . a fa" co co fc .*. fa LO fc r ■• r~i fa o >-3 m fa ■- C3 fa 03 O y-H 'S © a © -tf CN 7. r- o 1-1 © 1-1 rH o CM ■* © o LO CO O O O fa rH fa fa CM fa" fa 00 fa O r.s a CM fa" CO ©" « fa CM •- . fa fa ^ LO •- © rH CN rH a co fa" LO LO 71 fa © "5 © 3 C c c U .c •a a> c^ > ^ JD E 3 X. a, o > ^ o o. o g CO 1 CD £> E 3 z /: °- E Q oooooooooooooo ■.o o — CO — i CM O p p p p ^ in 00 — ' lO 300000003000000000 + + fa." b 6u CM o o o o pppppppppppppppp lO^^^iomioioiomioioiitoii^m [i, -s . "-a i-a m "* bu "" ~ h « £ co © CD 2S 00 — ' 00 d CO 2 - s -s 22 T N 5 .. 1H .- U. El, O U3 ^ Ex. t- CM oooooooooooooooooooo CO d - - ppppppppppppppppppp- inio^int^r~r~ininininioioioioci>cocoao — m r~ i- i- ifl ic ic in i- i- i- ic cc co co ao o b. — Ex. O CM — CM Ex." s Ex.' N Tf u. PS s a 2 £ s -i Li.' ~ s in Ex." u. — - — - — N t u. t '- ct — ©©©©:3SO©§eM^«Dacc355S5?S8 Nn^mtoi-ccS- — — — • — m m ~ r S - S 8 K2t-ce35o?iTrSaEoS5S5 — — — — — cm cm co co -r seven cover slips were used for appendages from one specimen in standard groups on one microscope slide. Slides carefully sealed have endured without notice- able change for 2 yr. Body length was measured from the anterior border of the prosome to the posterior edge of the caudal rami. All figures were drawn with the aid of a Wild M20 drawing tube. Unless otherwise stated, illustrated specimens are from Arctic collections. Except for a few figures of the left swimming legs and one of the left first maxilla gnathobase, all illustrated append- ages are from the specimen's right side. First anten- nae are shown in ventral view. Maxillipeds are shown in anterior view. All other appendages are shown in posterior view. The letter after each figure legend refers to the 0.1 -mm scale to which the figure was drawn (Fig. 225). Counts of Arctic spinocalanids from below 500 m are from entire samples; most estimates of total specimens from above 500 m are based on counts from subsamples, one-half to one-twentieth of whole samples (Table 2). Many specimens were enumerated, but only the relatively few specimens in good condition were examined for descriptive pur- poses (Table 1 and lists of specimens studied). Material Studied During the systematic study of Arctic spinocalanids it was necessary to obtain specimens from other collections for comparison and for supplementing published descriptions. An effort was made to ex- amine a maximum number of critical type specimens and specimens on which widespread records are bas- ed. Arctic specimens of the present collection are listed in Table 1 and borrowed specimens are listed in Table 3. Locations for samples from dates listed in Table 1 can be interpolated from Figures 1, 2; the sample from 5 May 1968 was taken at about lat. 82°19'N, long. 158°16'W. Abbreviations used in the descriptions are listed below: Ce cephalosome; head region including the somite of the maxillipeds Thl-Th5 somites of legs 1-5 A1-A2 antennae 1-2 Mn mandible Mxl-Mx2 maxillae 1-2 Mxp maxilliped P1-P5 legs 1-5 B1-B2 basipod segments 1-2 Table 3. — Sources of borrowed specimens. Sam- Depth ple Source Program, station Location Date (m) Catalog number 1. BM RESEARCH Bay of Biscay 00 1926.12.6.125-127 2. BM Farran off Mayo 06 900 1911.11.8.37514-518 3. BM Farran off Mayo 06 1,260 1911.11.8.37508-513 4. BM Farran SR470 56°16'N, 11°27'W 26 VIII 07 720-900 1908.7.6.5 5. BM Farran W. of Ireland 09 1911.11.8.37492-499 6. BM TERRA NOVA-172 66°38'S, 178°47'W 10X1110 0-400 1930.1.1.293-404 7. BM TERRA NOVA-270 or 282 66°30'-76°S, ca. 165°W 10 0-1,000 1930.1.1.405-414 8. NZOI A452 75°35'S,173°18'W 12159 0-1,300 9. NZOI A453 75°09'S,171°00'W 13159 0-1,000 10. NZOI A455 74°22'S,178°35'W 15159 0-300 11. NZOI A459 75°17'S,172°20'E 16159 0-500 12. NZOI A462 71°15'S,176 o 30'E 20159 0-1,000 13. NZOI F945 31°20'S,165°19'E 22X68 500-1,000 14. NZOI F946 34°32'S,157°32'E 3X168 0-1,000 15. OSLO NORTH POLAR 81°27'N,ca.l23°E 22 V 94 100 F2481, F2550, F6511 16. OSLO Sars, Osterfjord 60°45'N, 5°15'E ca. 1900 400-600 F7847 17. OSLO Sars, Osterfjord 60°45'N, 5°15'E ca. 1900 400-600 F7850 18. OSLO Sars, Osterfjord 60°45'N, 5°15'E ca. 1900 400-600 F4923a 19. OSLO Sars, Osterfjord 60°45'N, 5°15'E ca. 1900 200-400 20. OSLO Sars, Osterfjord 60°45'N, 5°15'E ca. 1900 400-600 F7848-F7849 21. OSLO Sars, Orsten 62°10'N, 6°10'E ca. 1900 200-1,000 22. OSLO MICHAEL SARS-4 49°38'N, 11°35'W 11 IV 10 23. OSLO MICHAEL SARS-34 28°52'N, 14°16'W 14 V 10 24. SIO CCOFI5206-90.28D 33°28'N,117°47'W 13 VI 52 300 XVm-775 25. SIO CCOFI5504-120.50 27°34'N,115 39'W 12 IV 55 0-140 XVHI-785 26. SIO NORPAC-33C1 41°45'N,124°29'W 6 VIII 55 0-140 XVm-777 27. SIO EASTROPAC-H41 4°03'N, 81°45'W 12 XI 55 0-300 XVm-1788 28. SIO CCOFI5804-137.30 25 14'N,112 44'W 4 IV 58 0-140 XVm-776 29. SIO CCOFI5804-137.40 24°55 , N,113°24 , W 4 IV 58 0-140 xvm-i80i 30. SIO TETHYS-7 13°13'N,127°06'W 23 VI 60 0-1,000 31. SIO TETHYS-12 1°45'S,133°44'W 29 VI 60 0-1,000 Re exopod Ri endopod Rel, etc. exopod segments Ril, etc. endopod segments The prosome of most spinocalanids is fragile; many specimens are broken between Thl and Th3. There is also a tendency for the Ce ventrolateral margin to curl outward (see Spinocalanus usitatus). The sutures between Ce and Thl and Th4 and Th5, are clear on most females. Sometimes, as with S. magnus, the specimen must be cleared in glycerine to trace the un- broken suture; this suture is more easily seen in un- stained material. The caudal setae are extremely fragile, and in some species have probably never been seen intact. Usually the caudal setae are represented by short broken bases, and even the bases may be missing. All appendages are fragile. Probably most damage is caused by the net in sampling; if the loss of append- ages were due to the fixative, great numbers of detached appendages would be found in the samples, and such is not the case. Some damages probably oc- cur naturally, and the frequent abnormalities of the swimming legs are probably due to these damage below). Al of females is most often broken, usually at a suture distal to segment 12. This appendage is intact, however, on most males, where the segments beyond 10 are more or less fused. Certain extraordinarily large setae on Al are nearly always broken at the base. Esthetes are fragile and usually only indicated by their short bases. In the Al armature (Table 4) there are probably no specific differences between females of Spinocalanus and Mimocalanus (except perhaps in S. antarcticus and S. magnus: segment 15 with only 1 seta). In males, only segment 14 separates Mimocalanus and Spinocalanus species; 18 other segments have equivalent armature; 6 other segments show specific differences, and therefore do not consistently separate the 2 genera (segment 8 could never separate the genera). Al armature could not be completely verified for all species, since some specimens had only partial or no Al. Apparent deviations from Table 4 are noted and should be reexamined at each opportunity. These ap- parent deviations, except in the two species noted Table 3. Continued. Sam- Depth ple Source Program, station Location Date (m) Catalog number 32. SIC) CCOFI6106- 120.45 27°42'N,115°35'W 15 VI 61 300 XVII1-778. 779 33. SIO CCOFI6106- 120.50 27°33'N,115°50'W 17 VI 61 400 WlII-788 34. SIO CCOFI6106-120.60 27°30'N,116°31'W 13 VI 61 400 XVUI-787 35. SOSC IPHC-235C 56°31N,136°00'W 2 III 30 700-900 36. sosc IPHC-410E 55°45'N,149°50'W 30 V 31 1,300-1,500 37. SOSC IPHC-429E 55°25'N,141°12'W 7 VI 31 1,300-1,500 38. use TENCATI-155 79°42'N,170°00'W 9 VIII 67 1.000 39. USNM ALBATROSS-5120 13°46'N,120°30'E 21108 0-630 74126 40. USNM ALBATROSS-5246 6°29'N,126°19'E 15 V 08 0-180 74125 41. USNM CARNEGIE-87 18°05'S,145°33'W 1 1 III 29 UK) and/or 152 10°05'N,139°44'W 27X29 100 42. USNM ANTON BRUUN-338 2°38'S, 65°01'E 29 V 64 1,000-2,000 113514 43. USNM ANTON BRUUN-349 26°03'S, 64°58'E 25 VI 64 1.000-2,000 113513 44. USNM ANTON BRUUN-351B 29°45'S,64°58'E 28 VI 64 350-1, 710 113512 46. USNM TRIDENT-36CB5 24°33'N,69°30'W 8X66 2,000-4,000 122646 46. USNM Ellesmere Island ca.83°N,74°W 15 VI 67 0-285 122224 47. USNM ATLANTIS II- 10 36°01 N, 17°16'E 20 V 69 1,000-1,150 136818 48. WHO! DISCOVERY 11-3668 32°29'N, 20°09'W 19 III 58 3,000-4. IMHI 1334 49. WHOI DISCOVERY 11-4768 40°04'N, 19°57'W 12X61 2,000-2,800 1419 50. WHOI ANTON BRUUN-108 13°50'N, 70°07'E 25 V 63 1,000-2,000 L890 51. WHOI ANTON BRUUN-347 22°06'S,64°55 E 23 VI 64 L,000-2,000 2394 52. WHOI ANTON BRUUN-349 26°03 S, 64°58'E 25 VI 64 1,000-2,000 279S 53. WHOI CHAIN 60-1 11°03N,78°43'W 3 VI 66 505-l.tXH) 54. WHOI USNM CHAIN 60-1 I1°03N,78°43'W 3 VI 66 1.004-1.850 123780, 123784 55. WHOI USNM CHAIN 60-2 19°02N.81°57W 9 VI 66 187 960 123771. L23783, 123 - 56. WHOI CHAIN 60-2 19°02N,81°57W 9 VI 66 980 1,900 57. WHOI CHAIN 60-4 21°55N,95°26W is vi t;<; 203-600 58. WHOI CHAIN 60-4 21°55N.95°26 W 18 VI 66 500 1,000 59. WHOI CHAIN 60-4 21°55N.95°26'W IS VI 66 1,003-1,900 60. l'\V THOMPSON 50°00N,145°00W « VIII 73 330 - Table 4. — Spinocalanus and Mimocalanus, Al armature. Al segments' Genus; sex I n III IV V vi vn VIII ix x xixnxni XIV XV XVI XVII XVIII XIX XX XXI XXII xxm xxw xxv Spinocalanus, 3s 6s 2s 2s 2s 2s 2s 2s 2s Is 2s 2s Is 2s 2s 2s 2s 2s 2s Is Is 2s 2s 2s 5s Mimocalanus J le le le le le le le Female Spinocalanus; Is 6s 2s 2s 2s 2s 2s 2s 2s Is 2s 2s Is 2s Is Is Is Is Is (1) Is 2s 2s 2s 5s Male le 4e 2e le 2e or 2s 2e le 2e le or 2s 2e 2e le le le le le le or 2s le le le le le or 2s le or Is (2) Is or le le le le le Mimocalanus; Is 6s 2s 2s 2s 2s 2s 2s 2s Is Is 2s Is Is Is Is Is Is Is rt Is 2s 2s 2s 5s Male le 4e 2e le 2e le 2e le or 2s 2e le le le or 2s le le le le le le le and It le le le le 2e le le Males, genera equivalent Males, genera differ Males, species differ 's = seta; e = esthete; side (1) may be right (rt) or left (It), side (2) is the remaining side. above, occurred only in female segment 1, which is easily damaged during dissection. There are slight specific differences in Al between some Spinocalanus males, especially in relation to the merging of segment 20 to 21, which occurs on either the left or the right side, depending on the species (see Figs. 29 and 59). The merged, and apparently "lost," segment 20 may or may not be represented by 1 seta, but never by an esthete. If there is no seta on the "lost" segment 20, the complete and distinct segment 20 on the opposite Al bears either 1 seta or 1 esthete. If there is 1 seta on the "lost" segment 20, the com- plete segment 20 on the opposite Al bears 1 esthete. Other specific differences are noted in the descrip- tions. In Table 4 and the discussion of deviations, Roman numerals are Al segments; Arabic numerals are numbers of setae (s) and numbers of esthetes (e). The armature of segment 2 in females is distributed as follows (see Fig. 13): proximal, 2 setae; midlength, 2 setae, 1 esthete; distal, 2 setae. The armature of seg- ment 2 in males is distributed as follows (see Fig. 29): proximal, 2 setae, 1 esthete; midlength, 2 setae, 2 es- thetes; distal, 2 setae, 1 esthete. In addition to anterior armature, segments 22-24 have 1 posterior seta; segment 25 has 1 posterior and 2 terminal setae. Esthetes are only on the anterior edge, somewhat ven- tral to setae. Not enough material of Monacilla and Teneriforma was available to warrant generalizations for Al ar- mature; published information is also insufficient. A2 usually is unbroken, although some setae may detach. In dissecting it is difficult not to detach these setae, both while cutting the appendage and while un- tangling it from the remaining appendages. Mn palp has very long setae, especially on Re, and these are easily broken while removing the append- age. The blade is "capped" by thin, extremely fragile teeth. The main body of the blade, which is much stronger, has a foundation of "teeth," but the actual biting surfaces are the fragile "caps." The 90° twist of the Mn blade requires that the blade itself be cut across at about half its length, so that it will lie flat when mounted, but successful cutting is difficult because of the fragile teeth. Mxl setae are long and fragile. The female gnathobase is strong and seldom damaged. Mx2 has some very small setae which are difficult to see in all preparations, although most setae are strong and not too fragile. The outer seta, if present, is undoubtedly very fragile and was intact on only a few specimens. Mxp usually is present and provides important characters. Some of the armature is fragile, however, especially Bl-2 transverse spine-combs, if present, and the Ri4-5 outer setae in most species. The fragility of the swimming legs has held back the study of the Spinocalanidae. Specimens usually have complete PI, but only infrequently P2, and specimens with P3 or P4 are rarities. Ri are less often detached than Re, so that if there are a few specimens of a species in a sample, the probability is high that there will be P2 Ri or even P3 Ri. Usually the rami are detached between Rel-2 and Ril-2. The setae of the swimming legs are also fragile, and usually broken at their own joints which occur at about two-fifths the length of the seta. Spinocalanus and Monacilla species have strongly developed rows of spines and spinules on the surface (especially the posterior surface) of certain segments in the swimming legs. These spines are also rather fragile and are sometimes detached, usually leaving small scars. Sometimes a spine becomes detached while the appendage is being mounted on a slide, and the spine may lie in an unnatural position. The posi- tion, size, and shape of these spines and spine rows are rather constant within species, but there is some variability, especially in groups of smaller spines, spinules, or denticles. Sometimes there are slight differences between right and left legs in the same specimen, generally involving only a numerical difference in one or a few spines. About 10' "<• of the specimens examined had actual abnormalities of the swimming legs: a typical seta in- stead of the modified sawlike terminal seta; a short Re3 with a terminal arc of typical setae; loss of one or more outer spines (Fig. 207); or an extra seta. The other leg of the pair is usually typical, and both should be examined. Certain of these abnormalities could arise, if the fragile appendage is broken in a late juvenile stage and does not develop normally in time for the adult stage. Male P5 are usually not detached, although some species have relatively long and fragile rami. In this study, males with left Bl longer than right Bl are termed "left-handed"; conversely, "right-handed" males have right Bl longer than left Bl. Handedness appears to be species-specific, and is also correlated with the merging of segment 20 to 21 in the opposite Al. The most important diagnostic characteristics of the family and genera are in italics in the descrip- tions. The species within each genus are numbered and discussed in chronological order, including junior synonyms which are placed in parentheses. surface layer is most responsive to local conditions; it is cold (varying seasonally from -1.85° to 0°C) and dilute (varies around 30%o), and usually well mixed. The subsurface layer shows increasing salinity (to about 33°/°°). The lower Arctic Water is transitional to Atlantic Water. Freshwater, mainly from Eurasia, and low salinity water entering the Arctic from Bering Strait (sill depth about 100 m) form the Arctic Water. This layer, nearly 200 m thick, flows under the influence of the wind toward the basin exit between Spitsbergen and Greenland. Some Arctic Water also leaves through the channels between the Canadian Arctic Islands and Greenland, flowing into Baffin Bay and Davis Strait. There is a more or less closed clockwise move- ment of this layer in the Canadian Basin (Beaufort Sea), probably also a consequence of the wind pattern. The sampling platform, T-3, has been caught in this gyre. Under the general surface outflow is a subsurface inflow of rather uniform Atlantic Water. This layer has been defined as the warmer water (up to 2°C) ly- ing between the upper and lower 0°C isotherms, which occur at about 200 and 900 m. Most Atlantic Water enters the Arctic between Spitsbergen and Greenland (maximum sill depth 600-700 m, average 450-500 m) and flows eastward along the continental slope of the Eurasian Basin. There is little seasonal change in the temperature or salinity (about 35%o ). More than half of the volume of the Arctic Ocean is Arctic Bottom Water, characterized by low temperature (less than 0°C) and rather high salinity (nearly 35% o ). Most of this water is probably formed during the winter in the Norwegian Sea. The Bottom Water in the Canadian Basin is slightly warmer than that in the Eurasian Basin, since the source of the former is at or above the level of the dividing ridge (about 1,300 m). The pattern of movement of the bot- tom water is unknown. There are no really sharp boundaries between these simplified layers, especially at the far side of the Arc- tic Ocean, in the Beaufort Sea, where the Atlantic Water has had the maximum loss of its character and where water intrudes from the Pacific. THE ARCTIC SPINOCALANIDAE Arctic Hydrography Coachman (1963) has reviewed the hydrography of the Arctic Ocean. In general, there are three layers (Fig. 3), based primarily on temperature and second- arily on salinity: (1) Arctic Water, from the surface to about 200 m, (2) Atlantic Water, from 200 to 900 m, and (3) Arctic Bottom Water, from 900 m to the bot- tom (maximum depth, over 5,000 m). The uppermost layer, the Arctic Water, is itself composed of three layers: (1) surface, 0-25 or 50 m; (2) subsurface, to 100 m; and (3) lower Arctic Water. The Vertical Distribution of Arctic Spinocalanidae The low standing crop of phytoplankton in the Arctic Ocean is attributed to the low submarine light intensities; the photosynthetic production is probably the lowest of any comparable ocean area (English. 19631. Most early work on Arctic plankton, with the excep- tion of Nansen's Norwegian North Polar Expedition (1893-1896), was confined to sporadic, seasonal in- vestigations of shallow waters of peripheral sen- pecially in the Atlantic Sector, the Barents Sea. and TEMPERATURE *C 2 -1 ♦! - 1 1 1 200 400 Vi M V •OO — iff 'It '.II METERS ill H' i / / 1 / * i 1 i g i n i // i IP i /I i/l ~ KXX) z »- ft. g i i i / / / i soo - 1 1 1 1 1 1 1 1 ■ I MOO i / i / i / i / •7 = 1 I 1 1 •/ woo if 3 f j 1 I i i 1 * '» woo — i 1 £000 i 1 i 1 1 1 50 SURFACE ( ARCTIC ) WATER ATLANTIC WATER CHARLIE (I) • '77^ I60°W ^ FRAM (19) 82°N II8 W E • N.P. I (12) 87°N 0°E • BRAVO (14) 82°N I04°W BOTTOM WATER i # - I 1 i i i Figure 3. — Vertical distribution of temperature and salinity for four stations and the principal water masses of the central Arctic Ocean (from Coachman, 1963). 10 to a lesser degree, the Siberian Seas. Since World War II more attention has been given to the Beaufort and Chukchi Seas and, with the use of floating ice stations, the central Arctic. Johnson (1963a, b) and Grainger (1965) have reviewed zooplankton investigations in the central Arctic. As in other oceans, the bulk of the net- plankton consists of crustaceans, with copepods predominating. The relative quantities of copepod species depend on the depth at the localities sampled, i.e., whether in shallow coastal seas or deep basins, and also on the depth of the sample. In shallow seas the most abundant species are Pseudocalanus minutus s. 1., Acartia longiremis, and several species of Oithona and Oncaea. The predominant copepods in the surface layers of the deep basins are Calanus glacialis, C. hyperboreus, Metridia longa, Microcalanus pygmaeus s. 1., Paraeuchaeta glacialis, and Pseudocalanus minutus s. 1. With increasing depth in the central basins other species become numerically more important, es- pecially Scaphocalanus magnus, Temorites breuis, and Spinocalanus species. The number of species varies with depth, though the number of specimens decreases. About 30 species are found in the upper 300 m, nearlv 50 from 300 to 1,000 m, and about 20 below 1,000 m. Seven species of Spinocalanidae are reported from the Arctic samples of the present study: Spinocalanus longicornis, S. antarcticus, S. horridus, S. elongatus, S. polaris, Mimocalanus crassus, and M. sulcifrons. Numbers of each species in 142 samples from various depth intervals are given in Table 2. No differences in numbers of specimens or species of Spinocalanidae were attributed to differences in the mesh sizes of nets; the data from the 2 yr are not separated. None of the samples from above 80 m contained Spinocalanidae. Spinocalanus longicornis — The depth range of S. longicornis was 80-2,500 m. Adults were found from 90 to 2,500 m, juveniles from 80 to 500 m. The average concentration of total specimens was fairly uniform between 90 and 400 m, decreasing sharply above and below (Table 2). Other Arctic workers (see systematic review) have included depths less than 80 m in the vertical range of S. longicornis, but most of these records are based on sampling intervals which included greater depths. Brodsky and Nikitin (1955) reported only one specimen in each of two samples from 25 to 50 m, and two specimens from to 10 m, although the species was common below 50 m. The possibility of contamination in the 0-10 m sample should not be overlooked; many surface samples have been reported by other workers, and no other occurrences between and 10 m have been noted. Brodsky (1957) reported S. longicornis in low numbers from 25 to 50 m, but indicated that it was not present from to 10 m. Minoda (1967) reported a "very small number" of specimens, which were probably S. longicornis, from to 60 m. Hur (1968), in 399 samples collected with a plankton pump, from 5 to 185 m at 5-m intervals, found low numbers of S. longicornis in only one sample from each of the following depths: 25, 40. 55, 60, and 75 m. Spinocalanus longicornis has been considered en- demic to the Arctic, although in the present review it is reported from the Atlantic, including the Caribbean and Mediterranean, as well as from the Pacific, with a depth range of 50-2,700 m. Spinocalanus longicornis can be earned into the Atlantic in the Arctic Water; it can also be brought into the Arctic in the deeper Atlantic Water. Perhaps there is some limited exchange ol S longicornis in the near-surface water between the Pacific and Arctic. Spinocalanus antarcticus — The depth range of S antarcticus was 140-2,500 m. Adults were found from 200 to 2,500 m, juveniles from 140 to 2,000 m. The highest average concentration occurred between 180 and 300 m, with uniformly low numbers to 2,500 m (Table 2). Reports of S. antarcticus are summarized in the following systematic review. Most records are based on sampling intervals which included depths below 140 m. Brodsky and Nikitin (1955) reported one fe- male from 10 to 25 m. Johnson (1963a) found one female between and 25 m. Dunbar and Harding (1968) reported few between 0-50 m and 50-100 m. Therefore its vertical range in the Arctic is at least 10-2,500 m, in low numbers in the Arctic Water and Arctic Bottom Water and in greatest numbers in the intermediate Atlantic Water. There would seem to be no physical barriers to its movement in and out of the Arctic. Its distribution into the At- lantic has not been noted, but the species occurs in the Pacific and Indian Sectors of the Antarctic systematic review). Spinocalanus horridus — The depth range ol S horridus was 500-2,500 m. This species was found only in very low concentration (Table 2). Reports of S. horridus are summarized in the following systematic review. Most records are based on very large vertical sampling intervals. Brodsky and Nikitin (1955) reported several females between 100 and 270 m and one male between 26 and 50 m; Minoda (1967) found one probabK S horridus between and 200 m. These occurrences are rare; most records indicate an upper depth limit between 300 and 500 m. Spinocalanus horridus can freely enter the Arctic in the Atlantic Water, and a few specimens are probably carried to the Atlantic in the Arctic Water. Spinocalanus horridus has a virl worldwide distribution and a great vertical rai everywhere. 11 Spinocalanus elongatus — The depth range of S. elongatus was 300-2,500 m. Adults were found from 350 to 2,500 m, juveniles from 300 to 2,000 m. The highest average concentrations occurred between 1,500 and 2,500 m, with uniformly low numbers up to 300 m (Table 2). Brodsky (1957) reported that this species was found in low numbers from 25 to 200 m, but that it was the most numerous of all copepods between 800 and 3,000 m, where it comprised over 10% of the numbers of copepods. Dunbar and Harding (1968) reported no S. elongatus above 175 m, greatest numbers from 900 to 2,000 m, and none below 2,000 m. Spinocalanus elongatus is apparently the only species of Spinocalanidae limited to the Arctic. Its rarity in the outflowing Arctic Water would definitely reduce its opportunities to enter the Atlantic. Possibly S. elongatus occurs in the North Atlantic in small numbers; it could be confused with the superficially similar S. brevicaudatus, found in the North Atlantic but apparently not in the Arctic. Spinocalanus polaris — The depth range of S. polaris was 1,500-3,000 m. Only adults were found. The highest average concentration occurred between 2,000 and 2,500 m (Table 2). Brodsky and Nikitin (1955) reported one juvenile S. polaris between 50 and 100 m and one female between and 265 m, although most records place the bulk of this species below 1,000 m. Spinocalanus polaris is found in the North Atlan- tic, including the Caribbean, below 980 m; if it does not occur above 980 m, it could not be carried into the Arctic. However, the few Arctic S. polaris reported above 265 m provide a possibility for this species to be transported out of the Arctic. Mimocalanu8 crassus — The depth range of M. crassus was 400-2,500 m, where it was found only in low concentration (Table 2). Only two other records of M. crassus are known: from the Caribbean Sea (Park, 1970) and the Indian Ocean (Grice and Hulsemann, 1967) with a vertical distribution similar to that found in the Arctic. Mimocalanus crassus has not yet been reported from the North Atlantic, but it probably occurs there at times, from where it could enter the Arctic in the Atlantic Water. Mimocalanus sulcifrons — The depth range of M. sulcifrons was 100-2,500 m. Adults were found from 350 to 2,500 m, juveniles from 100 to 2,500 m, all in low concentration (Table 2). Only a few other specimens of this species have been reported from the Arctic, in each case as M. distinctocephalus. Hughes (1968) found only juveniles, in two samples from 80 and 90 m. Other records are based on sample intervals of 0-2,000 m (Johnson, 1963a; Dunbar and Harding, 1968) and 0-520 m (Minoda, 1967). Johnson, and later Dun- bar and Harding, in spite of the absence of this species in strictly near-surface samples, suggested that it is brought into the Arctic from the Pacific. However, M. distinctocephalus (= M. nudus) in the North Pacific has not been reported with cer- tainty above 200 m (see systematic review); therefore, it probably can not enter the Arctic by this route. Mimocalanus nudus has not been reported from the Arctic. Only a few specimens of M. sulcifrons have been reported from the North Atlantic, and only between 2,000 and 4,000 m. Juveniles might be carried out of the Arctic in low concentration in the Arctic Water. If adults in the Atlantic live only below 2,000 m, there would be no possibility for them to enter the Arctic. The most abundant of the seven Arctic species were Spinocalanus longicornis and S. antarcticus. The other species were in about equal, low Concentration. None of these species have definitely been reported at the Arctic surface, but most have been taken, though rarely, as shallow as 25 or 50 m. The depth of highest concentration of each species varies: S. longicornis (100-400 m), S. antarcticus (180-400 m), M. sulcifrons (200-400 m), M. crassus (400-2,500 m), S. horridus (500-2,500 m), S. elongatus (900-2,500 m), and S. polaris (1,500-2,500 m). All have been collected at least as deep as 2,500 m. The distribution of these species in seas adjacent to the Arctic is of great interest, although in most cases the distributions are not yet well defined. Surface layer copepods are transported from the Pacific into the Arctic (Johnson, 1963a, b) and perhaps inter- mittently in a southward surface flow, but there can be no exchange of strictly deepwater components because of the relatively shallow sill separating the two areas at Bering Strait. Both S. longicornis and S. antarcticus have vertical distributions in the Arctic which include relatively high numbers in Arctic and Atlantic Waters, and therefore there appear to be no physical barriers to their movement in and out of the Arctic. However, only S. longicornis is reported in the North Atlantic and North Pacific; S. antarcticus may yet be found there since it is known from the Antarctic. Spinocalanus horridus has been found only rarely in Arctic Water, and its movement out of the Arctic is probably restricted. It is found in the North Atlantic and undoubtedly enters the Arctic freely in the Atlan- tic Water. Spinocalanus elongatus has a vertical distribution similar to S. horridus; its movement out of the Arctic is probably restricted. It has not been reported from the North Atlantic and may be the only species of Spinocalanidae endemic to the Arctic. Spinocalanus polaris and Mimocalanus sulcifrons 12 enter the Arctic Waters in low numbers, and their movement out of the Arctic would be restricted. They have been found in the North Atlantic, but at too great depths to be transported into the Arctic. Mimocalanus erassus was not found in the Arctic Water, and therefore probably does not leave the Arc- tic. It has been found in the Atlantic (Caribbean), as well as Indian Ocean, at depths sufficient to be carried into the Arctic if it occurs in the North Atlan- tic at similar depths. Therefore, the Arctic Spinocalanidae could be characterized as follows: (1) Spinocalanus longicornis (Arctic, North Pacific, and North Atlantic, including Caribbean and Mediterranean); (2) S. antarcticus (bipolar; possibly a continuous distribution); (3) 6'. horridus (cosmopolitan); (4) S. elongatus (Arctic); (5) S. polaris (Arctic, with some transport to the deep Atlantic); (6) Mimocalanus erassus (probably cosmopolitan); and (7) M. sulcifrons (Arctic, with some transport to the deep Atlantic). In view of the great vertical ranges, and evident tolerances of these Arctic species, it is interesting to speculate on the limited distributions of some of them. If there are no physical boundaries, perhaps there are boundaries imposed by the interactions of closely related species. To examine this possibility one must consider which closely related species are present in the North Pacific and North Atlantic. Of the spinocalanids not found in the central Arctic, all North Pacific and North Atlantic Spinocalanus species except S. abyssalis, S. magnus, and S. breuicaudatus, all Mimocalanus species except M. cultrifer, all Monacilla species, and Teneriforma naso are reported at too great depths to be transported into the Arctic. Five Arctic spinocalanids have limited dis- tributions. Spinocalanus polaris has a rather unique morphology (see systematic review) and may not have an obvious counterpart in the North Atlantic. It may be more widely distributed than known at present, or it may, since it is deep living, be limited by interac- tions with other deep-living North Atlantic spinocalanids. The three other Arctic Spinocalanus species of limited distribution, S. longicornis, S. antarcticus, and S. elongatus, would encounter three very closely related species in the North Pacific and North Atlan- tic, 5. abyssalis, S. magnus, and S. breuicaudatus respectively. Spinocalanus longicornis and S. abyssalis have overlapping distributions in the North Atlantic (including the Caribbean and Mediterranean), but the other species apparently do not. The last Arctic species of limited distribution, Mimocalanus sulcifrons, may, like S. polaris, be restricted by interactions with other deep-living North Atlantic spinocalanids, especially M. cultrifer and M. nudus. FAMILY SPINOCALANIDAE VERVOORT, 1951 Description PI Bl without inner edge seta. B2 with minute .seta near outer distal edge; long anterior seta near Rj base. Re 3-segmented; Rel with or without outer distal spine, without inner seta; Re2 with 1 (niter distal spine and 1 inner seta; Re3 with I outer distal spine, 4 inner setae, and 1 bladelike terminal seta. Terminal seta outer edge lightly serrate, inner edge plumose. Ri 1 -segmented, with outer lobelike process near end. 2 terminal setae, and 1-3 inner setae; lobelike process more or less rounded, usually tipped with open, porelike points. P2 Bl with inner seta (except Mimocalanus erassus). Surfaces (except Spinocalanus polaris) and margins of B2 nude. P3 Bl with inner seta (except M erassus). B2 with surfaces and margins nude. P4 Bl with inner seta in females (except M. erassus). with or without inner seta in males. B2 margins nude. P2 Ri 2-segmented; Ril without surface armament, with 1 inner seta; Ri'2 with 2 inner, 2 terminal and 0-1 outer setae. P3-P4 Ri 3-segmented; Ril without surface arma- ment, with 1 inner seta; Ri2 outer distal corner spiniform, / inner edge seta; Ri3 with 2 inner. 2 ter- minal, and 0-2 outer setae. Number of setae on rami is identical in P3 and P4 in any one species. P2-P4 Re 3-segmented; Rel without posterior sur- face armament, with 1 outer spine and 1 inner seta; Re2 with 1 outer spine and 1 inner seta; Re3 without serrate margins, ivith 3 outer edge spines. 5 inner edge setae, and 1 serrate bladelike terminal seta. Inner caudal seta small, directed ventrally: ter- minal 4 setae well developed but very fragile; outer seta small, minute, or, usually, apparently absent. Female. — Length usually 0.8-3.58 mm, two spt 4.20-4.7 mm. Prosome in dorsal view slender to robust ovoid, widest at Thl; Ce and Thl separate: Thl-Th4 sometimes with lateral spinules in Spinocalanus; Th4-Tho usually separate, probably never completely fused; Th5 lateral corners sometimes prolonged, never pointed; rostrum 2 symmetrical or asymmetrical long strong points, or single long blunt cone, or absent; P5 absent (see Monacilla typica). Urosome 4-segmented; genital segment longest, protruding ventrally one- fourth to one-half depth of rest of urosome: anal nient length 2 .■?-! times caudal ramus: caudal ramus length 1-2 1 .' times width (one ramus sometimes longer and wider). Al extending to anal segment or beyond caudal rami by up to 4 segments (except S. angusticepa and S. validus, and Mimocalanus oralis): segments 8 and 9 usually fused, never completely separate; segment 10 not fused with 9 (partly fused in M heronae); segments 24 and 25, terminal segments, separo A2 Bl with proximal hairs or seniles and 1 long 13 plumose distal seta; B2 with 2 subequal setae; Re of equal or greater length than Ri. Re 7-segmented; Rel with 0-2 setae; Re2 and Re3 separate, partly fused, or completely fused; Re2 with or 2 setae; Re3-6 each with 1 seta; Re7 with 1 seta at about midlength and 3 terminal setae. Ri 2-segmented; Ril twice length of Ri2 (appearing much longer in M. major); Ril with 2 unequal setae; Ri2 with 2 lobes, inner lobe with 9 setae, outer with 7 setae and cluster of hairs along out- er edge. Mn B2 length about Ws times width (appearing longer in M. major); Re 5-segmented; Rel-4 each with 1 long seta; Re5 with 2 terminal setae. Ri 2- segmented; Ril without saclike appendage on inner edge, with 4 inner edge setae (S. antarcticus and S. magnus with 2, and perhaps Teneriforma naso with 3); Ri2 with 11 setae (S. antarcticus apparently with 10). Mxl B2 about size of gnathobase, with 5 inner setae (M. crassus with 7). Gnathobase with 9 strong, thick, short, spinulose marginal setae, 0, 3, or 4 proximal posterior surface setae, 1 midmarginal anterior sur- face seta, and with or without 1 small distal anterior surface seta; distal posterior surface with spinules and denticles. Inner lobe-1 (beyond gnathobase) and inner lobe-2 about equal size; lobe-1 with 4, 5, or 6 setae depending on genus; lobe-2 with 4 setae, margins with spinules or stiff hairs. Ri terminal segment usually clearly separate, with 7 setae (9 in M. crassus). Prox- imal part of Ri indistinctly segmented (probably 2 segments); setation varying with genus. Mxl outer lobe-1 usually prominent, rhomboidal; with 9 setae. Outer lobe-2 usually absent; on a few species small, nude or with minute seta. Re well-developed ovoid flap, smaller than B2, with 11 long setae; hairs on in- ner margin. Mx2 length IV2-2 times width. Lobe-1 (proximal) with terminal midmarginal hook (see Fig. 116) and 6 setae (proximal 2 setae usually small); lobes-2-4 each with 3 setae; lobe-5 with 4 setae (proximal seta sometimes much reduced); lobe-6 small or reduced, with 2 setae (1 seta small or much reduced). Ri 3- segmented; Ril -2 each with 1 long seta and usually 1 reduced posterior seta; Ri3 with 3 long setae. At least 1 seta on each of lobes-2-5, especially 4-5, thickened, shortened, and spinulose. Mx2 outer seta present or, apparently, absent. Mxp B2 equal to or somewhat longer than Bl; Bl and B2 together exceeding length of distal part. B2 with 5 setae and longitudinal row of stiff hairs, spinules, or denticles (see Teneriforma naso). Ri 5- segmented; length of Ri2 exceeding Ril; Ril-2 each with 4 setae (see T. naso); Ri3 with 3 setae (S. elongatus with 2; see also T. naso); Ri4 with 3 inner and 1 outer setae; Ri5 with 2 inner, 1 outer, and 1 ter- minal setae. Male. — Length usually 0.78-2.9 mm, one species 4.4-5.08 mm. Prosome form varying with genus and usually differing from females, widest at Ce and/or Thl; Ce and Thl separate or partly fused; Thl-Th4 without lateral spinules; Th4 and Th5 separate; Th5 at least slightly prolonged, never pointed; rostrum reduced knob, or long cone, or absent; P5 simple; B2's uneven, usually longer on left ("left-handed"), although 2 Spinocalanus spp. "right-handed"; uni- ramus or biramus. Urosome 5-segmented, segment 2 longest, segment 5 (anal segment) shortest (reduced, or length 2 /3-l times caudal ramus); caudal ramus length 1-2 times width. Al extending from distal Th4 to beyond caudal rami by up to 3 segments; segments 8 and 9 fused; seg- ment 10 separate from or fused partly or completely with segment 9; segment 20 merging with segment 21, with noticeably wrinkled margins, on right Al in left- handed species and on leftAl in right-handed species; segments 24 and 25, terminal segments, separate; seg- ment 25 reduced or not; except in Teneriforma naso, all segments beyond 10 to near end more or less fused. A2 segmentation and setation probably as in cor- responding female, although several setae reduced. Proximal hairs or setules on Bl, and hairs on Ri2, longer than on female. Re length about 2 times Ri because of short Ril. Mn B2 swollen, length about equaling width; reduced blade and reduced number and size of B2 setae; rami as female except Ri surface setules sometimes longer, and Ril setae sometimes reduced in size and number. Mxl gnathobase and other inner processes much reduced and fragile; outer processes similar to female. Mx2 lobe-sizes reduced, wider apart than in female (especially in Mimocalanus spp.); setae reduced and fragile. Mxp segmentation (see Monacilla typica) and inner setae usually as in female {Spinocalanus elongatus male with typical number of setae, and S. polaris male apparently with reduced number of setae). Bl-2 narrow, without transverse spine-combs; Ri4-5 outer setae usually reduced or absent. Swimming legs generally similar to those in cor- responding female, although surface armature (spines, spinules, setules, hairs, etc.) sometimes vary- ing slightly; number of setae on rami identical to cor- responding female. Remarks Vervoort (1951) and Farran and Vervoort (1951a) separate the Spinocalanidae from the Pseudocalanidae in which the spinocalanids had been placed by previous workers; these are the first designations of the family and include the genera Spinocalanus, Monacilla, and Mimocalanus, to which Grice and Hulsemann (1965) added Teneriforma (as Tanyrhinus Farran). The publication by Vervoort (1951) was "published in the first half of 1951," while the publication by Farran and Vervoort (1951a) was "published in the second half of 1951, possibly in Oc- tober" (W. Vervoort, pers. commun.). 14 KEY TO THE GENERA OF SPINOCALANIDAE 1. Pi Rel without outer distal spine; P2-P4 pos- terior surface of Ri2 without row of spines (3) Pi Rel with outer distal spine (reduced in male Monacilla typica); P2-P4 posterior surface of Ri2 with 2 rows of spines (1 row reduced to small spinules on a few species) .... (2) 2. Rostrum absent; male anal segment usually re- duced Spinocalanus Rostrum 2 strong points in female, reduced to knob in male; male anal segment half length of caudal ramus Monacilla 3. Rostrum absent. Female prosome length at least 4 times urosome. Caudal ramus length not more than 1 ' 2 times width. Male P5 uniramus, Re present on both legs .... Mimocalanus Rostrum 1 long cone. Female prosome length about 3 times urosome. Caudal ramus length about 2 times width. Male P5 uniramus, Re present on left leg only . . Teneriforma naso Genus Spinocalanus Giesbrecht, 1888 TYPE SPECIES: Spinocalanus abyssalis Giesbrecht, 1888. Description Rostrum absent. PI Rel with outer distal spine; Rel-2 together longer than Re3; Ri with 1-3 inner setae. P2 Ri2 with 0-1 outer seta. P3-P4 Ri3 with 2 outer setae (S. polaris with 1 outer seta). Posterior surface of certain segments of swimming legs with rows of spines (see below). Caudal rami and caudal setae sometimes asymmetrical. Female. — Prosome in dorsal view slender to robust ovoid. Th5 lateral corners sometimes prolonged; Thl- Th4 sometimes with lateral spinules, sometimes vary- ing in number left to right. Prosome length 2 3 /4-3% times urosome (S. angusticeps, S. antarcticus, and S. validus about 4Vi). Caudal ramus length 1-2 times width. Al segment 2 length about 1V& times segment 1; segments 8 and 9 fused. Al usually exceeding caudal rami by 1-4 segments (see S. angusticeps, S. magnus, and S. validus). Armature given in Table 4. A2 Re length exceeds Ri by 20 r r. Rel with 0-2 setae; Re2 and Re3 fused; Re2 with 2 setae. Mn B2 with 4 inner setae, usually also with distal row of long setules on anterior surface (and then also with row of long setules on Ril surface). Mxl gnathobase posterior surface with proximal clusters of spinules and denticles; proximal posterior surface with 3 or 4 setae; distal anterior seta present or absent. Inner lobe-1 with 6 setae. Proximal part ol Ri with 4 proximal and 5 distal setae. Outer lobe-2 ab- sent. Mx2 length IVz-VA times width; small to Strong proximal outer hump, usually armed with hairs, setules, and/or spinules. I^jbe-5 with 4 strong setae; posterior surface of base may have spines. Lobe-€ small, with 2 setae (1 seta small). Ri segments not reduced. Mx2 outer seta present or, apparently, ab- sent. Mxp Bl sometimes with transverse spine-comb midlength on outer anterior and posterior edge. B2 sometimes also with transverse spine-comb about one-third length on outer anterior and posterior edge. Bl with 3 groups of 3 setae each. B2 with longitudinal row of stiff hairs or stiff setules; 2 setae about midlength, 1 seta more distal, and 2 distal setae. Form of Ri4-5 outer setae varying with species. PI Re3 sometimes with posterior surface spines. P2-P4 Re2 distal anterior surface with 1 row of spinules. P2 Re2 posterior surface with 1 row of spines (perhaps 2 rows in S. hirtus; no spines in S. angusticeps or S. validus). P3-P4 Re2 posterior sur- face with 1-2 rows of spines. P2-P3 Re3 posterior surface with 1-3 rows of spines (none on P2 posterior surface in S. angusticeps or S. validus). P4 Re3 with or without posterior surface spines or spinules. P2-P4 Re terminal seta with moderately coarse serrate outer edge (20-50 primary teeth); outer flange narrow. P2-P4 Ri2 posterior surface with 2 rows of spines; 1 row sometimes reduced to small spinules on a few species. P3 Ri3 posterior surface usually with 2 rows of spines, but at least 1 row of spines or denticles. P4 Ri3 posterior surface with 2 rows of spines. P4 Bl with transverse row of setules (spinules on S. polaris) on posterior surface near inner seta. Male. — Prosome in dorsal view usually not ovoid: anterior irregular or rounded, with central knoblike protrusion; posterior usually somewhat rectangular. Ce expanded dorsolaterals, nearly covering Thl (only slightly in S. magnus and not at all in S. angusticeps and S. antarcticus). Ce and Thl separate or partly fused. Prosome length 1 2 /3 - 2 ^s times urosome. Anal segment usually reduced (except one-half to two- thirds length of caudal ramus in .S. angusticeps. S. ant- arcticus. S. magnus, and S. validus). Caudal ramus length 1-1 'i times width. Al segment 2 somewhat longer than segment 1; seg- ment 10 partly fused with segment 9; segment 26 reduced (except in S. angusticeps. S. antarcticu- s magnus, and perhaps S. validus). Armature given in Table 4. Swimming legs generally similar to those in cor- responding female, although surface armament sometimes variable; rows of opines on posterior sur- 15 face of some segments in female sometimes circular clusters in male. P4 Bl without inner transverse row of setules or spinules on posterior surface; with or without inner seta. P5 biramus or very asymmetrical uniramus, usually left-handed (S. polaris and S. similis right-handed); each Re essentially 3-segmented, usually right Rel and Re2 more or less fused on left-handed species. Key to the Species of Spinocalanus GROUP A. PI Ri with 2 inner setae; P2 Ri2 with 0-1 outer seta; P3-P4 Ri3 with 2 outer setae. A2 Rel with 1-2 setae. Female: Th5 lateral cor- ners prolonged or not; Thl-Th4 with or without lateral spinules. Mxp Bl with 0, 1, or 3 transverse spine-combs; B2 generally with transverse spine- comb, 2 species variable, 1 species without; Ri4-5 outer 2 setae fragile, bladelike, nude, tapered toward each end; Ri5 inner edge with spinules. Male: Anal segment reduced. Al seg- ment 25 reduced. Mxp Ri4-5 outer 2 setae reduced, may be bladelike as in female; inner setae of Ri2-5 not inflated; Ri5 inner edge with or without spinules. P5 biramus, left- or right-handed (14 species) (1) GROUP B. PI Ri with 3 inner setae; P2 Ri2 with 1 outer seta; P3-P4 Ri3 with 2 outer setae. A2 Rel without setae. Female: Th5 lateral corners prolonged; Thl-Th4 without lateral spinules. Mxp Bl-2 without transverse spine-combs; Ri4-5 outer 2 setae may be asymmetrically plumose and as large and strong as other large terminal setae, or lightly plumose, moderately fragile, thin, and relatively short; Ri5 inner edge without spinules. Male: Anal segment not reduced. Al segment 25 not reduced. Mxp Ri4-5 outer 2 setae reduced; inner setae of Ri2-5 not inflated; Ri5 inner edge without spinules. P5 left-handed; uniramus and very asymmetrical, or biramus (4 species) (17) GROUP C. PI Ri with 1 inner seta; P2 Ri2 with no outer seta; P3-P4 Ri3 with 1 outer seta. A2 Rel with 1 seta. Female: Th5 lateral corners prolonged; Thl-Th4 without lateral spinules. Mxp Bl-2 without transverse spine-combs; Ri4-5 outer 2 setae lightly plumose, moderately fragile, thin, and relatively short; Ri5 inner edge without spinules. Male: Anal segment reduced. Al seg- ment 25 reduced. Mxp Ri4-5 outer 2 setae reduced or absent; inner setae of Ri2-5 inflated; Ri5 inner edge without spinules. P5 biramus, right-handed (1 species) 26. S. polaris 1. P2 Ri2 with 1 outer seta. A2 Rel with 2 setae. Female Thl-Th4 with or without lateral spinules (10 species) (5) P2 Ri2 with no outer seta. A2 Rel with 1 seta. Thl-Th4 without lateral spinules (4 spe- cies) (2) 2. P3-P4 Re2 proximal posterior surface spines broad, leaflike. Female: Th5 rounded, not prolonged. Mxp Bl without transverse spine-comb. Male: P5 longest Ri about equal to or less than IV2 times length of other Ri (4) P3-P4 Re2 proximal posterior surface spines relatively narrow, not leaflike. Female: Th5 slightly prolonged. Mxp Bl with 1 or 3 transverse spine-combs. Male: P5 long- est Ri more than 1 V2 times length of other Ri (3) 3. Female: Al terminal 2 segments elongate. Mxp Bl with 3 transverse spine-combs; Ri3 with 2 setae. Male: Left-handed . . . 25. S. elongatus Female: Al terminal 2 segments of moderate length and width. Mxp Bl with 1 transverse spine-comb; Ri3 with 3 setae. Male: Right-handed 21. S. similis 4. Female: Prosome, slender ovoid, length great- er than 2.4 times width. Male: P5 Ri lengths very unequal .... 1. S. abyssalis Female: Prosome, somewhat robust ovoid, length less than 2.4 times width. Male: P5 Ri lengths about equal .... 2. S. longicornis 5. Female (8) Male (unknown for 7 of 10 species) .... (6) 6. P5 Ri not reaching base of Re3 terminal setae 10. S. horridus P5 Ri reaching beyond base of Re3 terminal setae (7) 7. Pi Re3 posterior surface without spines . . 27. S. breuicaudatus PI Re3 posterior surface with spines .... 43. S. terranouae, new species 8. Th5 lateral corners rounded (10) Th5 lateral corners more or less prolonged, not rounded (9) 9. Length less than 2 mm. Mxp B2 without transverse spine-comb. P1-P4 Bl surfaces without dense covering of fine hairs. (Male unknown.) 31. S. abruptus Length greater than 2 mm. Mxp B2 with transverse spine-comb(?). P1-P4 Bl sur- faces with dense covering of fine hairs. (Male unknown.) 8. S. hirtus 10. At least 1 segment Thl-Th4, right and/or left, with ventrolateral spinules (13) Thl-Th4 without spinules (11) 11. Mxp Bl with transverse spine-comb. (Male unknown.) 39. S. aspinosus Mxp Bl without transverse spine-comb . (12) 12. PI Re3 posterior surface without spines. P2 Bl surface without spines (spinules and/or setules only). P3 Re2 posterior surface with row of spines distal from row of strong 16 spines. P4 B2 surface nude or with thin setules 27. S. breuicaudatus PI Re3 posterior surface with spines. P2 Bl surface with strong spines. P3 Re2 posterior surface with row of denticles distal from row of strong spines. P4 B2 surface with spin- ules 43. S. terranouae, new species 13. Mxp Bl without transverse spine-comb. (Male unknown.) 9. S. spinosus Mxp Bl with transverse spine-comb . . . (14) 14. Mx2 posterior surface of lobe-5 base with spines. (Male unknown.) . . 37. S. usitatus Mx2 posterior surface of lobe-5 base without spines (15) 15. Mxp Bl transverse spine-comb on conspicu- ous protuberance. (Male unknown.) . . . 38. S. hoplites Mxp Bl transverse spine-comb not on con- spicuous protuberance (16) 16. Length greater than 1.7 mm. Prosome elon- gate ovoid 10. S. horridus Length less than 1.7 mm. Prosome robust o- void. (Male unknown.) 40. S. oligospinosus 17. Female: Al reaching at least to anal segment. Mxp Ri4-5 outer 2 setae asymmetrically plumose and as large and strong as other large terminal setae. P2 Re2-3 posterior sur- face with 1 row of spines. P4 Re3 posterior surface nude. Male: P4 Bl with inner seta (19) Female: Al not reaching beyond genital seg- ment. Mxp Ri4-5 outer 2 setae lightly plu- mose, moderately fragile, thin, and rela- tively short. P2 Re2-3 posterior surface nude. P4 Re3 posterior surface with many small spinules. Male: P4 Bl without inner seta (18) 18. Length greater than 3.3 mm. Female: Pro- some slightly robust ovoid. P2 Rel outer spine typical. Male: Ce expanded dorso- lateral^, nearly covering Thl. P5 biramus 12. S. validus Length less than 3.3 mm. Female: Pro- some long slender ovoid. P2 Rel outer spine forming strongly bent hook. Male: Ce not expanded. P5 essentially uniramus (ru- dimentary left Ri), very asymmetrical .... 13. S. angusticeps 19. Female: Prosome in dorsal view slender long ovoid; in lateral view with gentle, uneven forward slope. Prosome length 3 l A times urosome. Male: Ce slightly expanded dorsolaterally. P5 biramus . . 4. S. magnus Female: prosome in dorsal view robust ovoid; in lateral view with abrupt, even forward slope. Prosome length 4U times urosome. Male: Ce not expanded. P5 uniramus, very asymmetrical 5. S. antarctieus 1. Spinocalanus abyssalis Giesbrecht, 1888 (Figures 4-10, UH) Spinocalanus abvssulis Giesbrecht, 1888, p. 335. — Giesbrecht, 1892, p. 209-213, pi. 13, fig. 42-48; pi. 36, fig. 49. — Giesbrecht and Schmeil, 1898, p. 31. — ?Thompson, 1903, p. 16, pi. 1, fig. 6. —van Breemen, 1908 (part), p. 28-29, fig. 27d (only). — Brodsky, 1950, p. 129-130, fig. 49. —Park. 1970, p. 481-482, fig. 13-22. — Grice, 1971, p. 278- 280, fig. 31- J. Spinocalanus abyssalis var. pygmaeus. — Farran and Vervoort, 1951a (part), p. 2-3, fig. If, h(only). Not S. abyssalis var. pygmaeus Farran, 1926. Spinocalanus parabyssalis Park, 1970 (part), p. 483- 485, fig. 34-36; male only. TYPE SPECIMEN: Neotype, female (1.20 mm) USNM 142700. TYPE LOCALITY: Central Pacific, 1,000-4,000 m. Material Studied: 1 female (1.0 mm), sample 8 (Table 3); 1 female (1.13 mm), sample 11; 3 females (1.23, 1.25, 1.27 mm), sample 22; 1 female (0.875 mm), sample 24; 2 females (0.98, 1.025 mm), sample 25; 1 female (0.90 mm), sample 28; 1 female (1.025 mm), sample 29; 3 females (1.10, 1.23, 1.30 mm) and neo- type female (1.20 mm), sample 30; 2 females (1.08, 1.22 mm), sample 31; 1 female (1.30 mm) and 1 male (1.13 mm), sample 55. Description Pi Ri with 2 inner setae; P2 Ri2 with no outer seta. Female.— Length (0.8?-) 0.85-1.32 mm (Figs. 4. 5). Prosome in dorsal view slender ovoid; prosome length greater than 2.4 times width. Head rounded in dorsal view, in lateral view with flat, gentle forward slope. Th4 and Th5 may be separate or partly fused, es- pecially dorsolaterally; Th5 lateral corners rounded, not prolonged; Thl-Th4 without lateral spinules. Prosome length slightly more than 3 times urosome. Genital segment slightly longer than wide, protruding ventrally one-half depth of rest of urosome; caudal rami length nearly 2 times width, symmetrical. Caudal setae incomplete on all specimens examined, bases symmetrical. Supra-anal plate with fringe of long hairs. Al exceeding caudal rami by about 4 segments; ter- minal segments (Fig. 148) of moderate length and width. Apparent deviation: I-2s; remainder as Table 4. A2 Rel with 1 seta [Giesbrecht (1892) reported 2 setae]. Mn with row of setules on distal anterior surface of B2 and on surface of Ril. Blade with 4 strong, evenly spaced ventral teeth, 4 smaller closer spaced dorsal teeth, and dorsal seta. 17 Spinocalanus abyssalis, female Figure 4. — Habitus, dorsal view; sample 30 (Table 3); scale B. Figure 5. — Habitus, lateral view; sample 30; scale B. Figure 6. — Mxp; sample 30; scale D. Figure 7.— PI; sample 30; scale D. Figure 8.— P2; sample 30; scale D. Figure 9.— P3; sample 30; scale D. Figure 10.— P4; sample 30; scale D. 18 Mxl gnathobase with 3 proximal setae (Park (1970, fig. 17) shows 4] on posterior surface; without seta on distal anterior surface. Mx2 with moderately strong, setule-covered hump on proximal outer edge; outer seta present; posterior surface of lobe-5 base with spines. Mxp (Fig. 6) Bl without spine-comb; longitudinal row of stiff hairs on midanterior surface; outer edge with 2-3 clusters of short setules. Transverse spine- comb of 6-7 moderate spines present or not on B2. B2 with longitudinal row of stiff hairs. Two outer setae on Ri4-5 nude, bladelike, tapering toward each end; spinules on inner edge of terminal Ri5. Pi (Fig. 7) Bl surface with few setules; inner margin with hairs. B2 surface with few setules and hairs; inner margin with hairs. Rel anterior surface with outer distal row of spinules; Re2 surface nude; Re3 posterior surface with 0-4 spines. Rel outer spine reaching to or beyond base of Re2 outer spine; Re2 outer spine reaching beyond base of Re3 outer spine. Ri surface nude. P2 (Fig. 8) Bl surface nude or with few setules, in- ner margin with hairs. P3 (Fig. 9) Bl surface with few setules, inner margin with hairs. P4 (Fig. 10) Bl sur- face with few thin setules in addition to posterior transverse row; inner margin with hairs; B2 nude. Re2 posterior surface on P3 with proximal row of ca. 8 very broad, flat, leaflike spines and distal row of 5-6 denticles; on P4 with proximal row of ca. 9 sim- ilar leaflike spines and distal row of ca. 6 small spines. Re3 posterior surface on P2 with 3 rows of 2-4 spines each; on P3 without proximal row of spines, but with 2 distal rows of a few small spines; on P4 with 3 rows of up to 5 small spines. P3 Ri3 posterior surface with 2 rows of spines. Re terminal seta with following number of primary teeth: P2 (24-38); P3 (27-34); P4 (25-29). P2 terminal seta slightly longer than Re3; P3 terminal seta somewhat shorter than Re3. Male. — Length 0.90-1.2 mm. Prosome anterior irregular in dorsal view, posterior somewhat rounded; in lateral view with gentle forward slope. Ce and Thl partly fused. Prosome length 2 times urosome. Caudal rami length slightly more than width, symmetrical. Caudal setae unknown. Al reaching about end of Th4. P4 Bl proximal outer surface with setules; without inner seta. P5 biramus, left-handed; left Bl reaching one-third length of right B2; left B2 reaching middle of right Rel. Left leg longest; left Re (including terminal blade) longest; right Ri longest. Order of length, longest to shortest, of Re segments: left 1, 2. 3; right 1 = 2, 3. Left Ri reaching middle of left Re2; right Ri reaching middle of longest terminal seta of right Re. Each Re with 1 small and 1 moderate bladelike ter- minal setae. Remarks Giesbrecht (1888, 1892) examined "about a hall- do/.en" mutilated females on which he based his description of the first Spinocalanus sp< abyssalis. He did not illustrate the prosome; his figures of appendages, characteristically excellent, differentiated S. abyssalis from the other Pseudocalanidae with which it was placed. However, the lack of information on body form and probably also the lack of a figure of P2 (even though Giesbrecht stated that P2 Ri2 had only 4 setae) led subsequent investigators to unite more or less closely related species with S. abyssalis. It is a curious coincidence that Sars (1900) mistakenly illustrated P2 Ri2 of Arc- tic S. longicornis with 5 setae (rather than 4), and later (1903) mistakenly illustrated the same segment on Norwegian S. breuicaudatus with 4 setae (rather than 5). Sars also failed to illustrate the surface spines of the swimming legs of 6'. breuicaudatus, and when he claimed that these three species were identical, it became effectively impossible to reconcile the descriptions solely from the literature. Farran (1926) proposed Spinocalanus abyssalis var. pygmaeus for the smaller of at least two forms, thereby making the larger form the "typical" species, even though Giesbrecht's specimens were small. This was repeated by most investigators and served to further confound the concept of "Spinocalanus abyssalis." Specimens examined in this study agreed essential- ly with the description and figures of Giesbrecht (1892). Giesbrecht's unit in measuring Al segments should have read 0.005 mm, not 0.05 mm. The Mxp he illustrated lacked the fragile outer seta on Ri4. Giesbrecht stated that PI Re3 posterior surface may or may not have spines, though the illustrated Pi lacked these spines. References to S. abyssalis by Giesbrecht and Schmeil (1898), van Breemen (1908. part), and Brodskv (1950) are solelv repetitions of the work of Giesbrecht (1892). Thompson (1903) illustrated P5 of a juvenile male from the west of Ireland; no size was reported. The figure does not agree with P5 of male stage V S longicornis (see Fig. 34) or S. breuicaudatus (With. 1915. fig. 15e). Thompson's specimen could be S. abyssalis, although possibly it is not even a Spinocalanus species. Park's (1970) description and figures ofS abyssalis agree with Pacific specimens and include the first habitus illustrations of this species. Grice'a ( 1971 1 figures also agree with the description of S. abyssalis. The references to these figures are cor- rect in his key, but the figure legend transposed the caption for S. abyssalis and S. parabyssalis (G. Grice, pers. common.). Farran's (1926) first use of S. abyss pygmaeus referred at least to >'. \OTU\ but possibly also to N. abyssalis since hot i , ur in 19 the North Atlantic (see S. longicornis) . The descrip- tion of S. abyssalis var. pygmaeus by Farran and Ver- voort (1951a, part) is from Giesbrecht (1892). Park (1970) described a new species, S. parabyssalis, the female of which has been identified in the present review as S. longicornis. The male, however, does not agree with the male S. longicornis, but is consistent with what would be likely for the male of S. abyssalis. It is therefore believed that S. parabyssalis male is the male of S. abyssalis. Park reported one male S. parabyssalis; a male which he reported as S. breuicaudatus (1.38 mm) has been identified in the present study as a second male S. abyssalis (1.13 mm); both males were from the same sample, with numerous female S. abyssalis and S. longicornis (sample 55, Table 3). Giesbrecht's type specimens appear to have been lost (Bruno Scotto di Carlo, pers. commun.). In view of the high number of common characteristics of S. abyssalis and S. longicornis, a neotype of S. abyssalis has been deposited in the U.S. National Museum (USNM 142700). The neotype (sample 30, Table 3) is from nearly the same area and depth as Giesbrecht's specimens; this female is consistent with what is known of the original type material and corresponds to the description given here. Of the material examined in the present study, S. abyssalis females from samples 8 and 22 had separate Th4 and Th5 (4 specimens); those from samples 11, 24, 25, 28-31, and 55 had partly fused Th4 and Th5 (13 specimens). A Mxp B2 spine-comb was absent on 7 specimens from samples 30, 31, and 55; a B2 spine- comb was present on 9 specimens from samples 8, 11, 22, 24, 25, and 29. Distribution Many records of Spinocalanus abyssalis cannot be interpreted; unless at least the length of specimens was given, the record could reflect one or a mixture of several species. If lengths were given, a record might be presumed based on the known distribution of the species. Specimens reported from transitional areas should be reexamined. In reports of size groups and in specimens examined in this study, there is a clear dis- continuity between females of the "pygmaeus" form (S. abyssalis and S. longicornis) and the "typical" form (S. breuicaudatus or other species), the former smaller, and the latter larger, than 1.40 mm (see S. breuicaudatus for several records of "S. abyssalis" not mentioned here). Probable or confirmed occurrences of S. abyssalis (reported as S. abyssalis with length range given, and/or S. abyssalis var. pygmaeus) are listed below. Those with superscript probably or definitely include other species — see distribution of (a) S. breuicaudatus, (b) S. horridus, and (c) S. terranouae: Pacific Ocean North: Minoda (1971)\ 160-2,000 m. Northeast: Fleminger (1967), 0-140 m. —von Vaupel-Klein (1970) a , 0-1,200 m. Central: Giesbrecht (1888, 1892), 1,000-4,000 m. —Present study, 0-1,000 m. Northwest: Tanaka (1953, 1956) a , below 200 m. —Minoda (1971) b , 457-500 m. Southwest: Farran (1936), ca. 300 m. Indo-Pacific: Vervoort(1946) a , 340-2,500 m. Indian Ocean Northwest: Sciacchitano (1930), 1,000 m. West: Grice and Hulsemann (1967), 750-4,000 m. Atlantic Ocean Northeast: Present study. Eastern Mediterranean Sea: Grice (1971), 1,050- 1,200 m. Southeast: Unteruberbacher (1964)\ 0-100 m. Caribbean Sea: Park (1970), 203-2,800 m. Antarctic Pacific Sector: Farran (1929)° , 0-400 m. —Brad- ford (1971) c , 0-1,300 m. Indian Sector: Vervoort (1957, 1965)° , 250-1,000 m. —Tanaka (1960) c , 0-400 m. Atlantic Sector: Hardy and Gunther (1935), 500- 2,000 m. Records of small species (usually reported as S. abyssalis var. pygmaeus) from the North Pacific, North Atlantic, or Mediterranean are complicated because of the co-occurrence of S. abyssalis and S. longicornis. Farran (1908), Grice (1963), Grice and Hulsemann (1965), Wheeler (1970), Vives (1970, 1971), and Roe (1972a, b) may have examined either or both species. Some specimens as short as 0.8-0.83 mm may be either or both species. Some of these records also include larger forms, probably S. breuicaudatus. Some records of S. abyssalis contain no information which would help to determine the species: Wolfenden (1904, 1911), Farran (1905), Sars (1907), Lysholm and Nordgaard (1945), Furuhashi (1961, 1966), Shmeleva (1964), and Vinogradov (1970). Wilson's (1942) record of S. abyssalis appears to be erroneous, inasmuch as the samples referred to, and labeled by Wilson, contained no Spinocalanidae. Wilson's (1950) samples from Hawaii and Peru also contained no Spinocalanidae; the other two samples, from Bering Sea and North Pacific, were not located. 2. Spinocalanus longicornis Sars, 1900 (Figures 11-34, 149) Spinocalanus longicornis Sars, 1900 (part), p. 75-78, pi. 22, fig. 1-12; female only. —Mrazek, 1902, p. 521. — Brodsky, 1950, p. 133-134, fig. 54. — Vidal, 1971, p. 19, 24, fig. 48-49, 52-55. Spinocalanus schaudinni Mrazek, 1902, p. 509-512, 521, pi. 4, fig. 1-12; pi. 5, fig. 1. — van Breemen, 1908 (part), p. 29, fig. 28 (only). 20 Spinocalanu* longicornti, fimale Figure 11.— Habitus, dorsal view; scale B. Figure 12.— Habitus, lateral view; scale B. Figure 13.— Al; scale E. Figure 14.— Mn blade; scale F. Figure 15. — Mxl; scale E. Figure lti. — Mx2; scale E. Figure 17. — Mxp; scale E. 21 Spinocalanus longicornis Figure 18.— Female, PI; scale E. Figure 19.— Female, P2; scale E. Figure 20.— Female, P3; scale E. Figure 21.— Female, P4; scale E. Figure 22.— Male, A2; scale E. Figure 23.— Male, Mn blade; scale F. Figure 24. — Male, Mn blade; scale F. Figure 25. — Male, Mxl gnathobase; scale F. Figure 26.— Male, Mx2; scale E. 22 30 Spinocalanua longicornis. male Figure 27. — Habitus, dorsal view, scale B. Figure 28.— Habitus, lateral view, scale B. Figure 29.— Al; scale E. Figure 30. — Mxp; scale E. Figure 31.— P4. left; scale D. Figure 32.— P5; scale E. Figure 33.— P5; sample 23 (Table 3); scale D. Figure 34.— Stage V, P5; sample 15; scale D. 23 Spinocalanus abyssalis var. pygmaeus Farran, 1926, p. 243 (at least part). Spinocalanus parabyssalis Park, 1970 (part), p. 483- 485, fig. 23-33; female only. — Grice, 1971, p. 278- 280, fig. 3E-F. Type Specimen: Holotype, female, on slide F6511, OSLO. TYPE LOCALITY: Central Arctic Ocean, Eurasian Basin, ca. 100 m. Material Studied: 2 females (0.95, 1.08 mm), sample 1 (Table 3); holotype female, 38 paratype females (0.92- 1.05 mm, mean = 0.97 mm), 10 damaged females, 5 male stage V (0.88-0.95 mm, mean = 0.91 mm), and 2 damaged male stage V from sample 15; 6 females (0.95-1.10 mm, mean = 1.03 mm), 1 female stage V (0.88 mm), and 1 male (1.08 mm), sample 23; 1 female (0.95 mm), sample 46; 1 female (0.88 mm), "Spinocalanus parabyssalis Park, holotype, 0.98 mm," sample 55; 6 females (1.08-1.24 mm, mean = 1.18 mm), sample 60; 2 females (1.08, 1.1 mm), sam- ple Al (Table 1); 1 female (1.05 mm), sample A2; 1 female (1.15 mm), sample A4; 2 females (1.08, 1.30 mm) and 1 male (1.25 mm), sample A6; 1 male (1.28 mm), sample A21; 1 male (1.36 mm), sample A22; 4 males (1.24-1.40 mm, mean = 1.31 mm), sample A23; 1 male (1.24 mm), sample A27; 5 females (1.15-1.20 mm, mean = 1.17 mm), sample A33; 5 females (1.12- 1.24 mm, mean = 1.17 mm) and 2 males (1.36, 1.40 mm), sample A34; 2 males (1.36, 1.40 mm), sample A35; 3 males (1.28, 1.36, 1.40 mm), sample A36; 6 females (1.08-1.21 mm, mean = 1.14 mm), sample A48; 1 female (1.15 mm), sample A50. Description Pi Ri with 2 inner setae; P2 Ri2 with no outer seta. Female.— Length (0.8?-) 0.88-1.30 mm (Figs. 11, 12). Prosome in dorsal view somewhat robust ovoid; prosome length less than 2.4 times width. Head slight- ly truncate in dorsal view, in lateral view with gentle forward slope. Th4 and Th5 usually separate, sometimes partly fused; Th5 lateral corners rounded, not prolonged; Thl-Th4 without lateral spinules. Prosome length about 3 l A times urosome. Genital seg- ment slightly wider than long, protruding ventrally one-half depth of rest of urosome; caudal rami length IV2 times width, symmetrical; cluster of spinules on outer dorsal surface. Caudal setae symmetrical; inner setae moderately long. Supra-anal plate without fringe of hairs. Al (Fig. 13) exceeding caudal rami by nearly 4 segments; terminal segments (Fig. 149) of moderate length and width. Apparent deviation: I-2s; remainder as Table 4. A2 Rel with 1 seta. Mn palp with inner row of setules on distal anterior surface of B2 and on surface of Ril. Blade (Fig. 14) with 4 strong, evenly spaced ventral teeth, 4 smaller, closer spaced teeth, and dorsal seta; proximal surface with cluster of spinules. Mxl (Fig. 15) gnathobase with 3 proximal setae on posterior surface, without seta on distal anterior sur- face. Mx2 (Fig. 16) with small hump on proximal outer edge, armed with spinules, setules, and hairs; outer seta present (not seen on Arctic specimens); posterior surface of lobe-5 base with spines. Mxp (Fig. 17) Bl without spine-comb; longitudinal row of stiff hairs on midanterior surface; outer distal edge with 2 posterior rows of short setules. Transverse spine-comb of 5-6 short spines present or not on B2. B2 with longitudinal row of stiff hairs. Outer 2 setae on Ri4-5 nude, bladelike, tapering toward each end; spinules on inner edge of Ri5. PI (Fig. 18) Bl anterior surface with outer distal row of spinules. B2 inner anterior surface with a few spinules; inner margin with hairs. Rel anterior sur- face with outer distal row of spinules; Re2 surface appearing nude; Re3 surface nude. Rel outer spine usually reaching beyond Re2; Re2 outer spine reaching beyond base of Re3 outer spine. Ri surface nude. P2 (Fig. 19) Bl surface nude, or with few setules, in- ner margin with hairs. P3 (Fig. 20) Bl surface with spinules and denticles, inner margin with hairs. P4 (Fig. 21) Bl surface, except for typical transverse row of setules, nude; inner margin with hairs; B2 nude. Re2 posterior surface on P3 with proximal row of ca. 8 very broad, flat, leaflike spines and distal row of 5-6 denticles; on P4 with proximal row of ca. 7 similar leaflike spines and distal row of 5-6 small spines. Re3 posterior surface on P2 and P3 without prox- imal row of spines, but with 2 distal rows of 2-4 small spines; on P4 without surface spines. P3 Ri3 posterior surface with 2 rows of spines. Re terminal seta with following number of primary teeth: P2 (31-44); P3 (30-43); P4 (24-35). P2 terminal seta more than IV2 times length of Re3; P3 terminal seta slightly longer than Re3. Male.— Length 1-1.40 mm (Figs. 27, 28). Prosome anterior irregular in dorsal view, posterior somewhat rounded; in lateral view with gentle forward slope. Ce and Thl partly fused. Prosome length 2 times urosome. Caudal rami length slightly more than width, symmetrical. Caudal setae as on female. Al (Fig. 29) reaching about end of Th4. IV- 2s,2e; VIII-2s,2e; XV-ls,le; XIX-ls,le; right XX-ls; left XX-le; remainder as Table 4. A2 (Fig. 22) as on female except for typical dimorphic characteristics. Mn blade (Figs. 23, 24) typically reduced, showing some slight variability. Mxl gnathobase (Fig. 25) reduced, 3 or 4 sinuate prongs. Mx2 (Fig. 26) typically reduced; proximal setae 24 smaller than on female; setae of lobe-5 nude; large setae distal from lobe-5 wider and more coarsely spinulose than on female, especially 1 of terminal 3 setae and largest seta of lobe-6. No spines on lobe-5 base; no outer seta observed. Mxp (Fig. 30) somewhat reduced; several setae of reduced size, especially on Bl-2 and outer edge of Ri4- 5, but number as on female. Bl surfaces nude. B2 relatively thinner than on female, with longitudinal row of stiff hairs. Ri5 inner edge without spinules. Pi similar to female PI except Re outer spines wider, shorter, and smoother. Ri outer lobe rounded. P2 similar to female P2 except Bl outer surface with long thin setules, and Re outer spines longer than on female. P4 (Fig. 31) Bl nude; Ri as on female, except more spines in each row. Re2 posterior surface with row of ca. 8 leaflike spines followed distally by more or less circular patch of ca. 15 small spines; Re3 with ca. 17 denticles in 3 groups on posterior surface; serrations on terminal seta finer than on female, ca. 50 primary teeth. P5 (Figs. 32, 33) reaching to end of urosome seg- ment 2; biramus, left-handed. Left Bl reaching mid- dle of right B2; left B2 reaching two-thirds length of right Rel. Left leg longest; left Re, including terminal blade, longest; right Ri not much longer than left. Order of length, longest to shortest, of Re segments: left 2, 1, 3; right 1 = 2 = 3. Left Ri reaching one-third length of left Re2; right Ri reaching to right Re3. Probably each Rel with short, flat seta on outer distal edge (not seen on Arctic specimens). Right Rel -2 more or less fused. Inner edge of left Re 1-2 with long hairs. Each Re with 1 small and 1 moderate bladelike terminal setae. Male stage V.— Length 0.88-0.95 mm. P5 (Fig. 34) biramus, symmetrical. Re 1-segmented with trace of segmentation about one-third length; 2 unequal ter- minal setae. Ri 1-segmented, thin distal portion reaching about two-thirds length of Re. Remarks Sars (1900) did not describe the well-developed spines on the posterior surface of P2-P4 Re, even though the material he examined is as described above. Sars mistakenly illustrated an outer seta on P2 Ri2; his slide (OSLO F6511) only has 1 P2, which was mounted so that the inner seta of Rel met the Ri at the base of a surface spine, giving the appearance of a fifth seta on Ri2. Sars' slide was remounted and it was seen that there is no outer seta on P2 Ri2 of the holotype or on any other S. longicornis studied. Sars' (1900) description of a male S. longicornis was pointed out first by Mrazek (1902) not to be of the male, and later Sars (1903, p. 157) confirmed that it was a Microcalanus species. A vial from the Norwegian North Polar Expedition (F2482) labeled by Sars "SpinocalanilS longicomu" contained only one male Microcalanus species; sample 15 (Table 3) also contained, in addition to S. longicornis, one male, one male stage V, and two female Microcalanus species. Mrazek (1902) repeated Sars' record of S. longicor- nis. Mrazek also described the male and female of a new species, .S'. schaudinni, which is considered here as conspecific with S. longicornis. Mrazek's descrip- tion provided much of what Sars' lacked, and led Mrazek to believe that the two species were distinct. Brodsky (1950) was the first to state that S. longicornis was probably equivalent to S. schaudinni, and at the same time distinct from both S. abyssalis Giesbrecht and S. longicornis/ S. abyssalis. — Sars. 1901, 1903 (see S. brevicaudatus) . Brodsky reported that the outer spines on PI Re do not reach the base of the following spines; specimens in the present study had longer spines, and this feature is considered variable. Farran's (1926) S. abyssalis var. pygmaeus has been assumed by Park (1970) and Grice (1971) to be equivalent to S. abyssalis Giesbrecht. Some of Farran's specimens were examined in the present study (sample 1, Table 3) and were determined to be S. longicornis. It is possible, however, that Farran in- cluded both species in this variety, since they co-occur in the North Atlantic. Park's (1970) S. parabyssalis holotype was deter- mined to be 5. longicornis; Park's description and figures of the female are consistent with the above description of S. longicornis. The male S. parabyssalis, however, is believed to be S. abyssalis. Grice's (1971) figures of 5. parabyssalis also agree with the description of S. longicornis. The references to these figures are correct in his key, but the figure legend transposed the caption for S. abyssalis and S, parabyssalis (G. Grice, pers. commun.). Of the material examined in the present study, the suture between Th4 and Th5 was clear on most of the Arctic S. longicornis females, and on the two females from sample 1. These segments were partly fused on the S. parabyssalis holotype, from sample 55, and on the six females from sample 23. Mxp B2 spine-comb was present right and left on the specimens in sample 1, and on 4 Arctic specimens; on left but not right on 2 Arctic specimens; absent on the holotype from sample 15, the 6 females from sam- ple 23, and 12 other Arctic specimens. Distribution Some records of S. abyssalis or S. abyssalis var. pygmaeus may be based on either or both S. abyssalis and S. longicornis (see discussion under N. abyssalis): these include specimens as short as 0.8-0.83 mm. Probable or confirmed occurrences of S. longicornis are summarized below. Bogorov (1946a) recognized two species, based primarily on size. S. longicornis and S. abyssalis: presumably the latter was the larger (see S. clongalus). Bogorov identified the material 25 reported by Shirshov (1938). Grice (1962) reported S. abyssalis var. pygmaeus, which was probably S. longicornis. Scotto di Carlo (1968) and Hure and Scot- to di Carlo (1968, 1969) reported S. abyssalis var. pygmaeus which they later (Hure and Scotto di Carlo, 1971) equated with S. parabyssalis Park. Hughes' (1968) specimens were examined and found to be S. longicornis. G. A. Heron (pers. commun.) identified S. longicornis (females, males, and juveniles) from Weather Station "P" in the northeast Pacific, from four 55-m depth intervals between 220 and 440 m; six females were examined in the present study. Arctic Ocean Central: Bogorov (1946a), 10-750 m. — Brodsky (1950, 1957), 25-3,000 m. Eurasian Basin: Sars (1900), 100-500 m. — Mra- zek (1902), 0-100 m. —Shirshov (1938), 3-1,000 m. —Grice (1962), 110 m. Canadian Basin: Bogorov (1946b), 0-300 m. — Brodsky and Nikitin (1955), 0-3,260 m. —Hughes (1968), 25-185 m. — Vidal (1971). —Present study, 80-2,500 m. Pacific Ocean Northeast: von Vaupel-Klein (1970), 0-1,200 m. —Present study, 220-440 m. Atlantic Ocean Northeast: Farran (1926), 200-2,700 m. East: Present study. Mediterranean Sea: Scotto di Carlo (1968), 200- 1,000 m. —Hure and Scotto di Carlo (1968, 1969, 1971), 50-1,000 m. —Grice (1971), 750- 1,400 m. Caribbean Sea: Park (1970), 200-950 m. Spinocalanus abyssalis has been reported from the Arctic without size indications: Bernstein (1934) and Khmyznikova (1936) for the Kara Sea; Johnson (1963a) and Minoda (1967) for the Central Arctic; and Jaschnov (1948), repeating the Kara Sea and Sars' Polar Basin records. Jaschnov's figures are copied from Sars' (1901, 1903) figures of S. brevicaudatus . These records may represent S. longicornis, but they may also include S. elongatus or other species. Norman's (1905) reference to specimens of S. longicornis obtained from Sars could be either to Arc- tic S. longicornis or Norwegian S. brevicaudatus. Ostenfeld (1909) reported S. longicornis from the Atlantic (identified by Farran) and the North Sea (identified by T. Scott); both records could include S. brevicaudatus. Von Vaupel-Klein's (1970) record of a male S. longicornis from the northeast Pacific is considered here (see also S. abyssalis). (3. Spinocalanus schaudinni Mrazek, 1902) See Spinocalanus longicornis Sars, 1900. 4. Spinocalanus magnus Wolfenden, 1904 (Figures 35-42, 150) Spinocalanus magnus Wolfenden, 1904, p. 118. — Far- ran, 1905, p. 30-31, pi. 3, fig. 1-12. —Wolfenden, 1906, p. 41-43, pi. 14, fig. 1-5. —van Breemen, 1908, p. 29, fig. 29. —Wolfenden, 1911, p. 216-217, fig. 8; pi. 25, fig. 3-5. —Sars, 1924, 1925, p. 33-34, pi. 9, fig. 8-15. —Rose, 1933, p. 85, fig. 43. —Davis, 1949, p. 22-23, fig. 25-26. —Brodsky, 1950 (part), p. 123- 124, fig. 43; "female T" only. —Farran and Ver- voort, 1951a, p. 2-3, fig. 2b, g. — Tanaka, 1956, p. 391-392, fig. 15. —Grice, 1971, p. 275-280, fig. 1, 3G-H. Spinocalanus latifrons Sars, 1907, p. 5. Spinocalanus heterocaudatus Rose, 1937, p. 151-157, fig. 1-3. —not Rose, 1942, p. 315-318, 15 figs. Spinocalanus pacificus Mori, 1942, p. 568, pi. 14, fig. 1-6. Spinocalanus spinipes Brodsky, 1950 (part), p. 126- 127, fig. 45; male only. Type Specimen: Unknown. TYPE LOCALITY: Northeast Atlantic, 500-1,800 m. Material Studied: 2 females (2.77, 2.90 mm), sample 2 (Table 3); 1 female (2.1 mm), sample 27; 2 females (1.98, 2.05 mm), sample 30; 36 females (1.95-2.20 mm, mean = 2.08 mm) and 1 male (1.80 mm), sample 31; 3 females (2.2-2.4 mm), sample 33; 1 female (2.32 mm), sample 34; 6 females (1.87-2.42 mm, mean = 2.10 mm), 3 female stage V (1.85, 1.92, 1.95 mm), and 4 male stage V (1.88, 2.00, 2.00, 2.02 mm), sample 41. Description PI Ri with 3 inner setae; P2 Ri2 with 1 outer seta. Female.— Length 1.87-3.1 mm (Figs. 35-38). Prosome in dorsal view relatively slender, long ovoid. Head rounded in dorsal view, in lateral view with gen- tle, uneven forward slope. Th4 and Th5 separate; Th5 lateral corners prolonged, variable, reaching about midgenital segment; projections sometimes more or less closely surrounding genital segment (Fig. 38). Thl-Th4 without lateral spinules. Prosome length 3Vi times urosome. Area of attachment proximal to genital segment without dorsal spinules. Genital seg- ment about as long as wide, protruding ventrally one- half depth of rest of urosome; caudal rami length about l 2 /3 times width, left ramus longer and wider than right. Caudal setae also asymmetrical: left ter- minal seta second from inner, very thick, length ex- ceeding 3 mm. Al reaching midcaudal rami; terminal segments (Fig. 150) of moderate length and width. Apparent deviations: I-2s; XV-ls; remainder as Table 4. A2 Rel without setae. Mn palp Without setules on inner distal B2; Ril 26 38 41 Spinocalanus magnus Figure 35. — Female, habitus, lateral view; sample 3 1 (Tahlc :i | ; scale A. Figure 36. — Female, habitus, dorsal view; sample 31. scale A. Figure 37. — Female, habitus, lateral view; sample 11; scale A. Figure 38. — Female, Th5 and urosome, dorsal view; sample 11: scale A. Figure 39. — Male, habitus, dorsal view; sample 31; scale A. Figure 40.— Male, habitus, lateral view; sample 31; Kale \ Figure 41. — Male, P5; sample 31; scale I). Figure 42. — Male stage V, P5; sample 11; scale D. 27 with 2 setae and surface setules. Blade damaged in all preparations. Mxl gnathobase apparently without anterior distal seta, although very flat spiniform process suggested by some preparations; 4 proximal posterior setae. Mx2 with moderately strong nude hump on prox- imal outer edge; outer seta apparently absent; posterior surface of lobe-5 base without spines. Mxp Bl without spine-comb; transverse row of setules about midlength on anterior surface; distal in- ner anterior surface with transverse row of stiff hairs; few other spinules and setules over surface. B2 without spine-comb; longitudinal row of stiff hairs on inner edge. Two outer setae on Ri4-5 not fragile, as large as other large terminal setae, fringed with long hairs on inner edge, short hairs on outer edge; no spinules on inner edge of Ri5. PI Bl inner edge and surface with few hairs. B2 in- ner surface and margin with few setules and hairs; outer distal anterior edge with transverse row of small spinules. Rel-2 anterior surface with outer distal row of spinules; Re3 posterior surface without spines. Rel outer spine barely reaching Re3, although somewhat variable; Re2 outer spine reaching slightly beyond base of terminal spine. Ri inner lobe with row of spinules on anterior surface. P2-P3 Bl surface nude, inner margin with hairs. P4 Bl surface with proximal row of spinules on posterior surface, in addition to transverse row of setules near base of seta; inner margin without hairs; B2 nude. P2 Rel outer spine small, straight, not sharply curved. Re2 posterior surface on P3-P4 with 1 oblique row of long spines. Re3 posterior surface on P2-P3 with 1 distal row of short spines; posterior surface on P4 without spines, anterior surface with distal row of small spines. Ri2 posterior surface on P2 with proximal row of 3-7 small spines, distal row of 6-9 long spines. P3-P4 Ri3 posterior surface with proximal row of denticles and distal row of long spines. Re terminal seta with following number of primary teeth: P2 (24-29); P3 (20-25); P4 (24-25). Male.— Length 1.80-2.43 mm (Figs. 39, 40). Prosome in dorsal view long ovoid; anterior irregular, nearly rectangular; posterior somewhat rounded; in lateral view with gentle forward slope. Ce and Thl separate; Ce only slightly expanded dorsolaterally. Prosome length 2 2 /3 times urosome. Anal segment not reduced, two-thirds length of caudal ramus. Caudal rami length 1% times width, symmetrical. Caudal setae unknown. Al reaching end of caudal rami; segment 1 without cluster of spines or setules; segments 8 and 9 (fused) relatively long; segment 25 not reduced. IV-2s, le; Vm-2s, le; XV-ls, le; XIX-ls, le; right XX-0; left XX-le; remainder as Table 4. P4 Bl with inner seta reaching mid-B2. P5 (Fig. 41) biramus, left-handed; reaching nearly to distal edge of urosome segment 3. Left Bl reaching one-third length of right B2; left B2 reaching middle of right Rel. Right leg somewhat longer than left; right Re, including terminal blade, longest; left Ri longest. Order of length, longest to shortest, of Re segments: left 1, 2, 3; right 1 = 2, 3. Left Ri reaching left Re tip; right Ri reaching right mid-Re3. Each Rel with short flat seta on distal outer edge. Right Rel-2 nearly fused; line of fusion midlength on each Ri. Inner edge of left Re2 with long hairs. Only 1 bladelike terminal seta seen on each Re. Male stage V.— Length 1.88-2.02 mm. PI Ri lobe anterior surface with 2 close rows of spinules, upper row ca. 15 short spinules, lower row ca. 7 long spinules. P5 (Fig. 42) biramus, symmetrical. Re 1- segmented; small flat seta just proximal to trace of segmentation about one-third length; 1 flat terminal seta. Ri 1 -segmented with midlength trace of segmen- tation; terminal flat seta nearly reaching Re terminal seta. Remarks Wolfenden's (1904) original description of S. magnus was very brief and only distinguished it from known species. Farran (1905) provided the first detailed descrip- tion; he stated that the setation of the swimming legs agreed with S. abyssalis, which it does not. His il- lustration of PI, however, shows 5 setae on Ri; this was overlooked by Rose (1937) and Grice (1971). Farran mentioned the asymmetry of the caudal rami and setae; the large ramus was incorrectly stated as the right, but correctly shown in his figure. Wolfenden ( 1906) provided descriptive information on an Antarctic specimen, which he said agreed close- ly with northeast Atlantic specimens and those from the Strait of Gibraltar. This description was essential- ly reprinted in- 1911. Sars (1924, 1925) unfortunately described and il- lustrated S. magnus PI Ri with only 4 setae. Three of the four samples in which he found this species were from the Mediterranean Sea; this seems to have been overlooked by Sars himself in his summary of dis- tribution, and also by Rose (1933, 1937) and Scotto di Carlo (1968), but not by Massuti (1939). Sars reported that S. magnus is found throughout the Bay of Biscay; this is not supported by his data or earlier records. Perhaps he misread "west" for "east" longitude of his eastern stations, placing the three stations in the vicinity of the Bay of Biscay, albeit two on land. Van Breemen's (1908) description is from Farran. Rose's (1933) and Farran and Vervoort's (1951a) descriptions are from Sars. Brodsky (1950) included two species in his descrip- tion of S. magnus (see S. antarcticus) . Only his remarks on and illustration of the North Pacific form apply to S. magnus. Brodsky, however, was the first since Wolfenden to point out the existence of two 28 closely related forms differing in body type; Brodsky suggested that these might be subspecies. Tanaka (1956) was the first to recognize, and provide descriptive information on, the male of S. magnus. Sars (1907) reported S. magnus from the Atlantic and also briefly described a new species, S. latifrons, which he later (1924, 1925) equated with S. magnus. Rose (1937) described in detail a new species, S. heterocaudatus, from the Bay of Algiers. He seems to have had but one specimen, although the number was not stated. He agreed that it was close to S. magnus, with essentially only four differences: S. heterocaudatus was characterized by (1) 5 setae on Pi Ri, (2) left caudal ramus with an extraordinarily long and thick second from inner terminal seta, (3) symmetrical (sic) caudal rami, and (4) Th5 lateral corners prolonged to nearly the end of genital seg- ment. There is no doubt that 5. magnus has 5 setae on PI Ri, and it also has an enlarged left caudal seta; both of these characteristics were described by Farran (1905) and agree with the present material. The caudal rami might be variable in symmetry (see S. similis). The Th5 prolongation is definitely variable, due in part to the contraction of the urosome, although not to the degree shown by Rose; the viewing angle may make the prolongation appear more or less than it is, although Rose's lateral view also indicates a great prolongation. This specimen may have been ab- normal, as suggested by Grice (1971). Nearly 1 yr after collecting S. heterocaudatus, Rose found S. magnus at the same locality, but S. heterocaudatus was never again reported. Rose (1942) described a specimen as the male of S. heterocaudatus, also from the Bay of Algiers; that this specimen is definitely not in Spino- calanidae has been mentioned by Vervoort (1946). Mori (1942) described a female stage V S. magnus as the adult of a new species, S. pacificus. He was aware of the close relationship of the two species, but incorrectly considered S. magnus to have 4 setae on PI Ri, not 5 as did his specimen. Mori's figures essen- tially agree with female stage V examined in this study: PI Ri lobe anterior spinules longer than shown by Mori, extending beyond lobe; P2 as shown by Mori with 1 row of spines on Re2-3 and Ri2; P3 with 1 row of spines on Re2-3 and Ri2-3; P4 with 1 row of spines on Re2 and Ri2-3, and no posterior surface spines on Re3. Re3 on P2-P4 proportionally shorter and wider than on adult females. Mori's specimen was 1.9 mm; it was collected near Palao, "accurate positions unknown." Brodsky (1950) described a female (see S. horridus) and a male as a new species, S. spinipes. The descrip- tion of the male is consistent with that of S. magnus, except for a shorter, probably left, P5 Ri, which may have been broken. There are only four species of Spinocalanus known, including S. magnus, with male anal segment not reduced; Brodsky 's figure clearly in- dicates a short, nonreduced anal segment. The body form and size is unlike 5. ualidus male; S. antarcticus and 5. angusticeps males have uniramus P5. Distribution Except for Arctic records and some records in tran- sition areas, between the Atlantic and the Arctic, and around the Antarctic, records of S. magnus are accepted. Spinocalanus magnus is very widespread, horizontally and vertically: Pacific Ocean North: Brodsky (1950, 1957), 400-4,000 m. — Minoda (1971), 1,000-2,000 m. Northeast: Davis (1949), 1,100-2,300 m. — Flem- inger (1967), 0-140 m. East: Present study, 0-300 m. Central: Wilson (1942), 50-100 m. —Present study, 0-1,000 m. Northwest: Brodsky (1950; 1952a, b; 1957), 0- 4,000 m. —Tanaka (1953, 1956), 0-1,000 m. — Furuhashi (1961, 1966), 298-3,010 m. West: Mori (1942). Indo-Pacific: Vervoort (1946), 345-2,500 m. Indian Ocean North: Sewell (1929), 0-360 m. West: Grice and Hulsemann (1967), 1,000- 4,000 m. Southwest: De Decker and Mombeck (1965), 1,000-1,500 m. Atlantic Ocean North: Lysholm and Nordgaard (1921, 1945), 400-1,250 m. Northeast: Wolfenden (1904, 1906), 540-1,800 m. —Farran (1905, 1908, 1920, 1926), 0-3,600 m. —Sars (1912; 1924, 1925), 0-3,000 m. —With (1915), 0-1,000 m. — Lvsholm and Nordgaard (1945), 300-1,000 m. East: Lysholm and Nordgaard (1945), 1,200- 3,400 m. —Grice and Hulsemann (1965), 180- 5,000 m. —Roe (1972b), 550-960 m. Mediterranean Sea: Wolfenden (1906). 0-650 m. —Sars (1907; 1924, 1925), 0-2,300 m. —Rose (1937), 0-400 m. — Scotto di Carlo (1968), be- low 300 m. — Hure and Scotto di Carlo (1968), below 300 m. —Grice (1971), 850-1,400 m. Southeast: Unteruberbacher (1964), 0-300 m. West: Grice (1963), 600-1,200 m. —Wheeler (1970), 2,000-4,000 m. Caribbean Sea: Park (1970), 500-1.900 m. Antarctic Indian Sector: Wolfenden (1906, 1911). 0- 3,400 m. The records of Pearson (1906) and Massuti (1939) are previous records of Farran and Sars. Probably S. magnus enters the Arctic from the Atlantic, but its distribution there should be in- vestigated further, since the similar S. antarcticus is present over a large part of the Arctic. Reports of S magnus from the Greenland Sea (Damas and Koefoed, 1907; repeated by Jespersen. 1939b). 100- 1,350 m; Davis Strait (Stormer. 1929). 100-300 m; 29 West Greenland (Jespersen, 1934), 500-1,500 m; and the Norwegian Sea (Ostvedt, 1955), 600-2,000 m, could be either or both species. The investigation of 5. magnus in the Antarctic has likewise been hampered by the co-occurrence of the two closely related species. The material of Hardy and Gunther (1935) and Vervoort (1951, 1957, 1965) should be reexamined. Wilson (1942) combined S. magnus from two samples (sample 41, Table 3); the composite did con- tain S. magnus and both stations are considered valid records. The other three samples reported by Wilson to contain S. magnus were not located but were collected at intermediate positions. Wilson (1950) reported 5. magnus from the China Sea, but the sam- ple so labeled did not contain any Spinocalanidae and the record is considered invalid. Some of Farran's specimens of S. magnus are in the collection of the National Museum of Ireland, Dublin (O'Riordan, 1969). The holotype of S. latifrons Sars is on slides F5041-F5043, OSLO. 5. Spinocalanus antarcticus Wolfenden, 1906 (Figures 43-68, 151, 152) Spinocalanus antarcticus Wolfenden, 1906, p. 43, pi. 14, fig. 6-9. —Wolfenden, 1911, p. 217-218, fig. 9. Spinocalanus magnus. — Brodsky, 1950 (part), p. 123-124, fig. 43; not "female T". —Bradford, 1971, p. 18, fig. 28-29. — Vidal, 1971, p. 18, 23, fig. 34-35, 38, 40-41. Not S. magnus Wolfenden, 1904. Spinocalanus? sp. Johnson, 1963a, p. 91-94, fig. 2. Indian Ocean Sector, Type Specimen: Unknown. Type Locality: Antarctic 1,200 m. Material Studied: 7 females (2.38-2.56 mm, mean = 2.46 mm), sample 7 (Table 3); 7 females (2.55-2.62 mm, mean = 2.58 mm), sample 9; 9 females (2.52-2.83 mm, mean = 2.66 mm), 1 male (1.80 mm), and 2 male Spinocalanus antarcticus, female Figure 43. — Genital segment, ventral view; scale C. Figure 44.— A2; scale C. 43 stage V (1.75, 1.90 mm), sample A2 (Table 1); 9 females (2.5-2.75 mm, mean = 2.64 mm), 5 female stage V (1.9-2.1 mm, mean = 2.04 mm), and 3 male stage V (1.7, 1.8, 1.8 mm), sample A4; 2 females (2.60, 2.68 mm) and 3 males (1.83, 1.84, 1.90 mm), sample A5; 6 females (2.50-2.80 mm, mean = 2.59 mm), 3 female stage V (2.0, 2.15, 2.25 mm), and 1 male (1.70 mm), sample A6; 2 males (1.98, 2.05 mm), sample A23; 2 males (2.00, 2.32 mm), sample A27; 6 females (2.60-2.76 mm, mean = 2.67 mm), sample A33; 2 females (2.60, 2.65 mm) and 5 males (1.92-2.09 mm, mean = 1.99 mm), sample A34; 1 male (2.00 mm), sample A36; 7 females (2.76-3.00 mm, mean = 2.84 mm) and 1 male (2.08 mm), sample A48; 2 females (2.75, 2.82 mm), sample A50; 1 male (1.92 mm), sam- ple A51. Description PI Ri with 3 inner setae; P2 Ri2 with 1 outer seta. Female.— Length 2.25-3.00 mm (Figs. 45, 46). Prosome in dorsal view robust ovoid, in lateral view with abrupt, even forward slope. Th4 and Th5 separate; Th5 lateral corners prolonged, variable, reaching about midgenital segment. Thl-Th4 without lateral spinules. Prosome length 4Vi times urosome. Area of attachment proximal to genital segment with dorsal spinules. Genital segment slightly wider than long, protruding ventrally one-third depth of rest of urosome; genital segment relatively opaque, masking internal structure; genital field (Fig. 43) apparently similar in all Spinocalanus species. Supra-anal plate with row of spinules. Caudal rami about as long as wide, left ramus longer and wider than right. Caudal setae also asymmetrical: left terminal seta second 30 Spinocalanim antarcticus, female Figure 45. — Habitus, dorsal view; scale A. Figure 46. — Habitus, lateral view; scale A. Figure 47. — Al; scale C. Figure 48. — Mxl; scale C". Figure 49. — Mxp; scale C. 31 51 Spinocalanus antarcticus, female Figure 50. — Mn; scale C. Figure 51. — Mn blade; scale F. Figure 52.— Mx2; scale C. Figure 53.— PI; scale C. Figure 54.— P2; scale C. Figure 55.— P3; scale C. Figure 56.— P4; scale C. 32 from inner, very thick, length exceeding 6 mm. Small outer (sixth) seta on some specimens. Al (Fig. 47) reaching heyond caudal rami by about 3 segments; terminal segments (Figs. 151, 152) of moderate length and width. Apparent deviation: XV- ls; remainder as Table 4. A2 (Fig. 44) Rel without setae. Mn (Fig. 50) palp without setules on inner distal B2; Ril with 2 setae and surface setules; Ri2 ap- parently with 10, rather than 11, setae. Blade (Fig. 51) with cluster of setules on midanterior surface; cutting edge with 5 large teeth, outermost 1 -pointed, others more or less 2-pointed; 1 near-dorsal 4-pointed pro- cess; and dorsal seta. Mxl (Fig. 48) gnathobase with 4 proximal posterior setae and small distal anterior seta. Mx2 (Fig. 52) with moderately strong nude hump on proximal outer edge; outer seta apparently absent; posterior surface of lobe-5 base without spines. Mxp (Fig. 49) Bl without spine-comb; transverse row of setules about midlength on anterior surface; longitudinal row of spinules midlength on inner anterior surface; distal inner anterior surface with row of stiff hairs. B2 without spine-comb; short longitudinal row of stiff hairs on inner edge. Ri4-5 out- er setae not fragile, as large as other large terminal setae, fringed with long hairs on inner edge, short hairs on outer edge; no spinules on inner edge of Ri5. Pi (Fig. 53) Bl inner edge and surface with few setules. B2 inner surface with setules, margin with hairs; outer distal anterior edge with transverse row of small spinules. Rel-2 anterior surface with outer dis- tal row of spinules; Re3 posterior surface with 6-8 short spines. Rel outer spine reaching Re3, although length variable; Re2 outer spine reaching beyond base of terminal spine. Ri inner lobe with row of spinules on anterior surface. P2 (Fig. 54)-P3 (Fig. 55) Bl surface nude, inner margin with hairs. P4 (Fig. 56) Bl surface with prox- imal row of spinules on posterior surface, in addition to transverse row of setules near base of seta; inner margin without hairs; B2 nude. P2 Rel outer spine small, straight, not sharply curved. Re2 posterior surface on P3-P4 with 1 oblique row of long spines. Re3 posterior surface on P2-P3 with 1 distal row of spines; posterior surface of P4 without spines, anterior surface with distal row of small spines. P2 Ri2 posterior surface with proximal row of few denticles, distal row of long spines. P3-P4 Ri3 posterior surface with proximal row of denticles (sometimes minute) and distal row of long spines. Re terminal seta with following number of primarv teeth: P2 (36-41); P3 (32-40); P4 (31-44). Male.— Length 1.66-2.32 mm (Figs. 57, 58). Prosome in dorsal view robust ovoid; anterior and posterior irregular, somewhat rectangular; in lateral view with abrupt, even forward slope. Ce and Thl separate. Ce not expanded dorsolaterals. Prosome length 2' 2 times urosome. Anal segment not reduced, two-thirds length of caudal ramus. Caudal rami length 1 ' 2 times width, left ramus slightly larger than right. Inner caudal seta reduced; left terminal seta se- cond from inner somewhat longer than right; small outer (sixth) seta present. Al (Fig. 59) reaching beyond caudal rami hy 2-3 segments; segment 1 without cluster of spines or setules; segments 8 and 9 (fused) relatively long; seg- ment 25 not reduced. IV-2s, le; VIII-2s, le: XV- ls, le; XIX-ls, le; right XX-ls; left XX-le; remainder as Table 4. Mn blade (Fig. 60) reduced; 5 more or less blunt teeth, 3-pointed near-dorsal process, 1 short, nude dorsal seta. Mxl gnathobase (Fig. 61) reduced; 6 outer blunt processes, 3 proximal moderately sharp, nude setae. Mxp (Fig. 62) similar to that on female, perhaps with loss or marked reduction of some Bl-2 setae; Ri4- 5 setae nude, outer 2 and terminal setae markedly reduced. PI (Fig. 63) Bl inner surface with few short hairs; B2 inner margin with long hairs. Ri surface nude. Rel outer spine reaching beyond midlength of Re2, nearly to base of Re2 outer spine; Re3 surface without spines. P2 (Fig. 64)-P3 (Fig. 65) as on female, except P2 Ri2 without proximal spinules. P4 (Fig. 66) Bl inner seta shorter than on female, reaching beyond Ril. Ri3 proximal posterior surface with or without few minute or small spines, distal sur- face with 1 row of long spines. Re3 without distal anterior row of spines. P4 otherwise as on female. P2-P4 Re terminal seta with finer and more numerous (ca. 50) teeth than on female. P5 (Fig. 67) left-handed; uniramus, very asymmetrical. Right leg reaching midurosome seg- ment 3; left leg greatly exceeding length of urosome. Right leg not reaching end of left Bl. Order of length, longest to shortest, of (Re) segments: left and right 1. 2, 3. Left Re2 about one-half length of Rel. Right leg segments nude, except short, flat terminal seta. Left B2 with inner distal cluster of hairs; small distal cluster of hairs on Re2; Re3 short, with central cluster of hairs, terminating in digit iform process. Male stage V.— Length 1.7-1.90 mm. Pi Ri lobe anterior surface with 2 close rows of spinules. upper row ca. 15 short spinules, lower row ca. 7 long spinules. P5 (Fig. 68) left-handed; uniramus. asymmetrical; right leg reaching left Rel. B2 with inner distal spiniform process. He 2-segmented. with perhaps trace of small terminal segment; Rel with outer distal spiniform process; Re with short. Hat terminal process. 33 34 Remarks Wolfenden (1906), who also first described the closely related S. magnus, differentiated between that species and S. antarcticus because of the different body types. This is easily seen when the two species are side-by-side, as they were in Wolfenden's Antarc- tic collections. However, this difference in form is not so easily defined. Wolfenden's figures indicate only a slight prolongation of Th5; actually it is as prolonged as on S. magnus. His lateral figure (1911) indicates some hesitancy in this regard since it is clearly altered. The unusual male, described by Wolfenden (1906), probably led several workers (e.g., Vervoort, 1957) to discount, at least in part, the validity of S. antarcticus. Wolfenden's description of the male agrees with that given above, except in one feature: Wolfenden stated, and illustrated, that right P5 was longest; this could be an error, since left P5, when the male is viewed dorsally, actually curves out to the right of the specimen, while the shorter right leg is obscured. No males from the Antarctic were found for comparison during the present study. Brodsky ( 1950) described two body types of S. magnus, which he suggested might be subspecies. Only his Arctic specimens were S. antarcticus, while those from the North Pacific were 5. magnus. His figure of P3 appears to be a composite of P3 and P4. The description and illustrations by Bradford (1971) are consistent with those of S. antarcticus. Bradford's specimens (sample 9, Table 3) were found also to agree in all respects with the description of S. antarcticus. Distribution There are several records of S. magnus from high latitudes that may be of S. antarcticus (see also S. magnus): especially the Arctic records of Grice (1962) in the Eurasian Basin (110 m); Grainger (1965) in the Canadian Basin (0-500 m); and Dunbar and Harding (1968) in the Canadian Basin (0-2,000 m). Several records of S. magnus are interpreted or con- firmed as S. antarcticus. Some of Farran's (1929) specimens from the Antarctic (sample 7, Table 3) were found to be S. antarcticus Brod>ky and Nikitin's (1955) records of .S. magnus from the Arctic Canadian Basin are accepted as S. antarcticus John- Bon (1963a) and Minoda (1967) both indirectly in- dicated the presence of male S, antarctic us. bo that their records of S. magnus are also interpreted a- S antarcticus. The distribution of S. antarcticus is summarized below: Arctic Ocean Central: Brodsky (1950), 0-400 m or more. Eurasian Basin: Minoda (1967). 0-850 m. Canadian Basin: Brodsky and Nikitin (1955), 0-below 1,000 m. —Johnson (1963a). 0- 2,000 m. — Vidal (1971). —Present study. 140-2,500 m. Antarctic Pacific Sector: Farran (1929), 0-1,750 m. - Bradford (1971), 0-1,000 m. Indian Sector: Wolfenden (1906, 1911). 1,200 m. Although there have been no records of S. antarc- ticus outside of the Arctic and Antarctic, the possibili- ty of a continuous distribution should not be over- looked. Perhaps S. antarcticus inhabits greater depths in midlatitudes, or perhaps it has not been recognized in midlatitude collections. A specimen of Spinocalanus sp. Johnson was reported by Wheeler (1970) from the northwest Atlantic, but this is probably a male of S. angusticeps. (6. Spinocalanus major Esterly, 1906) See Spinocalanus brevicaudatus Brodsky, 1950. (7. Spinocalanus latifrons Sars, 1907) See Spinocalanus magnus Wolfenden. 1904. Spinocalanus antarcticus, male Figure 57. — Habitus, dorsal view; scale A. Figure 58.— Habitus, lateral view; scale A. Figure 59.— Al; scale C. Figure 60. — Mn blade; scale F. Figure 61.— Mxl gnathobase; scale F. Figure 62.— Mxp; scale C. Figure 63.— PI; scale C. Figure 64.— P2; scale C. Figure 65.— P3; scale C. Figure 66.— P4; scale C. Figure 67.— P5; scale C. Figure 68.— Stage V, P5; scale I). 8. Spinocalanus hirtus Sars, 1907 Spinocalanus hirtus Sars. 1907, p. 5-6. — Sars. 1925. p. 32-33. Spinocalanus hirtipes. — Sars. 1924. lapsus calami, pi. 9, fig. 1-7. TYPE SPECIMEN: Holotvpe. female, on Slide F5039, OSLO. TYPE LOCALITY: East Atlantic. 0-5,000 m. Description Pi Ri with 2 inner setae: P2 Ri2 with 1 a. 35 Female. — Length 2.12-3.58 mm. Prosome in dorsal view long ovoid, head rounded; in lateral view with gentle, then abrupt, forward slope. Th4 and Th5 separated by fine suture; Th5 lateral corners slightly prolonged, margins with stiff hairs. Prosome length 3V3 times urosome. Mxp details unknown. P1-P4 Bl densely covered with short hairs. P2 Re2 posterior surface possibly with two rows of spines. P2- P3 Re3 posterior surface with several small spines along length. Male. — Unknown. Remarks Sars (1907) briefly described this species from a single female; more information on the same specimen, with illustrations, was added later (1924, 1925). Sars' figure of PI is lacking an inner seta on Re3. Only three other specimens have been reported: Vervoort (1946), one female from the Indo-Pacific, 653-1,000 m; Grice and Hulsemann (1965), two females from the eastern North Atlantic, near the type locality, 3,000-4,750 m. 9. Spinocalanu8 spinosus Farran, 1908 (Figure 84) Spinocalanus spinosus Farran, 1908, p. 27, pi. 1, fig. 10. —Rose, 1933, p. 86, fig. 46. —Farran and Ver- voort, 1951a, p. 2-3, fig. 4b, d-e. —Park, 1970, p. 485-487, fig. 37-42. TYPE SPECIMENS: Syntypes, females, BM 1908.7.6.7- 10. Type LOCALITY: Northeast Atlantic, 600-1,800 m. Material Studied: 3 females (1.67, 1.85, 1.87 mm), sample 5 (Table 3); 2 females (1.90, 1.90 mm), sample 31. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 1.67-2.0 mm. Prosome in dorsal view somewhat robust ovoid, head slightly truncate; in lateral view with abrupt forward slope. Th4 and Th5 separate; Th5 lateral corners rounded, not prolonged. Thl-Th4 with lateral spinules, number decreasing posteriorly, approximately even right and left. Prosome length 2% -3 times urosome. Genital segment about as long as wide, protruding ventrally about one-third depth of rest of urosome; long hairs on ventral surface of urosome segments 1-3. Caudal rami length IV2 times width, symmetrical. Al broken in specimens examined. A2 Rel with 2 setae; smaller seta easily detached. Mx2 with moderate, setule-covered hump on prox- imal outer edge; outer seta present; posterior surface of lobe-5 base without spines. Mxp Bl without spine-comb; strong transverse spine-comb on B2. Ri4-5 outer setae nude, bladelike, tapering toward each end; spinules on inner edge of Ri5. PI (Fig. 84) Bl outer surface with few thin spinules; inner margin with hairs. B2 anterior surface with few thin spinules and setules. Outer distal anterior sur- face of Rel-2 with row of spinules. Posterior surface of Re3 with ca. 4 spines. Ri inner anterior surface with row of thin spinules about midlength. Male. — Unknown . Remarks Farran (1908) briefly characterized Spinocalanus spinosus by describing the lateral spinules on Thl- Th4. This differentiated S. spinosus from the species known up to that time. Since then, several closely related species have been described (S. horridus, S. usitatus, S. hoplites, and S. oligospinosus) which seem to be distinguishable essentially by the presence or absence of spines on the base of Mx2 lobe-5; the presence and size, or absence of a transverse spine- comb on Mxp Bl; the distribution of lateral spinules; body size, and to some extent body shape. Specimens of all of these "species" have been examined in the present study and these characteristics are described for each. They appear to be consistent and no transi- tion forms have been observed. The only described male is assigned to S. horridus; if these named-species with spinulose prosomes are valid, there must be four undescribed males. The co-occurrence of these spe- cies hampers the interpretation of records of S. spino- sus. Rose's (1933) description is from Farran (1908). Farran and Vervoort (1951a) illustrated Mx2 without spines on the base of lobe-5, consistent with Farran's specimens from sample 5 (Table 3). Farran and Ver- voort assigned some records of S. horridus to S. spinosus, but the two species are considered distinct in the present study (see S. horridus). Some records of very large "S. spinosus" (e.g., Farran, 1929; Tanaka, 1956; Grice and Hulsemann, 1965, part) are interpreted as records of S. horridus. Vervoort (1946) reported two female S. spinosus; he stated that the Mxp of both was as described by Wolfenden (1911) for S. horridus, i.e., with Bl transverse spine-comb. Therefore, the larger (2.40 mm) specimen is considered in the present study to be S. horridus, while the smaller (1.80 mm) is considered S. usitatus. Park (1970) did not describe S. spinosus Mx2, but the Mxp of his specimens was described as without Bl spine-comb, consistent with the above description. Park illustrated P2-P4; PI is illustrated in the present study (Fig. 84). 36 Distribution Accepted records of S. spinosus are: Pacific Ocean Central: Present study, 0-1,000 m. Atlantic Ocean Northeast: Farran (1908, 1920, 1926), 180-1,800 m. Caribbean Sea: Park (1970), 505-1,000 m. Probable records, at least in part, of S. spinosus are: Grice (1963), Grice and Hulsemann (1965), Wheeler (1970), and Roe ( 1972b) in the North Atlantic; and De Decker and Mombeck (1965) and Grice and Hulsemann (1967) in the Indian Ocean. Some of Farran's specimens of S. spinosus are in the collection of the National Museum of Ireland, Dublin (O'Riordan, 1969). 10. Spinocalanu8 horridus Wolfenden, 1911 (Figures 69-83, 153) Spinocalanus horridus Wolfenden, 1911, p. 216, fig. 7; pi. 25, fig. 1-2. —Park, 1970, p. 491, fig. 66-70. Spinocalanus abyssalis. — Tanaka, 1937 (part), p. 253-254, fig. 4a-c; female only. — Minoda, 1971 (part), p. 11, fig. 30(3), 32(1), 33(2); 1.95-2.2 mm only. Not S. abyssalis Giesbrecht, 1888. Spinocalanus stellatus Brodsky, 1950, p. 125-126, fig 44. —von Vaupel-Klein, 1970, p. 8-10, fig. lb-f. Spinocalanus spinipes Brodsky, 1950 (part), p. 126- 127, fig. 45; female only. Spinocalanus dorsispinosus Brodsky, 1950, p. 129, fig. 48. Spinocalanus sp.? Brodsky, 1950 (part), p. 130-131, fig. 50; female only. Spinocalanus longispinus Brodsky, 1950, p. 131, fig. 51. — Vidal, 1971, p. 18, 23-24, fig. 42-45, 58-59. Spinocalanus spinosus. — Tanaka, 1956, p. 390-391, fig. 14. Not S. spinosus Farran, 1908. Type Specimen: Unknown. TYPE LOCALITY: South Atlantic, 0-1,200 m. Material Studied: 3 females (2.23-2.30 mm), sam- ple 38 (Table 3); 2 females (2.10, 2.12 mm), sample 58; 1 female (2.12 mm), sample 59; 1 female (2.28 mm) and 1 male (2.10 mm), sample A2 (Table 1); 1 male (2.1 mm) and 1 male stage V (1.95 mm), sample A4; 2 males (2.18, 2.25 mm), sample A5; 1 male stage V (1.97 mm), sample A6; 1 male (2.28 mm), sample A21; 2 females (2.48, 2.52 mm) and 1 male (2.32 mm), sample A23; 4 females (2.28-2.48 mm, mean = 2.36 mm), sample A24; 1 male (2.24 mm), sample A33; 1 female (2.36 mm), sample A40; 1 female (2.36 mm), sample A41; 1 female (2.32 mm), sample A42; 1 female (2.36 mm), sample A45; 2 females (2/SZ. 2.58 mm) and 1 male (2.24 mm), sample A48. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female.— Length 1.95-3.0 mm (Figs. 69. 70). Prosome in dorsal view slightly robust, elongate ovoid; head rounded with slight anterior protuberance. Prosome in lateral view with irregular, gentle forward slope. Th4 and Th5 separate; Th5 lateral corners rounded, not prolonged. Thl-Th4 with lateral spinules, number decreasing posteriorly; right side sometimes completely devoid of spinules (Arctic specimens), or spinule distribution may be ap- proximately even right and left (Caribbean specimens). Prosome length 2 2 A times urosome. Genital segment somewhat longer than wide, protruding ventrally about one-half depth of rest of urosome; hairs on ventral surface of genital segment and probably urosome segment 2. Caudal rami length l'/2 times width, symmetrical. Al exceeding caudal rami by 1-2 segments; ter- minal segments (Fig. 153) slightly elongate. Appar- ent deviation: I-2s; remainder as Table 4. A2 Rel with 2 setae. Mn palp with central and distal cluster of long setules on anterior surface; Ril surface with setules. Mxl gnathobase (Fig. 71) with cluster of spines and 4 setae on proximal posterior surface; distal anterior surface with thin spinule; small cluster of setules near distal outer margin. Mx2 (Fig. 72) with small setule-covered hump on proximal outer edge; outer seta present; posterior sur- face of lobe-5 base without spines. Lobe-4 distal posterior seta stout, with row of close-set spines (Fig. 72 showing only alternate spines). Lobe-5 distal posterior seta spinulose, with slightly widened open- ing at tip of seta. Mxp (Fig. 73) Bl with strong transverse spine-comb on outer midmargin; irregular oblique rows of spines on central and distal anterior surface. One specimen with spine-comb only on left Bl. B2 with strong transverse spine-comb about one-third length on out- er margin; longitudinal row of spinules on inner margin. Ril and Ri5 with inner row of thin spinules. Ri4-5 outer setae nude, bladelike, tapering toward each end. PI (Fig. 74) Bl-2 surface with setules and hairs, in- ner margin with hairs. Rel-2 outer distal anterior sur- face with row of spinules. Posterior surface of Re:! with 5-6 spines. Ri inner anterior surface with row oi thin spinules about midlength. P2 (Fig. 75) Bl miter surface with 3-4 rows of setules; hairs and setules on inner margin. P3 I Fig Bl surface with 1 inner and 3-4 outer rows of spines; inner margin with hairs. P4 (Fig. 77) Bl inner surtace with several patches of long setules. outer surface with 37 Spinocalanua horridus, female Figure 69. — Habitus, dorsal view; scale A. Figure 70. — Habitus, lateral view; scale A. Figure 71. — Mxl gnathobase; scale F. Figure 72.— Mx2; scale D. Figure 73.— Mxp; scale C. Figure 74.— PI; scale C. Figure 75.- Figure 76.- Figure 77.- Figure 78.- Figure 79.- Figure 80.- Figurc 81. Figure 82. Figure 88.- Spinocalanus horridus Female, P2; scale C. Female, P3; scale C. -Female, P4; scale C. -Male, habitus, dorsal view- Male, habitus, lateral view; -Male, Mn blade; scale F. -Male, Mxp; scale C -Male, P5; scale C. -Male stage V, P5; scale I). scale A. scale A. 39 Spinocalanus spinosus, female Figure 84.— PI; sample 5 (Table 3); scale D. 3-4 rows of spines. B2 inner surface with cluster of thin setules. P2 Re3 with 2 distal rows of short spines on posterior surface. P3 Re2 with midlength oblique row of strong spines and distal row of denticles. Re3 with 3 evenly spaced rows of short spines. Ri3 with 2 rows of strong spines. P4 Re2 with midlength oblique row of strong spines and distal row of smaller spines. Re3 with 3 evenly spaced rows of spines, proximal spines longest, distal spines shortest. Re terminal seta with following number of primary teeth: P2 (30-36); P3 (28-33); P4 (29-32). Male.— Length 2.0-2.9 mm (Figs. 78, 79). Prosome anterior irregular in dorsal view, posterior somewhat rectangular; in lateral view with gentle forward slope. Ce and Thl partly fused. Prosome length 1% times urosome. Caudal rami about as long as wide. Al reaching end of prosome; segment 1 with con- spicuous cluster of setules on anterodorsal surface. IV-2s, 2e; Vm-2s, 2e; XV-2s, le; XIX-ls, le; right XX-0; left XX-le; remainder as Table 4. Mn B2 without distal setules; 2 short setae on inner margin. Ril surface setules longer than on female. Segmentation and setation of Re and Ri as on female. Blade (Fig. 80) reduced; ventral tooth in some specimens larger than figured. Mxl very fragile, gnathobase reduced to 1-2 small spines on knob. Mx2 not robust; short, thin outer seta. Some inner setae of reduced size, but probably number as on female. Mxp (Fig. 81) with reduced size of setae; relatively fragile. Ri4-5 outer setae short and flat; spinules on in- ner edge of Ri5 as on female. PI Bl surface without setules, but hairs longer than on female. B2 surface without setules, inner margin hairs shorter than on female. Re3 posterior surface spines relatively longer than on female. P2 Bl-2, Ri, and Re3 as on female. Re2 with circular patch of spines on posterior surface. Terminal spine with ca. 60 primary teeth. No specimens observed with P3 Re2-3; remaining segments as on female. No specimens observed with P4 Re2-3. Bl without setules on inner surface; inner seta reaching beyond Ril; remaining segments a on female. P5 (Fig. 82) biramus, left-handed; reaching nearly to end of urosome segment 2. Left Bl reaching one- third to one-half length of right B2; left B2 reaching middle of right Rel. Left and right legs about equal; right Re, including moderately long, flat terminal seta, longest; left and right Ri about equal. Order of length, longest to shortest, of Re segments: left 1 = 2, 3; right 2, 1, 3. Each Rel with short, flat seta on distal outer edge. Inner edge of left Re2-3 with hairs. Each Re3 with 2 unequal, bladelike terminal setae. Ri im- perfectly 2-segmented; left Ri reaching middle of left Re3; right Ri reaching middle of right Re3. Each Ri with long bladelike extension. Male stage V.— Length 1.95-1.97 mm. Left side of Th2-Th3 with spinules (Arctic specimens). P5 (Fig. 83) biramus, symmetrical. Re imperfectly 2-segmented; small flat seta on distal edge of Rel; Re2 with 2 small flat terminal setae. Ri 1-segmented, reaching nearly one-half length of Re, with small flat terminal seta. Remarks Wolfenden's (1911) original description of S. horridus included a figure of Mxp which clearly in- dicates a transverse spine-comb on Bl. Park (1970) likewise illustrated Mxp with spine-comb on Bl. These are the only two investigators who have described this species under this name. Tanaka (1937) described one female S. abyssalis, which, except for lateral prosomal spinules, which Tanaka did not mention, agrees with the description of S. horridus given above. The prosomal spinules are not particularly conspicuous unless viewed with proper lighting; perhaps also, other populations, besides the Arctic specimens of the present study, have spinules only on one side, and this characteristic might then not be noticed. The male described by Tanaka is considered as S. breuicaudatus. Brodsky (1950) recognized that Tanaka's specimens were not S. abyssalis Giesbrecht. Minoda (1971) reported large (1.95-2.2 mm) 5. abyssalis from the Bering Sea; his il- lustration of Mxp indicates a transverse spine-comb on Bl, and the scale to which it was drawn indicates 40 its origin in the larger specimens, probably 6'. horridus. Tanaka's (1956) description of S. spinosus is consis- tent with that of S. horridus given above. The large size and transverse spine-comb on Mxp Bl are not characteristic of S. spinosus. Brodsky's (1950) descriptions of two new species from the northwestern Pacific, S. stellatus (female and male) and S. spinipes (female only), are consis- tent with that of S. horridus except for lateral prosomal spinules on female, which Brodsky did not mention. Brodsky (1952a) said that S. spinipes was more transparent than S. stellatus. The male of S. spinipes is considered S. magnus. Brodsky (1950) described an additional new species, S. dorsispinosus, based on a male from the northwestern Pacific. Brodsky's description is consis- tent with that of male S. horridus except for the presence of a group of spines "on dorsal side of head." Perhaps these spines are present on males of other Spinocalanus species, as well as S. horridus, but are extremely fragile or only present for a short time. A similar feature was noticed in some male Mimocalanus sulcifrons, but not in any other species studied, including S. horridus. Brodsky's (1950) description of S. longispinus from the central Arctic was based, in part at least, on an abnormal specimen. His illustration of P4 indicates an inner seta on B2 (never normally present) and many spines on Ril (normally nude in Spinocalanus species). Aside from this, his remarks and other figures are consistent with the description of S. horridus female, except for lateral prosomal spinules, not mentioned by Brodsky. Specimens reported by Vidal (1971) to be S. longispinus were examined (sample 38, Table 3) and found to be equivalent in all respects to other Arctic specimens of S. horri- dus. Distribution Records of large S. spinosus by Farran (1929), cited by Vervoort (1965); Vervoort (1946); Johnson (1963a); and Grice and Hulsemann (1965; part, up to 2.7 mm) are considered as records of S. horridus. Probably S. spinosus reported by Furuhashi (1961, 1966) and Minoda (1967) are also S. horridus. Records of S. horridus and its synonyms are summarized below: Arctic Ocean Central: Brodsky (1950), 400-2,500 m. Eurasian Basin: Minoda (1967), 0-850 m. Canadian Basin: Brodsky and Nikitin (1955), 100-3,800 m. —Johnson (1963a), 0-2,000 m. —Dunbar and Harding (1968), 500-2,000 m. —Vidal (1971). —Present study, 500-2,500 m. Pacific Ocean North: Brodsky (1957), 100-200 m. —Minoda (1971), 95-2,000 m. Northeast: von Vaupel-Klein (1970). 0-1,200 m. Northwest: Tanaka (1937, 1953, 1956), 0-1,200 m. —Brodsky (1950; 1952a, b; 1957), 200- 4,000 m. —Furuhashi (1961, 1966), 425-3,010 m. —Minoda (1971), 160-1,800 m. Indo-Pacific: Vervoort (1946), 1,385-1,900 m. Atlantic Ocean Northeast: Grice and Hulsemann (1965), 500- 4,000 m. South: Wolfenden (1911), 0-1,200 m. Caribbean Sea: Park (1970), 509-1,900 m. Antarctic Pacific Sector: Farran (1929), 0-1,000 m. Atlantic Sector: Hardy and Gunther (1935), 1,000-2,000 m. (11. Spinocalanus giesbrechti Brady, 1918) Spinocalanus giesbrechti Brady, 1918, p. 17-18, pi. 8, fig. 21-25. TYPE SPECIMENS: The Hancock Museum, Newcastle- upon-Tyne, England. TYPE LOCALITY: Antarctic, Pacific Sector; surface? Remarks Vervoort (1951; p. 9-10, 70) studied the type specimens of 5. giesbrechti and, because of their poor condition, was unable to correctly identify the specimens. Vervoort stated that "the description is so vague and the figures are so bad that I am not even certain if the form is a Spinocalanus at all." Brady's male specimen could not have been an adult; Pi could not be of Spinocalanidae; the P4 appears abnormal; P5 is not from a Spinocalanus species. The female could not have been a Spinocalanidae. Since the type material is now useless, it may never be known what species S. giesbrechti represents, but Brady's description is definitely not of any Spinocalanus species. 12. Spinocalanus validus Sars, 1920 Spinocalanus validus Sars, 1920. p. 2. —Sars. 1924. 1925, p. 29-30, pi. 7, fig. 1-15. —Rose, 1933. p. 85. fig. 44. —Grice and Hulsemann. 1967, p. 21, fig. 28-31. Type Specimens: Syntypes, females (4.50 mm), Mu- see Oceanographique de Monaco (Belloc. 19601: on slides F5035-F5036, OSLO. TYPELOCALITY: Northeast Atlantic. 0-1,400 m. Material Studied: 1 male (4.4 mm), sample 44 (Table 3); 1 female (3.7 mm), sample 51. 41 Description Pi Ri with 3 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 3.7-4.7 mm. Prosome in dorsal view slightly robust ovoid. Head somewhat truncate in dorsal view, in lateral view with abrupt forward slope. Th4 and Th5 separate; Th5 lateral corners prolonged, reaching to or slightly beyond midgenital segment (Sars, 1924, fig. 15). Thl-Th4 without lateral spinules. Prosome length 4V6 times urosome. Al incomplete on specimen examined; Al scarcely exceeding length of prosome, according to Sars. Mxp without spine-comb on Bl-2. Ri4-5 outer setae missing on specimen studied. Ri5 inner edge without spinules. PI as shown by Sars, without posterior surface spines on Re3. P2-P4 Bl surface very setulose, as shown by Sars; specimen examined lacked Re2-3 and Ri. P2 Rel out- er spine small, straight, not sharply curved. Male. — Length 4.4-5.08 mm. Prosome anterior irregular in dorsal view, posterior somewhat rounded; in lateral view with fairly abrupt forward slope. Ce and Thl partly fused. Prosome length slightly over 2 times urosome. Anal segment not reduced, two-thirds length of caudal ramus. Al incomplete on specimen studied. Mxp with thick, not expanded, hirsute terminal setae; Ri4-5 with reduced outer setae. Ri5 inner edge without spinules. PI Rel outer spine very wide, straight; no posterior surface spines on Re3. P2-P3 Re and Ri missing on specimen examined. P4 Bl without inner seta; outer surface with proximal setules and distal spines. P4 Re missing. P4 Ri2 posterior surface with 1 proximal row and 1 distal cluster of spines; Ri3 with 2 clusters of spines. P5 biramus, about as described by Grice and Hulsemann (1967), except perhaps longest basal segments on left (left-handed) rather than right. Anterior/posterior slope of distal B2 edges suggesting this possibility; however, orientation should be reex- amined in situ. Each Re with small bladelike terminal seta in addition to long terminal seta shown by Grice and Hulsemann. Remarks Sars (1920) briefly described S. validus; his record of finding the species at "Stn. 1851" must have been an error, since S. validus is not on that station's list of species, and the station is not mentioned in 1925. Rose's (1933) description and records are repeated from Sars. Grice and Hulsemann (1967) briefly described the male of S. validus, but they also found females (un- published records); a male is in the USNM (113512) and was examined in the present study. Distribution There have been only three additional reports of S. validus, all of which are summarized below: Pacific Ocean Indo-Pacific: Vervoort (1946), 653-1,000 m. Indian Ocean West: Grice and Hulsemann (1967), 350-2,000 m. Atlantic Ocean Northeast: Sars (1920; 1924, 1925), 0-1,400 m. —Grice and Hulsemann (1965), 1,500-4,000 m. — Vives (1970), 300-700 m. 13. Spinocalanus angusticeps Sars, 1920 (Figure 85) Spinocalanus angusticeps Sars, 1920, p. 3. — Sars, 1924, 1925, p. 30-31, pi. 8, fig. 1-9. — Farran and Vervoort, 1951a, p. 2-3, fig. 3a-b, d. — Tanaka, 1956, p. 392-394, fig. 16. Spinocalanus longipes Tanaka, 1956, p. 394-395, fig. 17. —Grice and Hulsemann, 1967, p. 21, fig. 26- 27. Spinocalanus sp. — Wheeler, 1970, p. 9, fig. 23-26. Not Spinocalanus? sp. Johnson, 1963a. Type Specimens: Syntypes, females (2.10 mm), Mu- see Oceanographique de Monaco (Belloc, 1960); on slides F5037, F5047, OSLO. Type Locality: Northeast Atlantic, 0-2,000 m. Spinocalanus angusticeps, female Figure 85.— P3; sample 13; scale C. 42 Material Studied: 1 female (2.14 mm), sample 13 (Table 3); 1 female (on slide), sample 14; 1 male (1.60 mm), sample 50. Description PI Ri with 3 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 1.98-2.33 mm. Prosome in dorsal view long, slender ovoid; anterior tapering toward slightly truncate head. In lateral view with uneven, gentle forward slope. Th4 and Th5 separate; Thl-Th4 without lateral spinules; Th5 lateral corners protruding about one-third length of genital segment (somewhat more than shown by Sars, 1924, fig. 1). Prosome length slightly more than 4 times urosome. Genital segment as long as wide; protruding ventrally one-third depth of rest of urosome. Caudal ramus length slightly more than width. Caudal setae symmetrical; inner terminal seta omitted in Sars' figure. Al unknown; broken in all described specimens. [Sars (1925) stated that Al must be longer than the body, but his figure shows Al broken at segment 21 next to distal edge of Thl, so that a complete Al would probably not extend beyond Th4.] A2 Rel without setae, as shown by Sars. Mx2 without spines on base of lobe-5. Mxp about as shown by Sars, except 4 setae on Ri2 and only 4 on Ri5. Bl without spine-comb; transverse row of short spinules on distal inner edge. B2 without spine-comb; longitudinal row of stiff long hairs on in- ner edge. Pi as shown by Sars, except typical short, thin, out- er seta on B2 about two-thirds distal; row of spinules on distal outer anterior surface of Rel-2; no spines on Re3 surface. P2 as shown by Sars, except distal outer anterior spinules on Re2; Rel outer spine forming strongly bent hook; no surface spines on Re3, terminal seta with 30-32 primary teeth. P3 (Fig. 85) Ri only shown by Sars. Bl with several outer and inner surface spines; inner margin with hairs. Re2 posterior surface with irregular cluster of spines; Re3 posterior surface with 2 distal clusters of small spines; terminal seta with 31 primary teeth. P4 incomplete on specimens studied; Bl with setules, not spinules, in position shown by Sars. Male.— Length 1.59-1.76 mm. Prosome in dorsal view robust ovoid, anterior and posterior irregularly rounded; in lateral view with abrupt forward slope. Ce and Thl separate; Ce not expanded dorsolaterally. Prosome about 2Vs length of urosome. Anal segment not reduced, one-half length of caudal ramus. Al described by Tanaka (1956), typical left-handed appendage; length unknown. Segment 25 not re- duced. PI as figured by Wheeler (1970), except typical out- er seta on B2, and Ri lobe with 1-3 terminal points. Rel outer spine short, reaching slightly beyond midlength of Re2. P2 Rel outer spine typical, not forming strongly bent hook. P4 Bl without inner seta. P5 essentially uniramus, left-handed; similar to that on S. antarcticus male except rudimentary left Ri. Left Re2 nearly equaling length of Rel. Remarks Tanaka (1956) described a new species, S. longipes, for one male taken in a sample with three female S. angusticeps. The similarity of P5 to S. antarcticus male P5, and the 3 inner setae on PI Ri on S. angusticeps female (male otherwise unknown) and S. longipes (sample 50, Table 3) definitely relates S". longipes to the group including S. angusticeps, S. ant- arcticus, S. magnus, and S. validus. Males have been assigned to the latter three species, and S. longipes is considered in the present study to be the male of S. angusticeps. Spinocalanus longipes has also been reported by Grice and Hulsemann (1967), who found female S. angusticeps in adjacent areas. Wheeler (1970) reported one male Spinocalanus sp. Johnson; Wheeler's specimen lacked Re on the longest P5, which he said was the right. Wheeler's figure of P5 suggests a rudimentary Ri; the figure of Pi essentially agrees with male Pi described above, with short Rel outer spine. Wheeler reported female S. angusticeps from the sample which contained this male. Distribution Records of S. angusticeps are summarized below: Pacific Ocean Northwest: Tanaka (1953, 1956), 0-1,000 m. Southwest: Present study, 0-1,000 m. Indian Ocean West: Grice and Hulsemann (1967), 750-2.000 m. Atlantic Ocean Northeast: Sars (1920; 1924, 1925). 0-2.000 m. East: Grice and Hulsemann (1965), 450-3.000 m. —Roe (1972b), 660-950 m. Northwest: Grice (1963), 700-1.200 m. — Wheeler (1970). 2.000-4,000 m. (14. Spinocalanus caudatus Sars, 1920) See Spinocalanus oligospinosus Park. 197(1. (15. Spinocalanus abyssalis var. pygmaeus Farran, 1926) See Spinocalanus longicornis Sars. 1900. 43 (16. Spinocalanus heterocaudatus Rose, 1937) See Spinocalanus magnus Wolfenden, 1904. (17. Spinocalanus pacificus Mori, 1942) See Spinocalanus magnus Wolfenden, 1904. (18. Spinocalanus stellatus Brodsky, 1950) See Spinocalanus horridus Wolfenden, 1911. (19. Spinocalanus spinipes Brodsky, 1950) Female, see Spinocalanus horridus Wolfenden, 1911. Male, see S. magnus Wolfenden, 1904. (20. Spinocalanus pseudospinipes Brodsky, 1950) See Spinocalanus breuicaudatus Brodsky, 1950. 21. Spinocalanus similis Brodsky, 1950 (Figures 86-97, 154) Spinocalanus similis Brodsky, 1950, p. 128-129, fig. 47. Spinocalanus similis var. profundalis Brodsky, 1955 (part), p. 185-187, fig. lg; male only. Type Specimen: Unknown. TYPE LOCALITY: Northwest Pacific, 1,000-4,000 m. Material Studied: 2 females (1.08, 1.15 mm), sam- ple 30 (Table 3); 4 females (1.03-1.24 mm, mean = 1.12 mm) and 2 males (1.53 mm), sample 31. Description Pi Ri with 2 inner setae; P2 Ri2 with no outer seta. Female.— Length 1.03-1.5 mm (Figs. 86, 87). Prosome in dorsal view robust ovoid. Head rounded in dorsal view; in lateral view with irregular, gentle forward slope. Th4 and Th5 separate; Th5 lateral cor- ners slightly prolonged; Thl-Th4 without lateral spinules. Prosome length slightly more than 3 times urosome. Genital segment slightly wider than long, protruding ventrally one-half depth of rest of urosome; caudal rami about as long as wide; right ramus sometimes larger than left (Brodsky, 1950). Caudal setae incomplete on specimens examined. Supra-anal plate without fringe of long hairs. Al exceeding caudal rami by about 1 segment; ter- minal segments (Fig. 154) of moderate length and width. Apparent deviation: I-2s; remainder as Table 4. A2 Rel with 1 seta. Mxl gnathobase with 4 proximal setae thick, about one-half thickness of strong outer setae; probably thin seta distal from cluster of long spinules on midanterior surface; posterior surface with several clusters of long and short spinules and spines. Mx2 with outer seta; posterior surface of lobe-5 base with spines. Mxp (Fig. 88) with transverse spine-comb on midouter Bl; longitudinal row of stiff, short hairs on midanterior surface; proximal and distal row of den- ticles on posterior surface. B2 with strong transverse spine-comb and longitudinal row of stiff hairs. Ri4-5 outer setae nude, bladelike, tapering toward each end; spinules on inner edge of Ri5. PI (Fig. 89) Bl with few setules and spinules on dis- tal surface; inner margin nude. B2 anterior and posterior inner surface with hairs and setules. Rel-2 anterior surface with outer distal row of spinules; Re3 posterior surface with 3-4 spines (present material) or nude (Brodsky, 1950). Re outer spines reaching beyond base of following spines. Ri surface nude. P2 (Fig. 90) -P3 (Fig. 91) Bl outer surface with 3 rows of spines; inner surface with setules, inner margin with hairs. P4 (Fig. 92) Bl outer anterior sur- face with few short spines; inner surface with few long hairs. P2-P4 B2 nude. Re2 posterior surface on P3-P4 with proximal row of 12 "wide lamellar spines" (Brodsky, 1950), but ap- parently not as wide as those on S. abyssalis or S. longicornis (Brodsky, fig. 47); distal row of few small spines. Re3 posterior surface on P2-P4 with 3 rows of 4-5 spines (Brodsky, 1950, for P3-P4). P3-P4 Ri3 posterior surface with proximal row of thin spines, distal row of few larger spines. Male. — Length 1.4-1.6 mm (Figs. 93, 94). Prosome anterior irregular in dorsal view, posterior somewhat rectangular; in lateral view with gentle forward slope. Ce and Thl separate. Prosome length 1% times Spinocalanus similis Figure 86. — Female, habitus, dorsal view; sample 30 (Table 3); scale B. Figure 87. — Female, habitus, lateral view; sample 30; scale B. Figure 88.— Female, Mxp Bl-2; sample 30; scale D. Figure 89.— Female, PI; sample 30; scale D. Figure 90.— Female, P2; sample 30; scale D. Figure 91.— Female, P3; sample 30; scale D. Figure 92.— Female, P4; sample 30; scale D. Figure 93. — Male, habitus, dorsal view; sample 31; scale B. Figure 94. — Male, habitus, lateral view; sample 31; scale B. Figure 95. — Male, Mn blade; sample 31; scale G. Figure 96.— Male, Mxp Bl-2; sample 31; scale D. Figure 97. — Male, P5; sample 31; scale D. 44 45 urosome. Caudal rami length IV2 times width, symmetrical. Caudal setae incomplete on specimens examined; short stiff hair on outer margin of rami may be remnant of sixth seta. Al reaching about end of prosome, left segment 20 merging with segment 21, typical of right-handed males. IV-2s, 2e; VHI-2S, 2e; XV-ls, le; XIX-ls, le; right XX-le; left XX-0; remainder as Table 4. Mn blade (Fig. 95) reduced; 7-8 short, subequal teeth; short dorsal seta. Mxp (Fig. 96) Bl-2 without spine-combs, setae of reduced size; longitudinal row of stiff hairs on B2. Outer setae nude, reduced; no spinules on inner edge of Ri5; large inner setae of last segments not inflated. PI as on female, except Re outer spines shorter and wider; spinules not seen on Rel-2 distal anterior sur- face; 4-5 spines on Re3 posterior surface. P2-P3 Bl with longer, thinner, and more numerous spines than on female; B2, Rel, and Ril as on female. P2 Ri2 proximal row of spines of decreasing length from outer to inner edge, distal row of 5-6 large spines. Re2-3 lacking on specimens examined. P4 Bl nude inner surface and margin; few short setules on outer proximal surface, and minute spinules on outer distal anterior surface. Rel and Ril as on female. P5 (Fig. 97) biramus, right-handed: reaching mid- dle of urosome segment 2. Right Bl reaching one-third length of left B2; right B2 reaching middle of left Rel. Right and left legs about equal; left Re, including ter- minal blade, slightly longer than right; left Ri longer than right. Order of length, longest to shortest, of Re segments: left 3, 1 = 2; right 1 = 2, 3. Left Ri reaching two-thirds length of Re long terminal seta. Right Ri reaching middle of right Re3. Each Rel with short, flat seta on distal outer edge. Inner edge of right Re2 with long hairs. Each Re with 1 small and 1 moderate bladelike terminal setae. Remarks Brodsky (1950) did not describe Mxp nor P2 Ri; the present specimens agree with Brodsky's description of S. similis except for the symmetry of caudal rami and the posterior surface spines on PI Re3. Brodsky (1955) briefly redescribed the male S. similis as S. similis var. profundalis; the female which he included in this variety is considered in the present study as 5. brevicaudatus. Distribution The only published records of S. similis are by Brodsky, and the distribution of this species is sum- marized below: Pacific Ocean Northwest: Brodsky (1950; 1952a, b; 1955, 1957), 0-8,500 m. Central: Present study, 0-1,000 m. Brodsky (1957) also indicated that S. similis was found in "abyssal and lower bathypelagic" layers in the Bering Sea. (22. Spinocalanus dorsispinosus Brodsky, 1950) See Spinocalanus horridus Wolfenden, 1911. (23. Spinocalanus sp.? Brodsky, 1950) (Spinocalanus abyssalis. — Tanaka, 1937) Female, see Spinocalanus horridus Wolfenden, 1911. Male, see S. brevicaudatus Brodsky, 1950. (24. Spinocalanus longispinus Brodsky, 1950) See Spinocalanus horridus Wolfenden, 1911. 25. Spinocalanus elongatus Brodsky, 1950 (Figures 98-113, 155) Spinocalanus elongatus Brodsky, 1950 (part), p. 132- 133, fig. 52; female and "male variant" but not "male." — Vidal, 1971, p. 18, 23, fig. 46-47, 50-51, 56-57. Type Specimen: Unknown. TYPE LOCALITY: Central Arctic Ocean, 50 m. Material Studied: 8 females (1.50-1.80 mm, mean = 1.65 mm), sample Al (Table 1); 6 females (1.58-1.80 mm, mean = 1.64 mm), sample A2; 20 females (1.50- 1.80 mm, mean = 1.61 mm), 1 female stage V (1.37 mm), and 2 males (1.45, 1.55 mm), sample A4; 9 females (1.58-1.70 mm, mean = 1.64 mm), 1 female stage V (1.54 mm), and 1 male (1.62 mm), sample A5; 1 female stage V (1.70 mm), 11 female stage IV and 2 male stage IV (1.15-1.23 mm), 12 stage III (0.8-1.05 mm), and 2 male stage V (1.48, 1.53 mm), sample A6; 2 males (1.60 mm), sample A20; 1 male (1.68 mm), sample A21; 1 male (1.56 mm), sample A23; 7 males (1.60-1.76 mm, mean = 1.66 mm), sample A24; 2 males (1.60, 1.64 mm), sample A27; 1 female (1.68 mm), sample A31; 2 males (1.70, 1.72 mm), sample A33; 12 females (1.50-1.72 mm, mean = 1.61 mm) and 1 male (1.68 mm), sample A48. Description PI Ri with 2 inner setae; P2 Ri2 with no outer seta. Female— Length 1.50-1.80 mm (Figs. 98, 99). Prosome in dorsal view ovoid, head somewhat trun- 46 47 Figure 105.- Figure 106.- Figure 107.- Figure 108.- Figure 109.- Figure 110.- Figure 111.- Figure 112.- Figure 113. -Female, Mxp; scale D. -Male, habitus, dorsal view; scale A. -Male, habitus, lateral view; scale A -Male, Mxp; scale D. -Male, PI, left; scale D. -Male, P2, left; scale D. -Male, P3, left; scale D. -Male, P5; scale D. -Male stage V, P5; scale D. 48 cate. In lateral view with flat, gentle forward slope. Th4 and Th5 separate; Th5 lateral corners slightly prolonged; Thl-Th4 without lateral spinules. Prosome length ahout 3% times urosome. Genital seg- ment as long as wide, protruding ventrally one-third depth of rest of urosome. Caudal rami as long as wide, left ramus slightly larger than right. Left terminal sec- ond from inner seta thickest, length ca. 2.1 mm; left and right terminal second from outer seta relatively thick to near tip; right inner seta larger than left, with long plumes. Al exceeding caudal rami by about 4 segments; ter- minal segments (Fig. 155) elongate. Apparent deviation: I-ls; remainder as Table 4. A2 Rel with 1 seta. Mn palp with inner row of long setules on distal anterior surface of B2 and on surface of Ril. Blade (Fig. 100) with 5 outer subequal teeth, ventral tooth 1- pointed, others at least 2-pointed; 3 narrow teeth, followed by outer seta, dorsally; proximal anterior surface with clusters of stiff hairs. Mxl gnathobase with 4 proximal setae on posterior surface, without seta on distal anterior surface. Mx2 with setule-covered proximal hump and outer seta; posterior surface of lobe-5 base without spines. Mxp (Fig. 105) Bl with 3 transverse spine-combs on distal outer surface; proximal transverse row and dis- tal oblique row of setules on anterior surface; inner distal hump with row of denticles. B2 with strong transverse spine-comb about one-third length on out- er surface; inner longitudinal row of stiff hairs. Ril with distal row of stiff hairs; Ri3 with only 2 setae; Ri5 with inner spinules. Ri4-5 outer setae nude, bladelike, tapering toward each end. Pi (Fig. 101) Bl-2 anterior surface with few rows of setules; inner margin with hairs. Rel-2 anterior sur- face with outer distal row of spinules; Re3 surface nude. Rel-2 outer spines barely reaching bases of following spines. Ri distal anterior surface with few short setules. P2 (Fig. 102) Bl outer surface with few spinules and setules, inner margin with hairs; B2 nude. P3 (Fig. 103) Bl outer surface with few spines, spinules, and setules, inner margin with hairs; B2 nude. P4 (Fig. 104) Bl proximal inner posterior surface with row of long and short setules; inner margin without hairs; B2 nude. P3-P4 Re2 posterior surface with proximal row of large, fairly wide spines and distal row of smaller spines. P2 Re3 rarely with few small proximal spines, usually only with 2 distal rows of spines. P3-P4 Re3 with 3 evenly spaced rows of spines, spine-size decreasing distally. Re terminal seta with following number of primarv teeth: P2 (37-44); P3 (38-49); P4 (40-44). Male.— Length 1.4-1.76 mm (Figs. 106, 107). Prosome anterior irregular in dorsal view, posterior somewhat rectangular; in lateral view with gentle forward slope. Ce and Thl partly fused. Prosome length 2 1 i times urosome. Caudal rami about as long as wide, symmetrical. Caudal setae symmetrical. Al reaching distal edge of urosome segment 1; seg- ment 1 without cluster of spinules on anterodowil surface. IV-2s, 2e; VIII-2s, 2e; XV-2s, le; XIX le; right XX-0; left XX-le; remainder as Table 4. A2 Rel with reduced seta. Mn blade and Mxl gnathobase reduced. Mx2 with fragile outer seta; setae of reduced length, terminal setae thicker than on female; lobe surfaces nude. Mxp (Fig. 108) Bl setae of reduced length, surfaces nude. B2 with longitudinal row of long stiff hairs. Longest setae of Ri2-5 nude, not inflated; Ri4-5 outer setae reduced, nude; no spinules on inner Rio. Ri3 with 3 setae. PI (Fig. 109) Bl-2 with fewer outer setules on anterior surface than on female; Rel-3 and Ri surface nude. Rel outer spine reaching about one-half length of Re2. Re2 outer spine stronger than on female. P2 (Fig. 110) Re2 with cluster, not row, of ca. 9 spines on posterior surface. Re2-3 outer spines longer than on female, reaching to base of following spines. Spiniform terminal edge of Re3 longer than on female. P3 (Fig. Ill) Re2 posterior surface proximal spines wider than on female; distal spines in cluster. Re3 similar to P2 Re3. P4 Bl-2 nude. Rel and Ri as on female. Re2-3 miss- ing on specimens examined. Re terminal seta with finer serrations than on female: P2 (63-70); P3 (ca. 64). P5 (Fig. 112) biramus, left-handed; reaching to end of urosome segment 2. Left Bl reaching middle of right B2; left B2 reaching one-third length of right Rel. Left and right legs about equal. Right Re, in- cluding long terminal blade, slightly longer than left; right Ri much longer than left. Order of length, longest to shortest, of Re segments: left and right 1 = 2, 3. Left Ri reaching two-thirds length of left Re2. Right Ri reaching two-thirds length of longest right Re terminal blade. Each Rel with short seta on outer dis- tal edge. Inner edge of left Re2-3 with long hairs. Each Re with 1 small and 1 moderate bladelike terminal setae. Male stage V.— Length 1.48-1.53 mm. P5 (Fig. 113) biramus, symmetrical. Re imperfectly 2- segmented; outer distal edge of Rel with short, flat seta; 2 short, unequally wide, terminal setae. Ri with trace of segmentation; terminal seta reaches nearly to end of Re. Remarks In addition to the female of >». elongatus, Brodsky ( 1950) described two distinct males under the name >' elongatUS. The "typical" male is S. polans (see below); the "variant" male is S. elongatus. 49 Distribution Spinocalanus elongatus is the only species of the present study that is known only from the Arctic Ocean. Bogorov (1946a) reported two female Arctic S. abyssalis, which he distinguished, by their size, from S. longicornis; this is considered a record of S. elongatus in the present study, since no other likely species has been found in the Central Arctic. The records of S. elongatus are summarized below: Arctic Ocean Central: Brodsky (1950, 1957), 25-3,000 m. Eurasian Basin: Bogorov (1946a), 10-750 m. Canadian Basin: Brodsky and Nikitin (1955), 100-3,826 m. —Dunbar and Harding (1968), 175-2,000 m. — Vidal (1971). —Present study, 300-2,500 m. 26. Spinocalanus polaris Brodsky, 1950 (Figures 114-129, 156) Spinocalanus polaris Brodsky, 1950, p. 133, fig. 53. — Grice and Hulsemann, 1965, p. 229-230, fig. 7g. —Park, 1970, p. 493, fig. 71-77. —Vidal, 1971, p. 18, fig. 36-37, 39. Spinocalanus elongatus Brodsky, 1950 (part), p. 132- 133, fig. 52; "male" but not "male variant." Type Specimen: Unknown. Type Locality: Central Arctic Ocean, 800-2,500 m. Material Studied.- 1 male (1.60 mm), sample A17 (Table 1); 2 males (1.60, 1.64 mm), sample A26; 3 females (1.28, 1.32, 1.40 mm) and 1 male (1.68 mm), sample A31; 12 females (1.36-1.56 mm, mean = 1.47 mm) and 1 male (1.56 mm), sample A48; 7 males (1.44-1.68 mm, mean = 1.55 mm), sample A50; 6 females (1.30-1.42 mm, mean = 1.36 mm) and 7 males (1.45-1.55 mm, mean = 1.49 mm), sample A51. Description Pi Ri with 1 inner seta; P2 Ri2 with no outer seta; P3-P4 Ri3 with 1 outer seta. Female.— Length (1.02?-) 1.17-1.60 mm (Figs. 114, 115). Prosome in dorsal view fairly robust ovoid; head abruptly narrow, rounded. In lateral view with fairly abrupt forward slope. Th4 and Th5 separate; Th5 lateral corners prolonged, reaching midgenital seg- ment; Thl-Th4 without lateral spinules. Prosome length about 3% times urosome. Genital segment slightly longer than wide, protruding ventrally one- half depth of rest of urosome. Caudal rami length IV3 times width, symmetrical. Caudal setae broken in all specimens; bases symmetrical. Al exceeding caudal rami by about IV2 segments; terminal segments (Fig. 156) of moderate length and width. Armature as Table 4. A2 Rel with 1 seta. Mn B2 surface without setules; Rel outer distal sur- face with row of setules. Blade (Fig. 121) with 4 sub- equal outer teeth, ventral tooth 1-pointed, others 3- pointed; 4 shorter 2-pointed teeth, followed by dorsal seta; midanterior surface with transverse row of long spinules. Mxl gnathobase (Fig. 122) with 4 proximal setae on posterior surface, without seta on distal anterior sur- face. Posterior surface with proximal cluster of strong denticles, central cluster of smaller denticles, and out- er cluster of spinules. Mx2 with small setule- and hair-covered proximal hump; no outer seta seen (Park, 1970, fig. 74, shows outer seta). Distal hook on lobe-1 (Fig. 116) promi- nent. Posterior surface of lobe-5 base without spines. Mxp (Fig. 123) Bl-2 without transverse spine- combs. Bl anterior surface with 1 proximal and 3 dis- tal transverse rows of long setules; outer longitudinal row of setules and inner longitudinal row of spines about midlength; longitudinal row of short, stiff hairs near inner distal edge. B2 with longitudinal row of short, stiff hairs. Ril inner distal edge with row of stiff hairs. Ri4-5 outer setae unmodified, thin, relatively short, probably lightly plumose; Ri5 inner edge without spinules. PI (Fig. 117) Bl-2 anterior surface with few rows of setules; Bl inner margin with setules; B2 inner margin with setules and hairs. Rel anterior surface with outer distal row of spines; Re2 posterior surface with distal row of spines; Re3 surface nude. Rel -2 out- er spines reaching base of following spine. Ri inner midlength surface with spinules. Re terminal seta relatively coarsely serrate. P2 (Fig. 118) Bl outer surface with setules, inner margin with hairs. B2 inner posterior surface with few setules. Re2 distal posterior surface with straight row of spines; Re3 distal anterior and posterior surface with group of small spines. P3 (Fig. 119) Bl proximal outer surface with few setules; inner margin with hairs. Re2 posterior surface with 1 distal row of long spines; Re3 distal anterior and posterior surface with group of small spines. Ri3 proximal posterior surface with few small spines. P4 (Fig. 120) Bl inner margin with hairs; posterior surface with transverse row of long spinules. Re2 posterior surface with 1 distal row of long spines; Re3 posterior surface with 3 evenly spaced longitudinal groups of denticles. Re terminal seta with following number of primary teeth: P2 (23-29); P3 (ca. 20); P4 (ca. 20; secondary teeth relatively large, irregular.) Male.— Length 1.44-1.68 mm (Figs. 124, 125). Prosome anterior somewhat rounded in dorsal view, posterior somewhat rectangular; in lateral view with gentle forward slope. Ce and Thl partly fused. 50 Spinocalanua polaris, female Figure 114.— Habitus, dorsal view; scale B. Figure 1 15.— Habitus, lateral view; scale B Figure 116.— Mx2, lobe-1; scale E. Figure 117.— PI; scale E. Figure 118.— P2; scale E. Figure 119.— P3; scale E. Figure 120.— P4; scale E. 51 Figure 121.- Figure 122.- Figure 123.- Figure 124.- Figure 125.- Figure 126.- Figure 127.- Figure 128.- Figure 129.- Spinocalanu8 polaris -Female, Mn blade; scale F. -Female, Mxl gnathobase; scale F. -Female, Mxp; scale E. -Male, habitus, dorsal view; scale B. -Male, habitus, lateral view; scale B. -Male, Mn blade; scale F. -Male, Mxp; scale E. -Male, PI; scale E. -Male, P5; scale E. 52 Prosome length 2 times urosome. Caudal rami slightly longer than wide, symmetrical. Caudal setae in- complete on specimens examined, bases symmetri- cal. Al reaching middle of urosome segment 2; left seg- ment 20 merging with segment 21, typical of right- handed males; segment 25 reduced. IV-2s, le; VIII-2s, 2e; XV-ls, le; XIX-ls, le; right XX-ls; left XX-0; remainder as Table 4. A2 Rel with 1 seta. Mn blade (Fig. 126) reduced; dorsal tooth relatively long. Mx2 long terminal setae much thicker than on female. Mxp (Fig. 127) Bl setae of reduced length, surface nude. B2 with longitudinal row of stiff hairs. Ri2-5 in- ner setae inflated about one-third length; Ri2 setae and Ri3 proximal seta distally plumose; other inflated setae nude. Ri2 apparently with only 3 setae, Ri3-5 apparently with only 2 setae. Ri4-5 outer setae and Ri5 terminal seta apparently absent. No spinules on inner Ri5. PI (Fig. 128) Bl with only few hairs on inner margin. B2 anterior surface with outer distal row of spinules; inner edge with setules and hairs. Re and Ri surface nude. Re3 terminal seta not coarsely ser- rate. P2 as on female except Re2 surface spines longer, and Re3 terminal seta with 66-69 primary teeth. P3 as on female except Re3 outer spines longer, and terminal seta with ca. 73 primary teeth. P4 Bl nude (one specimen with rudimentary right inner seta). Re2-3 missing on specimens examined, remainder as on female. P5 (Fig. 129) biramus, right-handed; reaching end of urosome segment 2. Right Bl reaching three- fourths length of left B2; right B2 reaching left Re2. Right leg longer than left; right and left Re, including terminal blade, about equal length; right and left Ri about equal length. Order of length, longest to shortest, of Re segments: left 2, 3, 1; right 1 = 2, 3. Left Ri reaching middle of left Re2. Right Ri reaching one-third length of right Re2. Each Rel with short, flat seta on distal outer edge. Left Rel-2 more or less fused; inner edge of Rel with few setules. Inner edge of right Re and Ri with hairs. Each Re with 1 small and 1 moderate bladelike terminal setae. Remarks Brodsky (1950) based his description of S. polaris solely on females, and considered S. polaris male as one of two types of male S. elongatus. Brodsky and Nikitin (1955) reported male S. polaris, so that they may have recognized this error. Park's (1970) lengths of females (1.02-1.06 mm) have been questioned in the above description; these should be verified, in view of the differences between some of his published lengths and present measurements of the same specimens. Distribution Records of S. polaris are summarized below: Arctic Ocean Central: Brodsky (1950, 1957), 200-3,000 m. Canadian Basin: Brodsky and Nikitin (19 0-3,826 m. —Dunbar and Harding (1968), 900-3,000 m. — Vidal (1971). —Present studv, 1,500-3,000 m. Atlantic Ocean East: Grice and Hulsemann (1965), 1,500-5,000 m. Caribbean Sea: Park (1970), 980-2,800 m. Brodsky (1957) recorded 6'. polaris, with some doubt, from the north and northwest Pacific. Some Mimocalanus species look like S. polaris, and if specimens are damaged, they could easily be con- fused. 27. Spinocalanus brevicaudatus Brodsky, 1950 (Figures 130-136, 157) Spinocalanus longicornis. — Sars, 1901, p. 22-23, pi. 12. Not S. longicornis Sars, 1900. Spinocalanus abyssalis. —Sars, 1903, p. 157-158, suppl. pi. 3, fig. 2. — van Breemen, 1908 (part), p. 28-29, fig. 27a, c (only). —With, 1915, p. 69-71, fig. 15; pi. 1, fig. lOa-b. —Rose, 1933, p. 84-85, fig. 42. — Tanaka, 1937 (part), p. 253-254, fig. 4d-e; male only. —Davis, 1949, p. 21-22, fig. 23-24. — Farran and Vervoort, 1951a (part), p. 2-3, fig. la-d, i (only). — Grice and Hulsemann, 1965, p. 229, fig. 7a. Not S. abyssalis Giesbrecht, 1888. Spinocalanus major Esterly, 1906, p. 55. Spinocalanus pseudospinipes Brodsky, 1950, p. 127- 128, fig. 46. Spinocalanus sp.? Brodsky, 1950 (part), p. 130-131. fig. 50; male only. Spinocalanus brevicaudatus Brodsky, 1950. p. 134- 136, fig. 55. — Semenova, 1962, p. 1571-1574, fig. 1- 4. Spinocalanus similis var. profundalis Brodsky, 1955 (part), p. 185-187, fig. la-c; female only. TYPE SPECIMEN: Lectotype, female (1.60 mm) OSLO F4923a. TYPE LOCALITY: Northeast Atlantic (Osterfjord). 400- 600 m. Material Studied: 3 females (1.55, 1.57. 1.60 mm). sample 3 (Table 3); paralectotype, female (1.60 mm. after Sars, 1901), on slide, sample 16; paralectotype. female, on slide, sample 17: lectotype. female (1.60 mm) and 1 male stage V (1.40 mm), sample 18; 2 paralectotypes, females (1.60 mm), sample 19; paralectotype, male (1.60 mm, after Sars, 19031. on slides, sample 20; 1 female (1.42 mm), sample 21; 1 53 135 Spinocalanua brevicaudatus Figure 130.— Female, Mxp Bl-2; sample 17 (Table 3); scale D. Figure 131.— Female, P2; sample 17; scale D. Figure 132.— Male, PI, left; sample 20; scale D. Figure 133.— Male, P2, left; sample 20; scale D. Figure 134.— Male, P3, left; sample 20; scale D. Figure 135.— Male, P4, left; sample 20; scale D. Figure 136.— Male, P5; sample 20; scale D. female (1.45 mm), sample 25; 1 female (1.47 mm), sample 26; 1 female (1.68 mm), sample 30; 1 female (1.95 mm) and 1 male (1.90 mm), sample 31; 1 female (1.85 mm), sample 33. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 1.40-2.4 mm. Prosome in dorsal view ovoid, head slightly truncate. In lateral view with fairly abrupt forward slope. Th4 and Th5 separate; Th5 lateral corners rounded, not prolonged; Thl-Th4 without lateral spinules. Prosome/urosome ratio variable; prosome 3-4 times urosome. Genital segment as long as wide, protruding ventrally one-half depth of rest of urosome. Caudal rami as long as wide, symmetrical. Caudal setae symmetrical. Al exceeding caudal rami by about 2 segments; ter- minal segments (Fig. 157) of moderate length and width. Most segments damaged on material studied. A2 Rel with 2 setae. Mx2 posterior surface of lobe-5 base apparently without spines. Mxp (Fig. 130) with typical number of setae; Bl surface with few rows of setules. B2 with transverse spine-comb about one-third length on outer surface; inner longitudinal row of stiff hairs. Ri4-5 outer setae nude, bladelike, tapering toward each end; not as shown by Sars (1901). Ri5 with inner spinules. PI similar to that in S. elongatus, Re3 posterior sur- face without spines. Sars' (1901) figure lacks 1 inner seta on Re3. P2 (Fig. 131) Bl outer surface with few spinules; in- ner posterior surface with setules. P3 Bl outer surface with few setules, spinules, and weak spines; inner 54 margin with hairs. P4 Bl proximal inner posterior sur- face with row of long and short setules; inner margin without hairs. P4 B2 posterior surface nude or with several thin setules. P3-P4 Re2 posterior surface with proximal row of strong, fairly wide spines and distal row of smaller spines. Ri2-3 surface spines shorter than on S. elongatus. P2 Re3 with 2 distal rows of spines. P3 Re3 with 3 evenly spaced rows of spines as on S. elongatus. P4 Re3 posterior surface with 3 groups of spines, smaller and fewer than on S. elongatus. Outer spines of P2-P4 Re2-3 longer than on S. elongatus. Re terminal seta with following number of primary teeth: P2 (ca. 25); P3 (21-25); P4 (25-27). Male.— Length (1.3?-) 1.5-1.90 mm. Prosome anterior irregular in dorsal view, posterior somewhat rounded; in lateral view with fairly abrupt forward slope. Prosome length 2 l A times urosome. Caudal rami about as long as wide, symmetrical. Caudal setae symmetrical. Al reaching middle of urosome segment 2; segment 1 with cluster of short spinules on anterodorsal surface. IV-2s, 2e; VIII-2s, 2e; XV-ls, le; XIX- ls?, le; right XX-damaged; left XX-le; remainder as Table 4. Mxp Ri4-5 outer setae reduced; Ri5 inner edge without spinules. PI (Fig. 132) similar to that of S. elongatus male ex- cept Rel outer spine shorter, Re2 outer spine larger. P2 (Fig. 133) as on female, except no anterior spinules on Re2, other surface spines slightly stronger, Re2 spines in cluster, Re3 with 3 rows of spines, and terminal seta with ca. 45 teeth. P3 (Fig. 134) as on female except no anterior spinules on Re2, Re2 with distal cluster of spines, and Re3 terminal seta with ca. 48 teeth. P4 (Fig. 135) as on female except Bl without inner seta and without inner transverse row of setules, Re2 with distal cluster of spines, Re3 surface spines stronger, and Re3 terminal seta with ca. 54 teeth. P5 (Fig. 136) biramus, left-handed. Left Bl reaching one-third length of right B2; left B2 reaching middle of right Rel. Right leg, including terminal blade, longer than left. Right Re longer than left; right Ri longer than left. Order of length, longest to shortest, of Re segments: left 1, 2, 3; right 2, 1 = 3. Left Ri reaching two-thirds length of longest left Re terminal blade. Right Ri reaching about to end of longest right Re terminal blade. Probably each Rel with short seta on outer distal edge. Right Rel-2 more or less fused. Inner edge of left Re2-3 with long hairs. Each Re probably with 1 small and 1 moderate bladelike terminal setae. Remarks Sars (1901) described a Norwegian Spinocalanus species as 5. longicornis, later (1903) adding the male and changing the name to .S'. abyssalis. Sare (1901) neglected to describe the outer seta of P2 Ri2 of his Norwegian specimens; a remounting of his prepara- tion clearly showed this seta (Fig. 128). Sars also failed initially to describe the surface spination of P2- P4 Re on his female specimens. These omissions, and the incomplete descriptions of S. abyssalis Giesbrecht, masked the distinctness of Sars' specimens. Most subsequent workers seemed to follow the authority of Sars, who incorrectly equated the Norwegian species (common in the North Atlantic) with both S. abyssalis Giesbrecht and .S'. longicornis Sars, 1900. Farran (1926) recognized two size groups in the Northeast Atlantic, the larger representing Sars' Norwegian species. Unfortunately Farran retained the name abyssalis for this group, renaming the smaller forms abyssalis var. pygmaeus (see S. longicornis Sars, 1900). Sars' (1903) figure of male P5 was poorly printed, the end of left Ri not shown well; therefore, later copies of this figure (Rose, 1933; Brodsky, 1950; Farran and Vervoort, 1951a) failed to show the full length of left Ri. This specimen's appendage has been redrawn in the present study (Fig. 133). Van Breemen (1908), Rose (1933), and Farran and Vervoort (1951a) copied some of Sars' figures, although they included figures or descriptions of S. abyssalis Giesbrecht and S. longicornis as well. With (1915) included a brief description of male stage V. Tanaka (1937) apparently combined one female 5. horridus (see above) and one male S. brevicaudatus as S. abyssalis. Brodsky (1950) recognized that neither represented S. abyssalis Giesbrecht, and considered Tanaka's specimens "Spinocalanus sp.?" Davis (1949) correctly illustrated S. brevicaudatus P2 Ri2 with 1 outer seta, but failed to note its dis- tinctness from S. abyssalis. Grice and Hulsemann (1965) gave a lateral view of the female S. brevicaudatus; their unpublished sta- tion list (1966) indicated a size of 1.4-2.0 mm (female) and 1.3 mm (male). This small male might be S. longicornis, and the length is questioned in the size range of males given above. Esterly (1906) very briefly, and without il- lustrations, described a single female as a new species. S. major. There is nothing in Esterly's description that sets S. major apart from S. abyssalis. except S. mayor's larger size. A. Fleminger examined the Ester- ly Collection and located a single slide which was labeled "Spinocalanus." This slide, now deposited in the U.S. National Museum (USNM 143692), con- tained only A2. Mxl, and Mxp. which appeared con- sistent with those appendages of S. brevicaudatus. However, the lack of Pi and P2 makes this identifica- tion uncertain. Spinocalanus brevicaudatus is known from the area in which Esterly found S. major; his brief description is consistent with that oi S. brevicaudatus. which is considered in the present 55 study to be conspecific. Since S. major was not well described, since the specimen from the Esterly Collec- tion is probably but not certainly S. major, and since 8. major has remained unused in the primary zoological literature for more than 50 yr, "S. major" should probably be suppressed. Brodsky's (1950) description of S. pseudospinipes is consistent with that of S. brevicaudatus, with the ex- ception that male P5 (left?) Ri, perhaps broken, is somewhat shorter (Brodsky, fig. 46). The description of S. pseudospinipes precedes by pages that of S. brevicaudatus, but the latter name is well established by recent redescriptions and records, and S. pseudospinipes, by comparison, has been little used and not redescribed. Brodsky (1950) was the first to point out that Sars' Norwegian specimens were distinct from both S. abyssalis and S. longicornis, and renamed them S. brevicaudatus. Brodsky did not have specimens, but referred only to Sars' (1901, 1903) descriptions. Semenova (1962) redescribed S. brevicaudatus from fresh specimens; Ri of P1-P2 were not described, but the species was defined essentially by size, body shape, P3-P4 Re2 posterior surface spines, and male P5. Brodsky (1955) briefly described S. similis var. profundalis; the description of the female is consistent with that of S. brevicaudatus. Distribution Most of the records summarized below are of S. abyssalis that probably represent S. brevicaudatus. In some cases the specimens have been reexamined; in other cases, a size, description, or other reference was reported which suggested S. brevicaudatus. Records which include specimens smaller than 1.40 mm are probably also of S. abyssalis Giesbrecht and/or S. longicornis Sars. Tanaka's (1937, 1953, 1956) record also includes S. horridus female. Brodsky (1950) considered Jespersen's (1934) record of S. abyssalis as S. brevicaudatus. In this and sub- sequent publications, Jespersen (1939a, b; 1940) did not report specimen size, but indicated that they were equivalent to S. abyssalis Giesbrecht and S. longicor- nis Sars, as well as S. brevicaudatus (as its syn- onyms). Therefore, Jespersen's records are considered as a probable mixture of species. Likewise, Stormer's (1929) and Ostvedt's (1955) records of S. abyssalis from nearby areas cannot be strictly interpreted. However, since the known distribution of S. brevicaudatus includes most of the areas sampled by Stormer, Jespersen, and 0stvedt, their records are in the following summary. Some of this material, es- pecially that of Jespersen (1934) from Baffin Bay, should be reexamined. Vervoort's (1951) record is not included in the following summary (see S. terranovae). Three records of S. longicornis are considered to be of S. brevicaudatus: Gran (1902), identified by G. O. Sars, perhaps the specimen in sample 21 (Table 3); Hoek (1906), identified by C. Wesenberg-Lund and G. O. Sars; and Ostenfeld (1916), identified by G. P. Farran. The records of Spinocalanus sp. by Figueira (1971) and Roe (1972a, b) are considered to be of S. brevicaudatus. Pesta (1927) cites Sars' Norwegian record of S. brevicaudatus as both S. abyssalis and S. longicornis. Grainger (1965) listed S. brevicaudatus from the Central Arctic, but this was only a reference to Bogorov's (1946a) record of S. abyssalis (E. H. Grainger, pers. commun.). Bogorov's specimens are considered in the present study to be S. elongatus, since S. brevicaudatus has not been reported from the Central Arctic. Park (1970) reported S. brevicaudatus from the Caribbean; four of his five specimens were examined in the present study; two are considered S. abyssalis and two S. aspinosus. Grice (1971) listed S. brevicaudatus from the Mediterranean Sea, but this was a misinterpretation of Rose's (1933) summary (G. Grice, pers. commun.). Pacific Ocean North: Brodsky (1950), 1,000-4,000 m. Northeast: Esterly (1906), 360 m. —Davis (1949), 0-2,300 m. —Fulton (1968), below 200 m. —von Vaupel-Klein (1970), 0-1,200 m. — Figueira (1971). —Present study, 0-400 m. Central: Present study, 0-1,000 m. Northwest: Tanaka (1937, 1953, 1956), below 200 m. —Brodsky (1952a, 1955, 1957), 0-8,500 m. — Furuhashi (1966), 298-775 m. Indo-Pacific: Vervoort (1946), 355-2,500 m. Indian Ocean West: Grice and Hulsemann (1967), 750-3,000 m. Atlantic Ocean North: Stormer (1929), 50-1,500 m. Jespersen (1934; 1939a, b; 1940), 0-1,500 m. — 0stvedt (1955), 100-2,000 m. —Semenova (1962), be- low 300 m. Northeast: Sars (1901, 1903), 400-600 m. — Gran (1902), 200-1,000 m. —Hoek (1906). — Farran (1908, 1920, 1926), 180-3,600 m. — Nordgaard (1912), 200-600 m. —Ostenfeld (1916). — Lysholm and Nordgaard (1921), 600-1,000 m. —Sars (1924, 1925), 0-4,000 m. — Runnstrom (1932), 100-400 m. —Grice and Hulsemann (1965), 180-4,000 m. —Present study, 1,260 m. East: Roe (1972a, b), 350-960 m. Southeast: Unteruberbacher (1964), 0-300 m. Central: Wheeler (1970), 2,000-4,000 m. Northwest: With (1915), 0-360 m. —Semenova (1962), below 300 m. —Grice (1963), 620- 1,200 m. —Wheeler (1970), 2,000-4,000 m. Caribbean Sea: Park (1970), 1,004-1,850 m. Some of Farran's specimens of S. abyssalis (=S. brevicaudatus) are in the collection of the National Museum of Ireland, Dublin (O'Riordan, 1969). 56 (28. Spinocalanu8 similis var. profundalis Brodsky, 1955) Female, see Spinocalanus breuicaudatus Brodsky, 1950. Male, see S. similis Brodsky, 1950. (29. Spinocalanus longipes Tanaka, 1956) (S. longipes Tanaka, 1953, nomen nudum) See Spinocalanus angusticeps Sars, 1920. (30. Spinocalanus? sp. Johnson, 1963a) See Spinocalanus antarcticus Wolfenden, 1906. 31. Spinocalanus abruptus Grice and Hulsemann, 1965 Spinocalanus abruptus Grice and Hulsemann, 1965, p. 227-229, fig. 6m-p. TYPE SPECIMEN: Holotype, female, BM 1965.4.20.34. TYPE LOCALITY: Northeast Atlantic, 1,000-2,800 m. Material Studied: Paratypes, 2 females (1.30-1.32 mm), sample 49 (Table 3). Description PI Ri with 2 inner setae. P2 Ri2 with 1 outer seta (?). Female. — Length 1.29-1.47 mm. Specimens ex- amined in present study damaged. Prosome in dorsal view ovoid. Th4 and Th5 separate. Th5 lateral corners prolonged, variable, more prolonged than shown by Grice and Hulsemann (1965). Thl-Th4 without lateral spinules. Prosome length 4-4 '2 times urosome. Caudal rami symmetrical. Mx2 with small spinule- and setule-covered hump on proximal outer edge; no outer seta seen; posterior surface of lobe-5 base without spines. Mxp Bl-2 without transverse spine-comb. Bl with few transverse rows of short setules (Grice and Hulsemann, 1965, fig. 6n). PI as shown by Grice and Hulsemann except outer setules on Bl surface, and B2 outer edge with short seta, anterior surface with outer distal row of spinules, inner surface with setules. P2-P4 rami missing on specimens examined. P2 perhaps with 1 outer seta on Ri2, although not clear on illustration by Grice and Hulsemann (1965, fig. 6p); Bl with inner seta, not shown by Grice and Hulsemann. Male. — Unknown. Remarks Spinocalanus abruptus is the only Spinocalanus species in the group with 2 inner setae on Pi Ri and 1 outer seta on P2 Ri2 that also has prolonged Th5 or is without transverse spine-comb on Mxp B2. Distribution Indian Ocean West: Grice and Hulsemann (1967), 1,000- 3,000 m. Atlantic Ocean Northeast: Grice and Hulsemann (1965), 1,000- 2,800 m. (32. Spinocalanus ovalis Grice and Hulsemann, 1965) See Mimocalanus ovalis (Grice and Hulsemann, 1965). (33. Spinocalanus ventriosus Grice and Hulsemann, 1967) See Mimocalanus ovalis (Grice and Hulsemann, 1965). (34. Spinocalanus sp. Grice and Hulsemann, 1967) Spinocalanus sp. Grice and Hulsemann, 1967, p. 22, fig. 35. Remarks Grice and Hulsemann (1967) briefly described a single male from the Indian Ocean as Spinocalanus species. This specimen was examined in the present study (sample 52, Table 3). Pi Re3 with a total of 4 setae, and therefore the specimen is not a Spinocalanus species. P2-P4 rami missing; P2 B2 dis- tal edge spiniform, somewhat similar to Clausocalanus species. Al segments beyond 10 are much longer than in Spinocalanus species. (35. Spinocalanus parabyssalis Park, 1 970) Female, see Spinocalanus longicornia Sars. 1900. Male, see S. abyssalis Giesbrecht. 1888. (36. Spinocalanus pteronus Park, 1970) See Spinocalanus uaitatiM Park, 1970. 57 37. Spinocalanus usitatus Park, 1970 (Figures 137-140) Spinocalanus pteronus Park, 1970, p. 487, fig. 43-50. Spinocalanus usitatus Park, 1970, p. 487-489, fig. 51-54. Type Specimen.- Holotype, female (1.62 mm), USNM 123785. TYPE LOCALITY: Caribbean Sea, 487-950 m. Material Studied: 1 female (1.88 mm), sample 31 (Table 3); 1 female (1.65 mm), sample 32; 1 female (1.68 mm), sample 34; holotype, female (1.62 mm), sample 55; 2 paratypes, females (1.38, 1.52 mm), sam- ple 56. Also the following type specimens of Spinocalanus pteronus: holotype, female (1.93 mm) and 1 paratype, female (1.80 mm), sample 54; 5 paratypes, females (1.70-1.82 mm, mean = 1.76 mm), sample 59. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female.— Length 1.38-1.93 mm (Figs. 137, 138). Body similar to S. spinosus; prosome in dorsal view somewhat robust ovoid, head slightly truncate; in lateral view with abrupt forward slope. Th4 and Th5 separate; Th5 lateral corners rounded, not prolonged. Thl-Th4 with lateral spinules, number decreasing posteriorly, approximately even right and left. Prosome length 2V2 -3 times urosome. Al incomplete on specimens examined. A2 Rel with 2 setae. Mx2 (Fig. 139) as on S. spinosus except posterior surface of lobe-5 base with spines. Mxp (Fig. 140) as on S. spinosus except Bl with transverse spine-comb on outer edge; oblique row of setules on midanterior surface; distal hump with oblique row of spinules on anterior surface; posterior surface near distal outer edge with transverse row of setules. B2 with strong transverse spine-comb; longitudinal row of spinules. Remaining segments as on S. spinosus. Pi as on S. spinosus. Male. — Unknown . Remarks Park (1970) described a new species, S. pteronus, apparently differing from S. usitatus only by the presence, in S. pteronus, of laterally extended edges of Ce. Ce of many Spinocalanus species has a tendency to bend up along the lateral edge; sometimes only one side is affected. This may be a reaction to the fixative, or it may be a function of age of the specimen. Ce of the holotype S. usitatus is turned out in this way on the left side. Therefore, since the present study con- siders that S. pteronus is essentially defined by a possible artifact, the name S. usitatus ("usitatus" = ordinary, usual) is retained for the species. 139 Spinocalanus usitatus, female Figure 137. — Habitus, dorsal view; sample 31; scale A. Figure 138. — Habitus, lateral view; sample 31; scale A. Figure 139. — Mx2, inner and terminal setae omitted; sample 31; scale D. Figure 140.— Mxp Bl-2; sample 31; scale D. 58 Distribution Vervoort (1946) reported one female (1.80 mm) S. spinosus with Mxp Bl with transverse spine-comb, in that it agreed with Mxp described by Wolfenden (1911) for S. horridus. Vervoort's specimen is con- sidered to be S. usitatus. Roe (1972a, b) reported S. spinosus from the east Atlantic and remarked that some of them may be closely related species as described by Park (1970). Pacific Ocean Northeast: Present study, 300-400 m. Central: Present study, 0-1,000 m. Indian Ocean Northwest: Vervoort (1946), 555-1,000 m. Atlantic Ocean Caribbean Sea: Park (1970), 487-1,900 m. 38. Spinocalanu,8 hoplites Park, 1970 Spinocalanus hoplites Park, 1970, p. 489, fig. 55-58. Type Specimen: Holotype, female (1.72 mm), USNM 123781. TYPE LOCALITY: Caribbean Sea, 980-1,900 m. Material Studied: Holotype, female (1.72 mm) and 3 paratypes, females (1.52, 1.60, 1.62 mm), sample 56 (Table 3). Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 1.52-1.78 mm. Similar to S. spinosus except Mxp Bl with transverse spine-comb. Differs from S. usitatus in that Mxp Bl spine-comb on very large tubercle, and Mx2 posterior surface of lobe-5 base without spines. Male. — Unknown. Remarks The development of a transverse spine-comb on Mxp Bl, in those species where it occurs, is variable; it is sometimes reduced or even lacking on Mxp of one side. Likewise, the spine-comb base is more or less prominent, and in S. hoplites is very pronounced. If it were not for the absence of spines on base of Mx2 lobe- 5, S. hoplites would be considered in this study as equivalent to 5. usitatus. Distribution 39. Spinocalanus aspinosus Park, 1970 Spinocalanus aspinosus Park, 1970, p. 489, fig. 59-61. Type Specimen: Holotype, female (1.38 mm), USNM 123780. TYPE LOCALITY: Caribbean Sea, 1,004-1,850 m. Material Studied: l female (1.41 mm), sample 53 (Table 3); holotype, female (1.38 mm), sample 54; 1 female (1.48 mm, after Park, 1970), on slide, sample 57. Description Pi Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 1.32-1.48 mm. Similar to S. hoplites except Thl-Th4 without lateral spinules. Mxp Bl with transverse spine-comb on conspicuous tubercle. Male. — Unknown. Remarks Two of the specimens examined in this study had been reported as S. brevicaudatus by Park (1970). Distribution Atlantic Ocean Caribbean Sea: Park (1970), 980-1,900 —Present study, 203-1,000 m. m. Atlantic Ocean Caribbean Sea: Park (1970), 980-2,800 m. Grice and Hulsemann (1965) reported some S. spinosus without integumental spinules from the northeast Atlantic. Probably those specimens belong- ed to S. aspinosus. 40. Spinocalanus oligospinosus Park, 1970 (Figure 158) ? Spinocalanus caudatus Sars, 1920, p. 3. — Sars, 1924. 1925, p. 31-32, pi. 8, fig. 10-17. —Rose, 1933. p. 85- 86, fig. 45. Spinocalanus oligospinosus Park, 1970, p. 491. fig. 62-65. Spinocalanus paraoligospinosus Hure and Scotto di Carlo, 1971, p. 584-585, nomen nudum. Spinocalanus neospinosus Grice, 1971, p. 276-280. fig. 2A-I. Type Specimen: Holotvpe, female (1.28 mm), USNM 123782. Type LOCALITY: Caribbean Sea, 487-950 m. Material Studied: Holotype. female (1.28 mm). sample 55 (Table 3). Also the type specimen o\ S. neospinosus: holotype. female (1.30 mm), sample 47. 59 Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female. — Length 1.20-1.50 mm. Similar to S. usitatus except S. oligospinosus Mx2 lobe-5 base without spines, and lateral prosomal spinules less numerous on right side. Left Thl, many distinct spinules; Th2-Th3, ca. 25 spinules; Th4, no spinules. Right Thl, many distinct spinules; Th2 few spinules; Th3-Th4, no spinules. Al terminal segments (Fig. 158) of moderate length and width. Apparent deviation: I-2s; remainder as Table 4. Mxp Bl spine-comb sometimes somewhat reduced on one side; holotype with distinct spine-comb left and right. P3 Re3 with 5 inner setae, not 4 as shown by Park (1970, fig. 65). Male. — Unknown. Remarks Sars (1920; 1924, 1925) described S. caudatus from one female collected in the western Mediterranean. Rose (1933) and Massuti (1939) cited this single record. This species has not been reported from the Mediterranean since the original record. It is possible that S. caudatus is conspecific with S. breuicaudatus, which has not been reported from the Mediterranean. It is also possible that S. caudatus is conspecific with S. oligospinosus which is common at the type locality of S. caudatus. Sars' (1924, 1925) description does not mention lateral prosomal spinules or Mxp Bl spine- comb, both of which could have been overlooked. The prosome/urosome ratio of S. caudatus is consistent with that of S. oligospinosus but not of S. breuicaudatus. The description is not precise enough to separate S. caudatus from other species; in this study, Sars' specimen is considered S. oligospinosus. The holotype S. caudatus, in poor condition, is on slide F5038, OSLO. Hure and Scotto di Carlo (1971) equated their earlier use of S. abyssalis with Grice's manuscript name for S. neospinosus, S. paraoligospinosus (G. Grice, pers. comraun.). Grice (1971, fig. 2E, G) did not note the presence of 4 posterior surface spines on PI Re3, and P3 Ri3 has 6 setae. Distribution Atlantic Ocean Mediterranean Sea: ?Sars (1920), 0-2,595 m. —Hure (1965), 200-1,000 m. —Hure and Scotto di Carlo (1968, 1969, 1971), 300-1,000 m. —Scotto di Carlo (1968), 200-1,000 m. — Grice (1971), 750-2,200 m. Caribbean Sea: Park (1970), 203-1,000 m. Wilson (1942) reported S. caudatus in seven samples from the northeast and central Pacific. Wilson's samples 87 and 132 were examined in the present study and found not to contain any Spinocalanidae. The other samples were not located. Furuhashi (1966) reported six female S. caudatus from the northwest Pacific; he also reported S. breuicaudatus (as S. pseudospinipes) from the same area, and perhaps the S. caudatus are S. breuicaudatus. (41. Spinocalanus paraoligospinosus Hure and Scotto di Carlo, 1971, nomen nudum) See Spinocalanus oligospinosus Park, 1970. (42. Spinocalanus neospinosus Grice, 1971) See Spinocalanus oligospinosus Park, 1970. 43. Spinocalanus terranovae, new species (Figures 141-147, 159) Spinocalanus abyssalis. — Bradford, 1971 (part), p. 17, fig. 22-27; 1.7-2.0 mm only. Not S. abyssalis Giesbrecht, 1888. Type Specimen.- Holotype, female (1.88 mm), BM 1930.1.1.293-404.a. Type Locality: Antarctic, Pacific Sector, 0-400 m. Material Studied: Holotype, female (1.88 mm) and 3 paratype females (1.85, 1.95, 1.98 mm), sample 6 (Table 3); 1 female (1.9 mm, after Bradford, 1971), on slide, sample 8; 1 paratype female (1.75 mm), sample 9, USNM 142701; 1 paratype female (1.80 mm), sam- ple 9, NZOI P261; 1 paratype female (2.0 mm, after Bradford), on slide, sample 9, NZOI P261; 1 paratype female (1.8 mm, after Bradford), on slide, sample 10, NZOI P261; 1 paratype female (1.75 mm, after Brad- ford), on slide, sample 12, NZOI P261; 1 male (1.9 mm, after Bradford), on slide, sample 12, NZOI P261. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta. Female.— Length (1.65?-) 1.70-2.20 mm (Figs. 141, 142). Prosome in dorsal view ovoid, head slightly trun- cate. In lateral view with moderately gentle forward slope. Th4 and Th5 separate; Th5 lateral corners rounded, not prolonged; Thl-Th4 without lateral spinules. Prosome length about 3V3 times urosome. Genital segment as wide as long, protruding ventrally one-third depth of rest of urosome; caudal rami length slightly greater than width, symmetrical. Caudal setae incomplete on specimens examined; bases appearing symmetrical. 60 Spinocalanua terranovae, female Figure 141. — Habitus, dorsal view; sample 6 (Table 8); scale A. Figure 142.— Habitus, lateral view; sample 6; scale A. Figure 143.— Mxp BI-2; sample 6; scale D. Figure 144.— PI; sample 6; scale D. Figure 145.— P2; sample 6; scale D. Figure 146.— P3; sample 6; scale D. Figure 147.— P4; sample 6; scale D. C 1„ , 147 61 Al exceeding caudal rami by nearly 2 segments; ter- minal segments (Fig. 159) of moderate length and width. Apparent deviation: I-2s; remainder as Table 4. A2 Rel with 2 setae. Mn B2 inner distal anterior surface and Ril surface with long setules. Blade broken on specimens exam- ined. Mxl gnathobase posterior surface apparently with 3 proximal setae; without seta on distal anterior sur- face. Mx2 with proximal setule-covered hump and outer seta; posterior surface of lobe-5 base without spines. Mxp (Fig. 143) Bl without spine-comb; anterior surface with several rows of short to long setules; dis- tal lobe with ascending oblique row of denticles followed by descending row of short stiff hairs. B2 out- er edge with transverse spine-comb; longitudinal row of stiff hairs. Ri4-5 outer setae nude, bladelike, taper- ing toward each end; spinules on inner edge of Ri5. PI (Fig. 144) Bl outer distal edge with few spines, inner margin with hairs. B2 inner surface with spines and inner margin with hairs; anterior surface with few central setules and outer distal row of spinules. Rel outer spine reaching base of following spine; Re2 outer spine reaching beyond base of following spine. Rel anterior surface with outer distal row of spinules. Re3 midposterior surface with 4-7 spines. P2 (Fig. 145) Bl inner and outer surface with rows of strong spines; inner margin with hairs. Re3 without proximal row of spines; midlength row with 5-6 spines; distal row with 4-6 spines. P3 (Fig. 146) Bl inner and outer surface with rows of strong spines; inner margin with hairs. Re2 posterior surface with proximal row of strong wide spines and distal row of denticles. Re3 posterior sur- face with 3 evenly spaced rows of spines; spines in- creasing in width and decreasing in number distally. Ri3 posterior surface with proximal row of 7-9 spines and distal row of 3-5 stronger spines. P4 (Fig. 147) Bl proximal outer surface with few rows of spines, smaller than on P2-P3; inner surface with setules. B2 posterior surface with inner transverse row of spinules as on S. horridus and S. spinosus. Re2 posterior surface with proximal row of strong wide spines and distal row of small spines. Re3 posterior surface with 3 evenly spaced rows of spines. Re terminal seta with the following number of teeth: P2 (30-34); P3 (28-29); P4 (33-38). Male. — Length 1.70-1.9 mm. Some Al segments damaged on specimen examined: IV-2s, 2e; VIII-2s, 2e; XV-?ls, le; XIX-ls, le; right XX-0; left XX-le. Mxp Ri5 inner edge with spinules. Other details known only from one damaged specimen reported by Bradford (1971). PI Re3 posterior surface with spines. P4 Bl with inner seta (Bradford, fig. 26). P5 shown by Bradford (fig. 22), in- complete; biramus, left-handed. Left Bl reaching middle of right B2; left B2 reaching nearly to end of right Rel. Ends of Re long terminal setae and Ri broken. Both Ri presumably reaching beyond Re3. Remarks Farran (1929) reported a group of large (1.8-2.06 mm) S. abyssalis from the Pacific Sector of the Ant- arctic. Some of these specimens were examined in the present study (sample 6, Table 3) and were found to be a new species, S. terranouae, described above. Vervoort (1951) reported three female S. abyssalis from the Atlantic Sector of the Antarctic. His specimens are considered S. terranouae in the present study, although they might be S. breuicaudatus or other species. The smallest specimen, 1.65 mm, is ten- tatively included in the length range of females. Vervoort's (1957) specimens of S. abyssalis are con- sidered S. terranouae in the present study. The length range of males reported by Vervoort (1.70-1.85 mm) is included in the above description. Vervoort's list of specimens from each sample indicates one male 2.20 mm, but this most likely should read "female." Bradford (1971) reported two size groups of S. abyssalis; only the larger was S. terranouae, the smaller was S. abyssalis Giesbrecht. Distribution Probable records of S. terranouae, previously known as a large S. abyssalis, are summarized below: Antarctic Pacific Sector: Farran (1929), 0-1,750 m. —Ver- voort (1957, 1965), 250-1,000 m. —Bradford (1971), 0-1,300 m. Indian Sector: Vervoort (1957, 1965), 100-1,500 m. — Tanaka (1960), 0-400 m. Atlantic Sector: Vervoort (1951), 0-900 m. Hardy and Gunther (1935) reported a Spinocalanus species from the South Atlantic (300-2,000 m) which might be S. terranouae. Etymology: The specific name, terranouae, from the British Antarctic ("Terra Nova") Expedition, 1910. Genus Monacilla Sars, 1905 Oxycalanus Farran, 1908. Hypsicalanus Wolfenden, 1911. TYPE SPECIES: Monacilla typica Sars, 1905. Description Rostrum 2-pointed. Pi Rel with outer distal spine, reduced in male; Rel-2 together longer than Re3; Ri with 3 inner setae. P2 Ri2 with 1 outer seta. P3-P4Ri3 with 2 outer setae. Posterior surface of certain 62 148 =t 160 Terminal segments of first antenna, female Figure 148. — Spinocalanua abyasalis: sample 11 (Table 3); scale D. Figure 149. — Spinocalanua longicornia: scale D. Figure 150. — Spinocalanua magnua: sample 41; scale D. Figure 151. — Spinocalanua antarcticua: sample 9; scale D. Figure 152. — Spinocalanua antarcticua: scale D. Figure 153. — Spinocalanua horridua: scale D. Figure 154. — Spinocalanua aimilia: sample 30; scale D. Figure 155. — Spinocalanua elongatua: scale D. Figure 156. — Spinocalanua polaria: scale D. Figure 157. — Spinocalanua brevicaudatua: sample 16; scale D. Figure 158. — Spinocalanua oligoapinoaua: sample 47; scale D. Figure 159. — Spinocalanua terranovae: sample 6; scale D. Figure 160. — Mimocalanua ovalia: sample 48; scale D. Figure 161. — Mimocalanua craaaua: scale D. Figure 162. — Mimocalanua aulcifrona: scale D. Figure 163.— Mimocalanua heronae: sample 31; scale D. 63 segments of swimming legs with rows of spines (see below). Caudal rami and caudal setae symmetrical. Female. — Prosome in dorsal view slender to robust ovoid, head rounded or with prominent crest. Ros- trum 2 sharp, vertically directed, spines; sometimes asymmetrical. Th5 lateral corners prolonged; Thl-Th4 without lateral spinules. Prosome length 3- 3V2 times urosome. Genital segment sometimes asymmetrical; protruding ventrally one-half depth of rest of urosome. Caudal rami length lVfe-2 times width. Al segment 2 length at least IV2 times segment 1; segments 8 and 9 fused. A2 Re length exceeding Ri by about 20%. Published information on A2 segmentation contradictory; probably Re2 with 2 setae and at least partly fused with Re3. Mn B2 with 4 inner setae. Mxl gnathobase as in Spinocalanus; distal anterior seta apparently absent. Inner lobe-1 with 5 setae. Outer lobe-2 small, with 1 short seta. Mx2 length 1 V2 times width; proximal outer hump nude. Lobe-5 with 4 strong setae; posterior surface of base without spines. Lobe-6 small, with 2 setae. Ri segments not reduced. Mx2 outer seta apparently ab- sent. Mxp Bl with or, apparently, without transverse spine-comb midlength on outer anterior and posterior edge. B2 without spine-comb. Bl with proximal and midlength group of 3 setae each; distal group ap- parently with 4-6 setae. B2 with longitudinal row of stiff hairs; 2 setae about midlength, 1 seta more distal, and 2 distal setae. Ri4-5 outer setae relatively short and thin, probably lightly plumose; Ri5 inner edge without spinules. PI Re3 apparently without posterior surface spines. P2-P3 Re2 posterior surface probably with row of small spines; P4 Re2 with 2 rows of spines. P2-P3 Re3 posterior surface probably with 2 distal rows of spines. P4 Re3 posterior surface with longitudinal series of small spines. P2-P4 Re terminal seta with finely serrate outer edge; outer flange narrow. P2-P4 Ri2 posterior surface with 2 rows of spines. P3-P4 Ri3 posterior surface with 2 rows of spines. P4 Bl with transverse row of long spinules on inner posterior surface; length variable, sometimes asymmetrical or lacking on right. Male. — Adult known only for M. typica. See also Monacilla sp. (= M. typica. — Wheeler, 1970). Key to the Species of Monacilla 1. Female: Prosome slender ovoid, anterior crest. (Male unknown.) 2. M. tenera Female: Prosome robust ovoid, no anterior crest (2) 2. Female: Genital segment symmetrical, not pro- truding on right; P4 Bl inner posterior surface with transverse row of spinules, equal length left and right. (Male unknown.) 7. M. gracilis Female: Genital segment asymmetrical, pro- truding on right; P4 Bl inner posterior surface with transverse row of spinules, longer on left, sometimes absent on right. Male: Length 1.59-2.3 mm. PI Rel outer spine reduced to mi- nute seta; PI Re2 outer spine forming sharply curved hook; P5 left Ri styliform 1. M. typica 1. Monacilla typica Sars, 1905 Monacilla typica Sars, 1905, p. 9. —Sars, 1924, 1925, p. 38-40, pi. 11, fig. 1-15; pi. 12, fig. 1-10. —Rose, 1933, p. 87-88, fig. 49. — Vervoort, 1946, p. 158-163. —Wilson, 1950, p. 267, pi. 26, fig. 380. — Farran and Vervoort, 1951b, p. 2-3, fig. 3b-e, h-i. — Tanaka, 1956, p. 396-397, fig. 18. — Owre and Foyo, 1967, p. 42, fig. 88, 92, 226-227, 229-230. — Grice, 1971, p. 276-279, fig. 2J-K, 3A-B. Oxycalanus spinifer Farran, 1908, p. 25-27, pi. 1, fig. 11-17. Oxycalanus semispinus Scott, 1909, p. 33-34, pi. 2, fig. 9-21. Monacilla dubia Scott, 1909, p. 35-36, pi. 3, fig. 17-29. Monacilla semispina. — Wilson, 1950, p. 266-267, pi. 26, fig. 379. Monacilla typica var. asymmetrica Tanaka, 1956, p. 398, fig. 19. Type Specimens: Syntypes, females (2.30 mm), Musee Oceanographique de Monaco (Belloc, 1960); on slides F5052-F5058, OSLO. Type Locality: East and northeast Atlantic, 1,000- 1,200 m. Material Studied: 1 male (1.75 mm), sample 39 (Table 3); 1 male (1.72 mm), sample 40. Description Female. — Length 1.95-2.5 mm. Prosome in dorsal view robust ovoid; head rounded, without crest. In lateral view with fairly abrupt forward slope. Rostrum 2 sharp, vertically directed spines, symmetrical. Prosome length 3V2 times urosome. Genital segment asymmetrical, somewhat protruding on right. Caudal rami length IV2 times width. Al reaching anal segment. Mxp Bl with transverse spine-comb midlength on outer edge. B2 with cluster of spinules about one-third length on outer edge. P4 Bl transverse row of long spinules on inner posterior surface more developed on left, sometimes lacking on right. [Sars (1905, 1925) reported P5 as reduced to simple 64 spine or absent. No other worker has mentioned female P5, and most have stated P5 absent. | Male. — Length 1.59-2.3 mm. Prosome ovoid, anterior irregular in dorsal view, posterior somewhat rounded; in lateral view with abrupt forward slope. Rostrum knoblike protuberance, faintly divided. Ce and Thl separate; Ce slightly expanded dorsally but not posteriorly. Prosome length 2Vi times urosome. Anal segment reduced. Caudal rami length slightly greater than width. Caudal setae relatively thick. Al reaching urosome segment 3. Segment 2 somewhat longer than segment 1; segments 9 and 10 fused. Right segment 20 merging with segment 21, as on left-handed species; segment 25 reduced. Mx2 fragile, surface nude; setae of reduced length, terminal setae thicker than on female. Mxp Bl-2 surface nude, setae of reduced size. Ril partly fused to Ri2. Ril with 4 short, plumose setae. Ri2 twice as long as on female, setae of reduced size. Ri3 proximal seta reduced; other 2 setae long, thick, with rough and shortly plumose inner edge. Ri4 inner setae as Ri3, outer seta reduced. Ri5 reduced, all setae reduced. PI Rel outer distal spine reduced to minute seta. Re2 outer distal spine forming sharply curved hook. P4 Bl posterior surface without inner transverse row of spinules or setules; inner margin with seta. P5 biramus, left-handed; left Bl reaching one-third length of right B2; left B2 reaching one-third length of right Re2. Left leg slightly longer than right; left and right Re about equal length; left Ri much longer than right Ri. Order of length, longest to shortest, of Re segments: left 3, 2, 1; right 2 = 3, 1. Left Ri reaching about to end of left Re longest terminal seta. Right Ri reaching one-third length of right Re2. At least right Rel with distal outer seta. Inner edge of left Re2-3 with hairs. Each Re with 1 small and 1 moderate bladelike terminal setae. Remarks Sars (1905) originally reported female M. typica length as 2.20 mm; apparently this was an error, cor- rected to 2.30 mm (Sars, 1925; Belloc, 1960). Sars' (1924, 1925) later description is thorough; however, he undoubtedly failed to note spines on the posterior surface of P2-P4, described by other workers. The descriptions by Rose (1933), Farran and Ver- voort (1951b), and Owre and Foyo (1967) are from Sars (1924, 1925). Distribution Pacific Ocean Northwest: Tanaka (1953, 1956), 0-1,000 m. -Furuhashi (1961, 1966), 369-3,010 m. — Minoda (1971), 87-164 m. West: Wilson (1950), 0-180 m. Indo-Pacific: Scott (1909), 0-2,000 m. — Ver- voort (1946), 355-1,800 m. —Wilson (1950). 0-1,080 m. Indian Ocean Northwest: Vervoort (1946), 555-1,000 m. West: Grice and Hulsemann (1967), 1,000- 2,000 m. Southwest: De Decker and Mombeck (1965), 500-1,500 m. Atlantic Ocean North: Grice and Hulsemann (1965), 850-1,400 m. Northeast: Sars (1905; 1924, 1925), 0-2,500 m. —Farran (1908, 1926), 450-1,800 m. — Lysholm and Nordgaard (1945), 0-1,700 m. —Grice and Hulsemann (1965), 1,000-2,800 m. — Vives (1970), 300-900 m. East: Sars (1905; 1924, 1925), 0-5,000 m. —Ly- sholm and Nordgaard (1945), 0-3,400 m. — Vervoort (1963), 0-100 m. —Roe (1972a, b), 830-950 m. Mediterranean Sea: Sars (1924, 1925), cited by Massuti (1939), 0-2,595 m. — Hure (1965), 300-1,000 m. —Hure and Scotto di Carlo (1968, 1969, 1971), 300-1,000 m. — Mazza (1968), 250-700 m. —Scotto di Carlo (1968), below 300 m. —Grice (1971), 850-2,200 m. Southeast: Unteruberbacher (1964), 0-52 m. Central: Lysholm and Nordgaard (1945), 0- 1,250 m. West: Grice (1963), 600-1,000 m. Caribbean Sea: Owre and Foyo (1964), 584- 877 m. —Park (1970), 487-1,900 m. 2. Monacilla tenera Sars, 1907 Monacilla tenera Sars, 1907, p. 6. —Sars, 1924, 1925, p. 40-41, pi. 12, fig. 11-18. —Farran, 1926, p. 245- 246, pi. 6, fig. 13-14. —Rose, 1933, p. 88, fig. 50. — Farran and Vervoort, 1951b, p. 2-3, fig. 4a-b. — Owre and Foyo, 1967, p. 42, fig. 224-225, 228, 231- 232. Hypsicalanus gracilis Wolfenden, 1911, p. 219-220, pi. 25, fig. 6-11. TYPE Specimkns: Syntypes, females (2.30 mm), Mu- see Oceanographique de Monaco (Belloc, 19601: on slides F5059-F5061. OSLO. Type Locality: Central North Atlantic, 0-3.000 m. Description Female. — Length 1.8-2.30 mm. Prosome in doreal view slender ovoid; head with prominent crest. In lateral view, crest with abrupt forward slope. Rostrum 2 sharp, vertically directed spines, left longer than right (Sars, 1926; Sars. 1907, first reporting right longer than left). Prosome length 3 times urosome. Genital segment symmetrical. Caudal rami length 2 times width. 65 Al reaching anal segment. Mxp Bl apparently without transverse spine-comb. P4 Bl reported by Sars (1925) to lack transverse row of spinules on inner posterior surface; row of spinules reported by Wolfenden (1911). Male. — Adult unknown. [Owre and Foyo (1967) briefly described one male stage V (size not reported). Urosome was reported with 5 segments, which would indicate an adult specimen, but the illustrated P5 suggest stage V male. The crest and rostrum were as on adult female.] Remarks Sars (1907) originally reported female M. tenera length as 2.20 mm; apparently this was an error, cor- rected to 2.30 mm (Sars, 1925; Belloc, 1960). The descriptions by Rose (1933) and Farran and Vervoort (1951b) are from Sars (1924, 1925). Distribution Indian Ocean Northwest: Sewell (1929), 1,260 m. West: Grice and Hulsemann (1967), 1,000- 2,000 m. Southwest: De Decker and Mombeck (1965), 1,000-1,500 m. Atlantic Ocean Northeast: Farran (1926), 900-3,600 m. East: Grice and Hulsemann (1965), 1,000-2,800 m. Central: Sars (1907; 1924, 1925), 0-3,000 m. — Wolfenden (1911). —Wheeler (1970), 2,000- 4,000 m. Northwest: Wheeler (1970), 2,000-4,000 m. Caribbean Sea: Owre and Foyo (1964, 1967), 877-1,500 m. —Park (1970), 203-1,900 m. (3. Oxycalanu8 spinifer Farran, 1908) See Monacilla typica Sars, 1905. (4. Oxycalanus semispinus Scott, 1909) See Monacilla typica Sars, 1905. (5. Monacilla dubia Scott, 1909) See Monacilla typica Sars, 1905. (6. Hypsicalanus gracilis Wolfenden, 1911) See Monacilla tenera Sars, 1907. 7. Monacilla gracilis (Wolfenden, 1911) Oxycalanus gracilis Wolfenden, 1911, p. 221-222, fig. 10; pi. 25, fig. 12-14. Monacilla gracilis. — Tanaka, 1956, p. 398-399, fig. 20. Type Specimen: Unknown. Type Locality: Central Atlantic, 0-3,000 m. Description Female. — Length 1.8-2.25 mm. Similar to Mona- cilla typica, except genital segment symmetrical, and transverse row of long spinules on inner posterior sur- face of P4 Bl equal length left and right, reaching beyond distal edge of Bl. Male. — Unknown. Remarks M. gracilis had been considered a synonym of M. typica. Tanaka (1956) reported one specimen which he considered M. gracilis, a distinct species. Vervoort (1963) pointed out that Tanaka's specimen may be distinct, but that it can only be referred to Wolfenden's M. gracilis with some reservations. Distribution Pacific Ocean Northwest: Tanaka (1953, 1956), 0-1,000 m. Atlantic Ocean Central: Wolfenden (1911), 0-3,000 m. (8. Monacilla typica var. asymmetrica Tanaka, 1956) (M. typica var. asymmetrica Tanaka, 1953, nomen nudum) See Monacilla typica Sars, 1905. 9. Monacilla sp. Monacilla typica.— Wheeler, 1970, p. 9, fig. 21-22. Not M. typica Sars, 1905. Remarks Wheeler (1970) reported one male (1.01 mm) as M. typica. P5 were damaged, but apparently differed from M. typica P5. This specimen, as well as other small Monacilla males, should be reexamined. Ver- voort (1946) reported one male (also 1.01 mm) as M. typica; it may be the same species as Wheeler's specimen. No females can at present be assigned to this species. 66 Distribution Pacific Ocean Indo-Pacific: ?Vervoort (1946), 1,845-2,500 m. Atlantic Ocean West: Wheeler (1970), 2,000-4,000 m. Genus Mimocalanus Farran, 1908 TYPE SPECIES: Mimocalanus cultrifer Farran, 1908. Description Rostrum absent. Pi Rel without outer distal spine; Rel-2 together longer than Re3; Ri with 1-2 inner setae. P2 Ri2 with 0-1 outer seta. P3-P4 Ri3 with 0-1 outer seta. Surface of swimming leg segments without spines. Caudal rami symmetrical. Caudal setae usual- ly incomplete on specimens examined: no trace of out- er (sixth) seta; outer and inner terminal setae usually completely detached; middle 2 terminal setae usually broken at bases; inner seta directed ventrally, small or reduced. Female. — Prosome in dorsal view slender to robust ovoid. Most species with distinct indented sclerotiza- tion about midlength on Ce; sclerotization direct ex- tension of sclerotized supporting margin of labrum, and perhaps serving as strengthened attachment for mandibular muscles. (Sclerotization reduced in adult males, where mandible also reduced.) Th5 lateral cor- ners prolonged, at least to midgenital segment. Thl- Th4 without lateral spinules. Prosome length nearly 4 to nearly 5 times urosome. Caudal rami length 1-1 Vz times width. Al segment 2 length about equaling segment 1 (M. ovalis segment 2 length about IV2 times segment 1). Segments 8 and 9 partly fused (completely fused on M. ovalis). Al narrowing between segments 9-11. Al exceeding caudal rami by 1-4 segments (only reaching genital segment on M. ovalis). Armature as in Spinocalanus female (Table 4). Esthete bases smaller than on Spinocalanus female. A2 (Fig. 194) Re length exceeding Ri by 20%. Rel without setae. Re2 and Re3 separate; Re2 without setae. Mn B2 with 2 or 3 inner setae, surface nude. Ril surface nude. Mxl gnathobase proximal surface without spinules; distal anterior seta present. Inner lobe-1 with 4 setae; inner lobes- 1-2 relatively large. Proximal part of Ri with 3 or 7 proximal and 3 or 7 distal setae. Small nude outer lobe-2 sometimes present. Mx2 (Fig. 197) length 2 times width; without prox- imal outer hump. Lobe-5 with 3 strong and 1 reduced setae; posterior surface of base without spines. Lobe-6 reduced, apparently with only 1 seta. Ri segments reduced. Mx2 outer seta apparently absent. Mxp Bl-2 without transverse spine-comb. Bl with proximal group of 2 setae, midlength group of 3 setae, and distal group of 3 or 4 setae. B2 with longitudinal row of reduced spinules or denticles (M. ovalis with stiff hairs); 3 setae about midlength and 2 distal setae (M. ovalis with 2 setae midlength, 1 seta more distal, and 2 distal setae). Ri4-5 outer setae usually moderately long, with very long plumes each side; Rio inner edge without spinules. P2-P4 Re terminal seta with finely serrate outer edge (48-90 teeth); outer flange wide. P4 Bl transverse row of thin setules on posterior surface proximal to inner seta (M. crassus without transverse row of setules and without inner seta). Male. — Prosome in dorsal view elongate to robust ovoid; anterior varying with species. Reduced sclerotization midlength on Ce in adult; male stage V sclerotization as on adult female. Ce not expanded. Ce and Thl separate. Prosome length 2*4 -3' 4 times urosome. Anal segment not reduced, length 2 /j-1 times caudal ramus. Caudal rami length 1-1 Vi times width. Al segment 2 length about equaling segment 1; seg- ment 10 partly fused with segment 9; right segment 20 merging with segment 21, as on other left-handed species; segment 25 not reduced. Al narrowing between segments 8-10. Esthete bases very small. Ar- mature given in Table 4. Swimming legs generally similar to those on cor- responding female. P4 Bl without inner transverse row of setules on posterior surface; some species with inner seta. P5 uniramus, left-handed, variably asymmetrical; Re essentially 3-segmented, Rel-2 may be partly fused. Key to the Species of Mimocalanus 1. P2 Ri2 with 1 outer seta (4) P2 Ri2 with no outer seta 1 2 1 2. P2-P3 Bl without inner seta; P3-P4 Ri3 with no outer seta 8. M. crassus P2-P3 Bl with inner seta; P3-P4 Ri3 with 1 outer seta (3) 3. Female: Al not reaching beyond genital seg- ment; Al segment 2 length 1 ! .- times segment 1. Pi Ri outer lobe reduced. (Male unknown.) 6. M. ovalis Female: Al reaching beyond caudal rami; Al segments 1 and 2 lengths about equal. Pi Ri outer lobe not reduced. (Male unknown.) . 1. M. cultrifer 4. Length greater than 4.0 mm. (Male unknown.) 3. hi. major Length less than 4.0 mm (5) 5. P2-P4 Ri2 with broadly inflated inner edge. (Male unknown.) 4. M. inflatus P2-P4 Ri2 not inflated (6) 6. Female: length less than 2.0 mm; Th4 and Th5 partly fused; inner caudal seta small, not re- 67 duced; Mxp Ri2 length more than l'/2 times Ril. Male: Length less than 1.5 mm . . 10. M. heronae, new species Female: Length greater than 2.0 mm; Th4 and Th5 separate; inner caudal seta reduced; Mxp Ri2 length less than IV2 times Ril. Male: Length greater than 1.5 mm (7) 7. Female: Prosome robust ovoid, Th5 lateral cor- ners reaching midgenital segment; PI Re2 outer spine extending two-thirds length of Re3; P2-P4 Re3 spiniform distal process about equaling size of Re3 outer distal spine; P2-P4 Re terminal seta with less than 75 teeth. Male: Prosome robust ovoid; P5 left Rel longer than Re2 2. M. nudus Female: Prosome ovoid, Th5 lateral corners reaching beyond midgenital segment; PI Re2 outer spine reaching distal edge of Re3; P2-P4 Re3 spiniform distal process about one-half size of Re3 outer distal spine; P2-P4 Re ter- minal seta with more than 75 teeth. Male: Prosome elongate ovoid; P5 left Re2 longer than Rel 9. M. sulcifrons 1. Mimocalanu8 cultrifer Farran, 1908 (Figures 164-168) Mimocalanus cultrifer Farran, 1908, p. 23-24, pi. 1, fig. 5-9. —Rose, 1933, p. 86, fig. 47. —Davis, 1949, p. 24. — Farran and Vervoort, 1951b, p. 2-3, fig. la- b, d. —Vervoort, 1957, p. 42-44, fig. 11-15. — ?Grice, 1971, p. 278-279, fig. 3C-D. Type Specimen: Holotype, female, BM 1908.7.6.4. TYPE LOCALITY: Northeast Atlantic, 720-1,800 m. Material Studied: 1 female (1.57 mm), sample 37 (Table 3). Description Pi Ri with 2 inner setae; P2 Ri2 with no outer seta; P3-P4 Ri3 with 1 outer seta. Female.— Length 1.0-1.70 mm(?) (Figs. 164, 165). Prosome in dorsal view fairly robust ovoid; head rounded, with slight bulge. In lateral view with fairly Mimocalanus cultrifer, female Figure 164. — Habitus, dorsal view; sample 37; scale A. Figure 165.— Habitus, lateral view; sample 37; scale A. Figure 166.— PI; sample 37; scale D. Figure 167.— P2; sample 37; scale D. Figure 168.— P4; sample 37; scale D. 168 68 abrupt forward slope. Th4 and Th5 separate. Prosome length 4 times urosome. Genital segment slightly wider than long, protruding ventrally one-third depth of rest of urosome. Caudal rami slightly longer than wide. Inner caudal seta reduced. Al exceeding caudal rami by 3-4 segments. Specimen examined lacked segments 24-25; remainder as Table 4. Mn B2 with 2 inner setae (Farran, 1908). Mxl gnathobase apparently without proximal posterior setae. PI (Fig. 166) B2 and Rel outer distal edge with 3-4 small teeth. P2 (Fig. 167)-P4 damaged on specimen examined; described by Vervoort (1957); P4 (Fig. 168) Bl with inner transverse row of thin setules. Male. — Undescribed. Grice and Hulsemann (1965; unpublished station list, 1966) reported a male of M. cultrifer (1.2 mm). Roe (1972b) reported 1 male (1.14 mm). Remarks Rose's (1933) and Farran and Vervoort's (1951b) descriptions are from Farran (1908); Farran and Ver- voort (1951b) included length data (up to 1.91 mm) from Vervoort (1946) which probably include lengths of M. heronae or another species. The above descrip- tion limits the length of female M. cultrifer to the most probable records. Vervoort (1957) did not in- clude his earlier (1946) record in the description of M. cultrifer. Possibly records of small M. cultrifer (1.0- 1.17 mm) by Vervoort (1946) and Grice and Hulsemann (1965; unpublished station list, 1966) are of M. crassus. Grice's (1971, fig. 3C) lateral view of M. cultrifer is not as robust as other descriptions of this species (see M. heronae). Grice did not indicate the lengths of his specimens. Distribution Minoda's (1971) specimens smaller than 1.70 mm are included in the summary below; the larger specimens are considered M. heronae. Pacific Ocean North: Minoda (1971), 743-1,230 m. Northeast: Davis (1949), 1,100-1,900 m. — Flem- inger (1967), 0-140 m. —Present study, 1,300- 1,500 m. Northwest: Furuhashi (1966), 252-512 m. —Mi- noda (1971), 195-485 m. Southwest: Vervoort (1957), 500-750 m. Indo-Pacific: Vervoort (1946), 340-1,000 m. Indian Ocean West: Grice and Hulsemann (1967), 750-4.000 m. Atlantic Ocean Northeast: Farran (1908, 1926), 360-1,800 m. — (irice and Hulsemann (1965), 250-4,750 m. East: Roe (1972a, b), 400-800 m. Mediterranean Sea: ?Grice (1971), 1,190-1,350 m. Northwest: Grice (1963), 620-1,200 m. —Wheel- er (1970), 2,000-4,000 m. West: Wheeler (1970), 2,000-4,000 m. Caribbean Sea: Park (1970), 208-1,900 m. Antarctic Pacific Sector: Vervoort (1957, 1965), 250-500 m. Indian Sector: Vervoort (1957, 1965), .500-1. (XX) m. 2. Mimocalanu8 nudus Farran, 1908 (Figures 169-180) Mimocalanus nudus Farran, 1908, p. 24-25, pi. 1, fig. 1-4. —Farran and Vervoort, 1951b, p. 2-3, fig. 2b-c. e-g. Mimocalanus distinctocephalus Brodsky, 1950, p. 136-137, fig. 56. Type Specimens: Syntypes, 2 females (2.30, 2.64 mm), BM 1908.7.6.5. TYPE LOCALITY: Northeast Atlantic, 720-1,440 m. Material Studied: Syntype, female (2.30 mm), sample 4 (Table 3); 1 female (2.68 mm), sample 35; 2 females (2.57, 2.65 mm) and 1 male (2.50 mm), sam- ple 36; 4 females (2.50-2.70 mm, mean = 2.61 mm), sample 37. Description Pi Ri with 2 inner setae; P2 Ri2 with 1 outer seta; P3-P4 Ri3 with 1 outer seta. Female.— Length 2.28-2.70 mm (Figs. 169, 170). Prosome in dorsal view robust ovoid; head somewhat truncate. In lateral view with fairly abrupt forward slope. Th4 and Th5 separate. Prosome length 4 times urosome. Genital segment slightly wider than long, protruding ventrally one-third depth of rest of urosome. Caudal rami length 1 ' 3 times width. Inner caudal seta reduced. Al terminal segments missing on all described specimens. Segments 20-25 missing on specimens ex- amined; armature of remaining segments as Table 4. Mn B2 apparently with 2 inner setae. Blade (Fig. 171) with large, sharp, slightly curved ventral tooth followed by 2 large and 1 slender tricuspids, a shorter bicuspid. 3 slender teeth, and a dorsal seta. Mxl gnathobase with 1 proximal posterior seta, nearly as large as smallest strong marginal setae; proximal anterior seta also relatively large: distal 69 174 175 Mimocalanua nudus Figure 169. — Female, habitus, dorsal view; sample 36 (Table 3); scale A. Figure 170. — Female, habitus, lateral view; sample 36; scale A. Figure 171. — Female, Mn blade; sample 35; scale D. Figure 172. — Male, habitus, dorsal view; sample 36; scale A. Figure 173. — Male, habitus, lateral view; sample 36; scale A. Figure 174. — Male, Mn blade; sample 36; scale H. Figure 175.— Male, P5; sample 36; scale D. 70 Mimocalanus nudu*, female Figure 176.— PI; sample 4 (Table I); stale 1) Figure 177. — PI Re2-.1; sample IS; scale C. Figure 178. — P2; sample 4; scale I). Figure 179.— P.I; sample 4; scale 1). Figure ISO. — P4; sample 1; scale 1). 71 anterior seta reaching beyond margin. Proximal part of Ri with 3 proximal and 3 distal setae; Ri terminal segment with 7 setae. Mxp Bl anterior surface with midlength cluster of many thin spinules and distal series of short thin setules; distal knob with 3 setae and short truncate process. B2 proximal anterior surface with short longitudinal row of short spinules of decreasing size distally. Ri2-5 with nude inner and terminal setae. Ri4-5 outer setae moderately long, with long plumes on each side. PI (Figs. 176, 177) B2 and Rel outer distal edge with few small teeth. Main body of Re2 outer spine not reaching beyond two-thirds length of Re3; some spines with a threadlike terminal part. Re3 outer spine relatively wide. Ri outer lobe large, rhomboi- dal. P2 (Fig. 178) Rel outer spine about two-thirds size of Re3 outer spines. Re3 spiniform distal edge nearly equaling size of outer distal spine. P3 (Fig. 179) B2 outer distal edge with 2-3 sharp teeth. Rel outer spine smaller than Re2-3 outer spines. Re3 spiniform distal edge as large as outer dis- tal spine. P4 (Fig. 180) Bl posterior surface with inner transverse row of very thin setules. B2 distal outer edge with dull tooth. Re outer spines relatively small; Rel outer spine smallest. Re3 outer spines not serrate. Re3 spiniform distal edge as large as outer distal spine. Re terminal seta with following number of teeth: P2 (52-65); P3 (56-67); P4 (ca. 61). Male.— Length 2.50 mm (Figs. 172, 173). Prosome in dorsal view robust ovoid; anterior irregular, with slight forward protrusion. In lateral view with irregular, fairly abrupt forward slope. Prosome length 3V2 times urosome. Caudal rami length lVfe times width. Al exceeding caudal rami by 2 segments. VIII- 2s, le; XI-ls, le; left segments 17-25 missing; remainder as Table 4. Mn blade (Fig. 174) relatively complex; 5 strong ventral teeth, 3 dorsal setiform processes. Mxl gnathobase reduced to 2-3 short, narrow teeth; cluster of spinules on distal anterior surface. Mxp Bl distal lobe with 3 unequal setae. B2 with 2 small followed by 1 large midlength setae. Ri4-5 outer setae reduced or absent. PI Re2 outer spine length as on female; Re3 outer spine narrower than on female. P2 Rel outer spine as on female. P5 (Fig. 175) reaching middle of urosome segment 3. Left Bl reaching just beyond right Bl; left B2 reaching one-third length of right Rel. Left leg longer than right by more than length of Re3. Left Re longer than right. Order of length, longest to shortest, of Re segments: left 1, 2, 3; right 1 = 2, 3. Right Re3 small. Each Rel with flat seta on outer distal edge, left 2 times length of right. Rel-2 nearly fused left and right. Inner edge of left and right Re2-3 with long hairs, those of left Re3 in tufts. Each Re with short bladelike terminal process. Remarks Farran (1908) reported only one female in the original description; however, at least one other specimen was taken in that survey and labeled Mimocalanus nudus by Farran; this second specimen was examined during the present study (sample 4, Table 3). Farran separated M. nudus from M. cultrifer essentially by size, but failed to note es- pecially the differences in armature of the swimming legs. Farran and Vervoort (1951b) illustrated P2 Ri2 with 1 outer seta. Brodsky (1950) first reported the distinctive sclerotization on Ce and used this characteristic to define a North Pacific species, M. distinctocephalus. In the present study, this sclerotization was found to be more or less developed on all Mimocalanus species. Brodsky referred to this feature as a "band of pigment"; however, it appeared in the present study not to be pigmented. Brodsky (fig. 56, Nl) illustrated PI Ri with 3 inner setae; this is considered in the pres- ent study as an error, or an abnormality. Brodsky's (fig. 56, N2) illustration of P2 Ri2 shows 1 outer seta. No differences were found between North Pacific "M. distinctocephalus" and Farran's M. nudus, and the two forms are considered conspecific. Distribution Pacific Ocean North: Brodsky (1950, 1957), 200-4,000 m. — Mi- noda (1971), 170-2,000 m. Northeast: von Vaupel-Klein (1970), 0-1,200 m. —Present study, 700-1,500 m. Northwest: Brodsky (1950, 1952a, 1955, 1957), 0-8,500 m. Indian Ocean West: Grice and Hulsemann (1967), 1,000-2,000 m. Atlantic Ocean Northeast: Farran (1908), 1,440 m. —Present study, 720-900 m. East: Roe (1972a, b), 700-960 m. Caribbean Sea: Park (1970), 980-2,800 m. 3. Mimocalanus major Sars, 1920 Mimocalanus major Sars, 1920, p. 4. — Sars, 1924, 1925, p. 35-36, pi. 10, fig. 1-16. Type Specimen: Holotype, female (4.20 mm), on slide, OSLO. Type LOCALITY: Central North Atlantic, 0-3,000 m. 72 Description Pi Ri with 2 inner setae; P2 Ri2 with 1 outer seta; P3-P4 Ri3 with 1 outer seta. Female. — Length of only known specimen 4. 20 mm. Prosome in dorsal view slender ovoid; head somewhat truncate. In lateral view with abrupt forward slope. Th4 and Th5 separate. Prosome length nearly 5 times urosome. Caudal rami about as long as wide. Labrum with 3 large protuberances on midventral surface. Al segment 1 with 2 strong setae. Male. — Unknown. Remarks Sars' (1924, 1925) description is thorough, but should be confirmed with additional specimens. Sars states that Ce and Thl are fused. He did not describe an inner transverse row of setules on P4 Bl, present on most Mimocalanus species. The 2 strong setae on Al segment 1 are unusual, and perhaps there is a third strong seta. Distribution Known only from the original record, Central North Atlantic, 0-3,000 m. 4. Mimocalanus inflatus Davis, 1949 Mimocalanus inflatus Davis, 1949, p. 24. — Grice and Hulsemann, 1965, p. 227, fig. 5o-w. — ?Grice and Hulsemann, 1967, p. 20, fig. 16-18. TYPE SPECIMEN: Unknown. TYPE LOCALITY: Northeast Pacific, 1,900-2,900 m. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta; P3-P4 Ri3 with 1 outer seta. Female.— Length (1.76?-) 2.3-2.4 mm. Known for only two specimens (Davis, 1949; Grice and Hulsemann, 1965). Male. — Unknown. Remarks Grice and Hulsemann (1967) reported two small specimens (1.76-2.01 mm) but the swimming feet were lacking, and they questioned this identification, which could be of M. ovalis or other species. Davis (1949) did not illustrate this species, and it is not certain whether his specimen had P1-P4 armature as described above (taken from Grice and Hulsemann, 1965). The only characteristic which set the two specimens apart from other Mimocalanus species was the inflated P2-P4 Ri2. Distribution Pacific Ocean Northeast: Davis (1949), 1,900-2,900 m. Indian Ocean West: ?Grice and Hulsemann (1967), 1.0O0- 2,000 m. Atlantic Ocean East: Grice and Hulsemann (1965), 1,900-3,000 m. (5. Mimocalanus distinctocephalus Brodsky, 1950) See Mimocalanus nudus Farran, 1908. 6. Mimocalanus ovalis (Grice and Hulsemann, 1965), new combination (Figure 160) Spinocalanus ovalis Grice and Hulsemann, 1965, p. 229, fig. 7b-f. ^Spinocalanus ventriosus Grice and Hulsemann. 1967, p. 21-22, fig. 32-34. TYPE SPECIMEN: Holotype, female, BM 1965.4.20.3. TYPE LOCALITY: Northeast Atlantic, 1,900-5,000 m. Material Studied: 2 paratypes, female (1.32 mm), sample 48 (Table 3). Also the holotype of Spinocalanus ventriosus, female (damaged), sample 43. Description PI Ri with 2 inner setae; P2 Ri2 with no outer seta; P3-P4 Ri3 with 1 outer seta. Female. — Length 1.16-1.52 mm. Prosome in dorsal view robust ovoid; head broadly rounded. In lateral view with abrupt forward slope. Th4 and Tho separate; Th5 lateral corners reaching midgenital seg- ment. Prosome length 3 1 ! times urosome. Genital seg- ment slightly longer than wide, protruding ventrally one-half depth of rest of urosome. Caudal rami length about equaling width. Al reaching only to midgenital segment; segment 2 length about 1 ' 2 times segment 1; segments 8-9 com- pletely fused; terminal segments (Fig. 160) relatively wide, short. Segments 8 through 24 with 1-2 rows ol minute denticles on ventral and dorsal surface, not noticed on any other Spinocalanidae. Segment 1 damaged; armature of remaining segments as Table 4. Mn B2 with 3 inner setae (Grice and Hulsemann. 1965. fig. 7c). 73 Mxp Bl with row of thin setules on midanterior sur- face, and 4 clusters of thin spinules on outer posterior surface; distal lobe anterior surface covered with setules; 1 relatively long and 1 small distal setae, and 1 long truncate distal process. B2 with longitudinal row of stiff hairs; 2 midlength setae, followed by more distal seta and 2 distal setae. Ri segments rhom- boidal; inner subterminal setae short (some probably missing from specimens examined), but terminal seta thick, not expanded, probably very long (broken). Ri4-5 outer setae relatively thick and long; each side with long plumes. PI B2 with small outer seta and long inner anterior seta, as in other Spinocalanidae. B2 inner edge without hairs; Rel nude. Ri outer lobe reduced to small lamella. P2-P3 Bl inner edge with hairs and 1 seta; outer anterior surface with few setules. Specimens exam- ined lacked P2-P4 Re2-3. Male. — Unknown. Remarks Mimocalanus ovalis has some characteristics of Spinocalanus species: relative lengths of Al segments 1-2 and complete fusion of segments 8-9, and arma- ment and spacing of setae on Mxp B2. However, the lack of outer distal spine on PI Rel, lack of surface spines on swimming legs, number of setae on Ri of swimming legs, 3 inner setae on Mn B2, form of Mxp Ri4-5 outer setae, and prolonged Th5 would indicate placement with Mimocalanus species. Specimens with complete Re of swimming legs should be exam- ined. Grice and Hulsemann (1967) briefly described females of a new species Spinocalanus uentriosus. Ex- cept for a slight knoblike protrusion at the tip of Th5, their description is consistent with that of Mimocalanus ovalis. The knobiike protrusion could be an artifact, caused by a contraction of the body walls at that point, as it appeared to be in the holotype examined in the present study. The holotype was lacking Al distal segments, P1-P4 Re2-3, P2 Ri2, and P3-P4 Ri2-3. PI Rel, PI Ri, P2-P3 Bl, Al segments 1-2 and 8-9, and Mxp B2 as in M. ovalis. There was no indication of an outer spine on PI Rel; usually a socket, or at least a torn thread, can be seen if the spine is detached. The holotype lacked the swimming leg segments that would have surface spines in Spinocalanus, so that placement of S. ven- triosus with Mimocalanus ovalis may still be questionable. Distribution Indian Ocean West: Grice 4,000 m. and Hulsemann (1967), 1,000- Atlantic Ocean Northeast: Grice and Hulsemann (1965), 1,900- 5,000 m. (7. Mimocalanus species Grice and Hulsemann, 1967) See Mimocalanus crassus Park, 1970. 8. Mimocalanus crassus Park, 1970 (Figures 161, 181-191) Mimocalanus species Grice and Hulsemann, 1967, p. 21, fig. 19-25. Mimocalanus crassus Park, 1970, p. 478-481, fig. 1-12. Type Specimen: Holotype, female (0.95 mm), USNM 123771. TYPE LOCALITY: Caribbean Sea, 487-2,800 m. Material Studied: Holotype, female (0.95 mm), sample 55 (Table 3); 1 female (1.08 mm), sample A5 (Table 1); 1 female (1.0 mm), sample A6; 1 female (1.20 mm), sample A23; 2 males (1.27, 1.36 mm), sam- ple A29; 3 females (1.16, 1.20, 1.24 mm) and 1 male (1.24 mm), sample A31; 3 males (1.08 mm), sample A32; 1 female (1.20 mm), sample A33; 1 female (1.20 mm), sample A36; 2 females (1.16, 1.24 mm), sample A45; 3 females (1.20, 1.26, 1.27 mm) and 1 male (1.22 mm), sample A48; 23 females (1.08-1.25 mm, mean = 1.12 mm) and 1 male (1.04 mm), sample A51. Description Right PI Ri with 2 inner setae; left PI Ri with 1 inner seta, distal. P2 Ri2 with no outer seta. P3-P4 Ri3 with no outer seta. P2-P4 Bl without inner seta. Female.— Length 0.95-1.27 mm (Figs. 181, 182). Prosome in dorsal view robust ovoid; head rounded. In lateral view with abrupt forward slope. Th4 and Th5 separate. Prosome length 4Va times urosome. Genital segment length three -fourths times width, protruding ventrally one-fourth depth of rest of urosome. Caudal rami length slightly greater than width. Mimocalanus crassus Figure 181. — Female, habitus, dorsal view; scale B. Figure 182. — Female, habitus, lateral view; scale B. Figure 183.— Female, Mn blade; scale D. Figure 184.— Female, PI; scale D. Figure 185.— Female, P2; scale D. Figure 186.— Female, P3; scale D. Figure 187.— Female, P4; scale D. Figure 188. — Male, habitus, dorsal view; scale B. Figure 189. — Male, habitus, lateral view; scale B. Figure 190.— Male, Mn blade; scale D. Figure 191.— Male, P5; scale D. 74 190 7;S Al exceeding caudal rami by about 2 segments; ter- minal segments (Fig. 161) of moderate length and width. Armature as Table 4. A2 Re2-3 not fused, otherwise as shown by Park (1970, fig. 3): Mn Bl with 3 inner setae. Mn blade (Fig. 183) relatively large; large rounded ventral tooth, 1 broad irregular midlength tooth followed by rounded bicuspid, and dorsal projection with 2 sharp, narrow teeth, 1 tricuspid, and dorsal seta. Mxl as shown by Park (1970, fig. 5); gnathobase with 4 proximal posterior setae and short spinelike distal anterior seta. B2 with 7 inner setae. Proximal part of Ri with 7 proximal and 7 distal setae; Ri ter- minal segment with 9 setae. Mxp as shown by Park (1970, fig. 7); Bl with spinules midlength on inner edge; distal lobe with short spinules, 3 setae, and short truncate process. B2 with short longitudinal row of spinules on proximal in- ner edge. Ri4-5 outer setae lightly plumose, thin and relatively short. PI (Fig. 184) Re2 outer distal spine reaching beyond base of Re3 outer distal spine. PI Ri with moderate lobe; setae asymmetrical as described above. P2 (Fig. 185) Rel outer distal spine smaller than outer spines of Re2-3. P3 (Fig. 186) B2 outer distal edge with 2-3 small teeth. P4 (Fig. 187) Bl inner margin with hairs, without inner seta and without transverse row of setules. Re outer spines smaller than on P2-P3. Re terminal seta with following number of teeth: P2 (55-63); P3 (ca. 55); P4 (ca. 48). distal edge, left wider than right. Right Rel-2 nearly fused. Inner edge of left B2 and Re3 with long hairs. Each Re with moderate bladelike terminal process, longer on right. Remarks Park (1970) indicated M. crassus PI Ri with "3 or 4 setae"; his illustration (fig. 8) shows right PI with 3 setae, as does the illustration of male right PI by Grice and Hulsemann (1967, fig. 24). Park's illustra- tion (fig. 11) of male left PI shows 3 setae, in agree- ment with specimens examined in the present study. PI of holotype had been removed prior to deposit. Mimocalanus crassus is the only Spinocalanidae with asymmetrical PI. Mimocalanus crassus is also the only Spinocalanidae with no outer seta on P3-P4 Ri3; no inner seta on P2-P4 Bl; Mxl B2 with 7 inner setae, Ri proximal part with 7 and 7 setae, and Ri ter- minal segment with 9 setae. Mimocalanus crassus is the only Mimocalanus species with short, lightly plumose outer setae on Mxp Ri4-5, and without inner transverse row of setules on P4 Bl (see M. major). Distribution Arctic Ocean Canadian Basin: Present study, 400-2,500 m. Indian Ocean West: Grice and Hulsemann (1967), 2,000- 3,000 m. Atlantic Ocean Caribbean Sea: Park (1970), 487-2,800 m. Male.— Length 0.78-1.36 mm (Figs. 188, 189). Prosome in dorsal view robust ovoid; anterior irregular, head rounded. In lateral view with irregular, abrupt forward slope. Prosome length 2 3 /4 times urosome. Caudal rami length equaling width. Al reaching middle of urosome segment 2. Al segments 8-9 relatively elongate. VTII-2s, le; XI- 2s, le; remainder as Table 4. Mn blade (Fig. 190) with 6 small teeth, decreasing in size dorsally, and minute dorsal seta. Mxl gnathobase reduced to 1 or 2 teeth. PI as on female, Ri asymmetrical. P2 as on female, except Re3 outer spines sharply curved near tip. P3-P4 missing Re2-3 on specimens examined; remainder as on female, except B2 outer distal edge without teeth, and Ri2 outer spiniform process re- duced. P5 (Fig. 191) nearly reaching distal edge of urosome segment 3. Left Bl reaching one-fourth length of right B2; left B2 reaching one-half length of right Rel. Left and right legs about equal length. Right Re, including terminal blade, slightly longer than left. Order of length, longest to shortest, of Re segments: left 2, 1, 3; right 1 = 2 = 3. Each Rel with short flat seta on outer 9. Mimocalanus wlcifrons Wheeler, 1970 (Figures 162, 192-215) Mimocalanus nudus. — Grice and Hulsemann, 1965, p. 227, fig. 6a-l. Not M. nudus Farran, 1908. Mimocalanus sulcifrons Wheeler, 1970, p. 7-9, fig. 5-20. Mimocalanus distinctocephalus. — Vidal, 1971, p. 17- 18, fig. 60-63. Not M. distinctocephalus Brodsky, 1950. Type Specimen: Holotype, male (1.90 mm), USNM 122646. TYPE LOCALITY: Western North Atlantic, 2,000-4,000 m. Material Studied: Holotype, male (1.90 mm), sam- ple 45 (Table 3); 3 females (2.20, 2.30, 2.30 mm), sam- ple A26 (Table 1); 3 females (2.24, 2.28, 2.32 mm), sample A48; 4 females (2.05-2.40 mm, mean = 2.23 mm), 4 males (2.15-2.60 mm, mean = 2.37 mm), and 1 stage V male (2.08 mm), sample A50; 4 females (2.17-2.30 mm, mean = 2.24 mm) and 1 stage V male (2.04 mm), sample A51; 1 female (2.24 mm), sample A52. 76 192 Mimocalanua aulcifrons, female Figure 192. Figure 193. Figure 194. Figure 195. Figure 196. Figure 197. Figure 198. Figure 199. Figure 200. Figure 201. -Habitus, dorsal view; scale A. -Habitus, lateral view; scale A. -A2, terminal setae omitted; scale C -Mn blade; scale D. -Mxl gnathobase, left; scale D. -Mx2, lobe-1 setae omitted; scale D. -PI; scale C. -P2; scale C. -P3; scale C. -P4; scale C. 77 Mimocalanu8 sulcifrons Figure 202.— Female, Mxp; scale D. Figure 203.— Male, Mn blade; scale D. Figure 204. — Male, Mxp; scale D. Figure 205.— Male, PI; scale D. Figure 206.— Male, P2; scale C. Figure 207.— Male, P2 Re3, abnormal; scale C. Figure 208.— Male, P3; scale C. Figure 209.— Male, P4; scale C. Figure 210.— Male, P5; sample 45 (Table 3); scale D. Figure 211.— Male, P5; scale D. Figure 212. — Male stage V, habitus, lateral view; scale A Figure 213.— Male stage V, P5; scale D. 78 213 Mimocalanus sulcifrons, male Figure 214. Figure 215. -Habitus, lateral view; scale A. -Habitus, dorsal view; scale A. Description Pi Ri with 2 inner setae; P2 Ri2 with 1 outer seta; P3-P4 Ri3 with 1 outer seta. Female.— Length 2.05-2.40 mm (Figs. 192, 193). Prosome in dorsal view ovoid; head broadly rounded, with slight anterior protrusion and 2 indistinct dor- solateral swellings. In lateral view with abrupt forward slope. Th4 and Th5 separate. Prosome length 4 ! 2 times urosome. Genital segment length two-thirds times width, protruding ventrally one-third depth of rest of urosome. Caudal rami length 1 Vi times width. Inner caudal seta reduced. Al exceeding caudal rami by 4 segments; terminal segments (Fig. 162) relatively wide, large. Armature as Table 4. Mn B2 with 3 inner setae. Mn blade (Fig. 195) with 1 long ventral tooth, followed by 4 bi- or tricuspids of decreasing length, slender dorsal bicuspid, and dorsal seta. Mxl gnathobase (Fig. 196) without proximal posterior setae; proximal anterior seta thin and finely plumose; distal anterior seta on left only, reaching beyond margin. Proximal part of Ri with 8 proximal and 3 distal setae. Mxp (Fig. 202) Bl inner anterior surface with midlength cluster of few denticles and distal series <>l short thin setules; distal knob with 4 setae and short truncate process. B2 proximal anterior surface with short longitudinal row of short, relatively thick spines of irregular length, paralleled by very short, thin spinules. Ri2-5 inner setae and terminal seta, nude. Ri4-5 outer setae moderately long, with very long plumes on each side. PI (Fig. 198) B2 and Rel outer distal edge with 2-8 small teeth. Main body of Re2 outer spine just reaching base of Re3 outer spine; Re3 outer spine narrow. Ri outer lobe moderate. P2 (Fig. 199) B2 outer distal edge with 3-4 small blunt teeth. Rel outer spine equaling size of Re3 outer spines. Re3 spiniform distal edge much smaller than outer distal spine. P3 (Fig. 200) similar to P2. P4 (Fig. 201) Bl posterior surface with inner transverse row of very thin, uniform setules. Re3 outer spines finely serrate. Re terminal seta with following number of teeth: P2 (79-90); P3 (80-89); P4 (73-83). Male.— Length 1.90-2.60 mm (Figs. 214, 215). Prosome in dorsal view elongate ovoid; anterior narrow, with anterior depression lined with short hairs. Mid-Ce with dorsolateral oblique rows of flat, prominent, fragile spines. Prosome in lateral view with irregular, fairly gentle, forward slope. Prosome length 3 :, /4 times urosome. Caudal rami length l'/« times width. Al exceeding caudal rami by 1 segment. VIII- 2s, 2e; XI-2s, le; remainder as Table 4. Mn blade (Fig. 203) reduced to 6-7 subequal teeth. Mxl gnathobase with 6-7 toothlike setae. Inner lobes-1-2 reduced (as Wheeler, 1970, fig. 14); outer lobe-1 with very long, fragile, plumose setae, 2 of which reach urosome; Re as on female. Mx2 overall appearance as illustrated by Wheeler (1970, fig. 15); lobe-1 with 3-4 reduced setae, lobe-2 with 3 longer setae; lobe-3-4 with 3 still longer setae; lobe-5 with 2 long setae. Lobe-6 and Ril-2 with 1 coarsely plumose seta, as long as Mx2; Ri3 with 3 long setae. Mxp (Fig. 204) Bl distal lobe with dense patch of setules; B2 with dense longitudinal row of stiff hairs; several setae apparently missing on specimens ex- amined. Ri 1-5 inner setae and terminal seta plumose; outer setae reduced or absent. PI (Fig. 205) Re2 outer spine reaching about midlength Re3; Re2-3 outer spines finely serrate. P2 (Fig. 206) Rel outer spine nearly as long as Re.'l outer spines. Re terminal seta with 69-76 teeth. P3 (Fig. 208)-P4 (Fig. 209) missing Re2-3 on specimens examined; P4 Bl inner seta reaching mid- dle of B2. P5 (Figs. 210, 211) reaching distal edge of urosome 79 segment 3. Left Bl reaching one-fifth to one-fourth length of right B2; left B2 reaching one-half length of right Rel. Left leg longer than right by length of Re3. Left Re longer than right. Order of length, longest to shortest, of Re segments: left 2, 1, 3; right 1 = 2 = 3. Right Re3 not small. Each Rel probably with short flat seta on outer distal edge. Right Rel-2 more or less fused. Inner edge of left Re3 with long hairs. Each Re with short bladelike terminal process. Male stage V.— Length 2.04-2.08 mm (Fig. 212). Ce sclerotization about as on adult female. P5 (Fig. 213) uniramus, symmetrical. Re incompletely 2- segmented; Rel with outer distal seta; Re2 with short bladelike terminal process. Remarks Mimocalanus nudus and M. sulcifrons females are similar, while the males are more distinct. Both species are found in the North Atlantic, but M. sul- cifrons has not been found in the Pacific, nor M. nudus in the Arctic. The present study considers the larger male Arctic Mimocalanus species to be con- specific with M. sulcifrons; therefore, the larger Arctic female is most likely also M. sulcifrons, described for the first time. The large North Pacific female Mimocalanus species is considered identical to M. nudus; the corresponding Pacific male is, therefore, most likely to be male M. nudus, described for the first time. Wheeler (1970) found one male specimen on which he based the description of M. sulcifrons. His descrip- tion of the "horn-shaped esthetes" was of the esthete bases only. Th4 and Th5 of the holotype are separate, as on other male Spinocalanidae. Wheeler stated that PI Ri had "1 subterminal, and 2 terminal setae," however his illustration (fig. 17) correctly shows a total of 4 setae. The anterior depression was much more pronounced on the holotype, and the holotype did not possess the conspicuous dorsolateral spines of the Arctic males. However, the other characteristics of the holotype of M. sulcifrons were consistent with those of the Arctic males. Perhaps the "sulcus" and the dorsolateral spines are characteristics of age of the adult male (see Spinocalanus dorsispinosus Brodsky, 1950). Grice and Hulsemann (1965) reported one male M. nudus from the northeast Atlantic. Their description is generally consistent with that of male M. sulcifrons. The illustration of dorsal view (fig. 6a) clearly shows a depressed anterior, while male M. nudus has a protruding anterior margin, and is more robust. Distribution Johnson (1963a), Minoda (1967), Hughes (1968), and Dunbar and Harding (1968) reported M. distinc- tocephalus from the Arctic; these records will be con- sidered as M. sulcifrons in the present study. Arctic Ocean Eurasian Basin: Minoda (1967), 0-520 m. Canadian Basin: Johnson (1963a), 0-2,000 m. —Hughes (1968), 80-90 m. —Dunbar and Harding (1968), 0-2,000 m. — Vidal (1971). — Present study, 100-2,500 m. Atlantic Ocean Northeast: Grice and Hulsemann (1965), 2,000- 4,000 m. West: Wheeler (1970), 2,000-4,000 m. 10. Mimocalanus heronae, new species (Figures 163, 216-223) Mimocalanus cultrifer. — Tanaka, 1956, p. 387-389, fig. 13. Not M. cultrifer Farran, 1908. Mimocalanus distinctocephalus. — Boucher and de Bovee, 1970, p. 527-534, fig. 1-2. Not M. distincto- cephalus Brodsky, 1950. Type Specimen: Holotype, female (1.23 mm), USNM 142702. Type Locality: Central and Northeast Pacific, 0- 1,000 m. Material Studied: Holotype, female (1.23 mm), sample 31 (Table 3); 2 paratypes, females (1.25, 1.27 mm), sample 32, SIO XVm-779; 1 paratype, female (1.28 mm), sample 32, USNM 142703. Description PI Ri with 2 inner setae; P2 Ri2 with 1 outer seta; P3-P4 Ri3 with 1 outer seta. Female.— Length 1.23-1.95 mm (Figs. 216, 217). Prosome in dorsal view slender ovoid; head rounded. In lateral view with irregular, fairly abrupt forward slope. Th4 and Th5 partly fused. Prosome length nearly 4 times urosome. Genital segment slightly wider than long, protruding ventrally one-third depth of rest of urosome. Caudal rami length slightly greater than width. Inner caudal seta small, not reduced. Al exceeding caudal rami by nearly 1 segment; ter- minal segments (Fig. 163) relatively wide. Segment 9 partly fused with segment 10. Armature as Table 4. Mn B2 with 2 short inner setae; proximal seta thick, nude, distal seta plumose. Blade (Fig. 218) with large ventral bicuspid, followed by 4 bi- or tricuspids of decreasing length, 2 narrow bicuspids, 1 thin sharp process, and dorsal seta. Mxl gnathobase without proximal posterior setae; left and right with short distal anterior seta. Proximal part of Ri with 3 proximal and 3 distal setae. Mxp (Fig. 219) Bl distal knob with long anterior surface hairs, 3 subequal setae, and short truncate process. B2 anterior surface with proximal row of short hairs and row of denticles. Ri2-5 inner setae nude; Ri4-5 outer setae moderately long with long plumes each side; terminal seta plumose. 80 217 223 Teneriforma naso, male Figure 224.— P5; sample 42; scale D. 224 Mitnocalanus heronae, female Figure 216.— Habitus, lateral view; sample 31 (Table 3); scale B. Figure 217. — Habitus, dorsal view; sample 31; scale B. Figure 218.— Mn blade; sample 31; scale G. Figure 219. — Mxp; sample 31; scale D. Figure 220.— PI; sample 31; scale D. Figure 221.— P2; sample 31; scale D. Figure 222.— P3; sample 31; scale D. Figure 223.— P4; sample 31; scale D. 225 I - Figure 225.— Scales A-H, 0.1 mm. A I C D E F G H 81 PI (Fig. 220) Rel outer distal edge with 1-3 small teeth. Re2 outer spine not reaching beyond two-thirds length of Re3. Re3 outer spine narrow. Ri outer lobe moderate, with small terminal point. P2-P4 (Figs. 221-223) variously incomplete on specimens examined in this study. P4 Bl with inner transverse row of thin, uniform setules. Male. — Length of only known specimen 1.28 mm (Tanaka, 1956). According to Tanaka: right P5 with 4 segments, but probably Rel-2 more or less fused; Re3 relatively large (Tanaka, fig. 13i); left P5 exceeding right by Re3 and one-half length of Re2. Remarks Tanaka (1956) described two females and one male M. cultrifer; his illustration (fig. 13f) of P2 Ri2 suggests an outer seta, not possessed by M. cultrifer. In other respects, also, his description is consistent with that of M. heronae. Boucher and de Bovee (1970) reported the first Mimocalanus species from the Mediterranean Sea, four females which were identified as M. distinc- tocephalus. Their description, including figures of complete P2-P4, is consistent with that of M. heronae, except they stated that Th4 and Th5 were separate, a characteristic that may be variable, as in some Spinocalanus species. They also stated that P2 Ri2 and P3-P4 Ri3 have 2 outer setae; their figure of P2 suggests 1 outer seta, and their figures of P3-P4 in- dicate 1 outer seta, consistent with M. heronae. Their Table 3 indicates 1 outer seta on P2 B2, which should instead be indicated for PI B2. Perhaps the Mediterranean specimens reported by Grice (1971) as M. cultrifer are also M. heronae (see M. cultrifer). Distribution Minoda (1971) reported two females (1.55, 1.95 mm) said to be equivalent to Tanaka's species. These are also considered M. heronae in the present study. Vervoort (1946) reported M. cultrifer females up to 1.91 mm, from the Indo-Pacific; it is possible that the specimens longer than 1.70 mm are M. heronae. Pacific Ocean North: Minoda (1971), 743-1,230 m. Northeast: Present study, 300 m. Central: Present study, 0-1,000 m. Northwest: Tanaka (1953, 1956), 300-500 m. —Minoda (1971), 195-485 m. Indo-Pacific: ?Vervoort (1946), 636-900 m. Atlantic Ocean Mediterranean Sea: Boucher and de Bovee (1970), 50-500 m. Etymology: I am pleased to name this species for Gayle A. Heron, Department of Oceanography, University of Washington, Seattle, in gratitude for her unfailing friendship over nearly 20 years and in recognition of her enthusiasm and skill in the study of the Copepoda. Genus Teneriforma Grice and hulsemann.1967 Tanyrhinus Farran, 1936. not Tanyrhinus Mannerheim, 1852. TYPE SPECIES: Teneriforma naso (Farran, 1936). Description Rostrum a single long blunt cone. PI Rel without outer distal spine; Rel-2 together only slightly longer than Re3. PI Ri with 2 inner setae. P2 Ri2 with 1 outer seta. P3-P4 Ri3 with 2 outer setae. Surface of swim- ming leg segments without spines. Caudal rami symmetrical, length 2-2 V% times width. Female. — Prosome in dorsal view slender ovoid; head rounded. In lateral view with' very abrupt forward slope. Th5 lateral corners prolonged, reaching one-third length of genital segment. Thl-Th4 without lateral spinules. Th4 and Th5 separate. Prosome length 3-3 l A times urosome. Genital segment protruding ventrally one -fourth to one -third depth of rest of urosome. Al reaching anal segment; segment 2 length about equaling segment 1; segments 8 and 9 fused. A2 Re length about equaling Ri. Rel with 1 seta; Re2 and Re3 fused; Re2 with 2 setae. Mn B2 with 3 inner setae. Mxp Bl-2 without transverse spine-comb; B2 ap- parently with only 2 midlength setae, and without longitudinal row of stiff hairs or other armament. Published illustrations of Ril-3 indicating less setae than typical for the family. Ri4-5 outer setae moderately long, plumosity unknown. P2-P4 Re terminal seta with moderately coarse serrate outer edges; outer flange narrow. P4 Bl posterior surface apparently without inner transverse row of hairs or setules. Inner seta present. Male. — Prosome in dorsal view elongate ovoid; head rounded. In lateral view with very abrupt forward slope. Th5 lateral corners prolonged, as in other male Spinocalanidae; nearly reaching urosome segment 2. Ce not expanded. Ce and Thl separate. Prosome length 3 times urosome. Anal segment not reduced, length two-thirds times caudal ramus. Al reaching to end of caudal rami; segment 2 length about equaling segment 1; segments 9 and 10 separate; segments beyond 10 not fused; right Al miss- ing from specimen examined; left segments 20 and 21 distinct, as in left-handed species; segment 25 not reduced. P5 (Fig. 224) uniramus, left-handed, very asymmetrical. Right B2 reduced to small knob; no 82 right Re. Left Bl reaching midlength of right B2. Left Re 3-segmented; Rel with short outer distal seta; Re3 inner edge with long hairs; short terminal seta. Order of length, longest to shortest, of left Re segments: 2, 1, 3. Only a single species is known, Teneriforma naso. Teneriforma naso (Farran, 1936) (Figure 224) Tanyrhinus naso Farran, 1936, p. 86-87, fig. 4. — Grice and Hulsemann, 1965, p. 231, fig. 8. Teneriforma naso. — Grice and Hulsemann, 1967, p. 22, fig. 36-38. —Wheeler, 1970, p. 9-10, fig. 27-28. TYPE SPECIMEN: Unknown. TYPE LOCALITY: Southwest Pacific, 0-600 m. Material Studied: 1 male (0.92 mm), sample 42 (Table 3). Description Female.— Length 0.89-1.20 mm. Male. — Length of only known specimen 0.92 mm. P5 (Fig. 224) redrawn from specimen of Grice and Hulsemann (1967), sample 42 (Table 3). Remarks Only about a dozen specimens of Teneriforma naso are known. Farran's (1936) only specimen had ap- parently lost the outer spine of PI Re2, which is pres- ent on this species. He also failed to note the small outer seta of P2 Ri2 and the 2 small outer setae of P4 Ri3. Grice and Hulsemann (1965) reported two females; lengths 0.9 and 1.1 mm (1966, unpublished station list). Distribution Pacific Ocean Southwest: Farran (1936), 0-600 m. Indian Ocean West: Grice and Hulsemann (1967), 750-3,000 m. Atlantic Ocean Northeast: Grice and Hulsemann (1965), 180- 3,000 m. West: Wheeler (1970), 2,000-4,000 m. Caribbean Sea: Park (1970), 505-1,900 m. SUMMARY Most copepods of the family Spinocalanidae are known to be widely distributed, and they often com- prise a large proportion, or even a majority, of the copepods in deep samples. However, because these copepods are fragile and therefore difficult to study, published records and descriptions of them have generally been vague or misleading. As a basis for a review of the Spinocalanidae, zooplankton samples were collected from Fletcher's Ice Island in the Canadian Basin of the central Arctic, in 1967-68. Both sexes of seven species of spinocalanids were identified and redescribed: Spinocalanus longicornis, S. antarcticus, S. horridus. S. elongatus, S. polaris, Mimocalanus crassus, and M. sulcifrons. The depth of greatest concentration varied for each species, but was always below 100 m. Two species were taken in relatively shallow water (80-100 m) and four only below 300 m, but all were found at least as deep as 2,500 m. Spinocalanus horridus and Mimocalanus crassus are probably cosmopolitan. The other five Arctic spinocalanids, in spite of great vertical ranges, have more or less restricted geographic distributions, and one species, S. elongatus, appears to be endemic to the Arctic. These restricted distributions may result from interactions with closely related species with similar vertical distributions. Spinocalanus longicor- nis, S. antarcticus, and 5. elongatus appear to be replaced geographically by the closely related species S. abyssalis, S. magnus, and S. brevicaudatus, respectively. Of these species-pairs only S. longicornis and S. abyssalis are known to have substantial overlapping distributions. The distributions of all spinocalanids in the Arctic/North Atlantic transition area need further study. All published records and descriptions of spinocalanids are reviewed. Type specimens of the following species have been examined and redescribed: Spinocalanus abruptus, S. abyssalis, S. aspinosus, S. brevicaudatus, S. hoplites, S. longicor- nis, S. neospinosus, S. oligospinosus, S. ovalis, S. parabyssalis, S. pteronus, S. usitatus, S. ventriosus, Mimocalanus crassus, M. nudus, and M. sulcifrons. Other critical specimens are redescribed, including four species identified by Farran as Spinocalanus abyssalis var. pygmaeus, S. magnus (two species), and S. spinosus. Many named species or forms are placed in synonymy, and two new species, Spinocalanus terranovae and Mimocalanus heronae are described. Spinocalanus ovalis is transferred to the genus Mimocalanus; Spinocalanus longipes is shown to be the male of S. angusticeps; males of S. abyssalis and 6". polaris are considered to be incorrectly iden- tified in the literature; and the male of Mimocalanus nudus and the female of M. sulcifrons are described for the first time. The 32 species of Spinocalanidae which are con- sidered valid are distributed as follows: Spinocalanus (19 species, in 3 groups containing 14, 4, and 1 species, respectively). Monacilla (4 species), Mimocalanus [B species), and Teneriforma (1 species). Species o\ spinocalanids are most easily identified by certain 83 characters of the swimming legs. Since these legs are missing from most specimens, characters of the swim- ming legs have been correlated with other, relatively stable, characters, especially the armature of Mxp and body form and size. Characters used in the keys to genera and species have been selected so that usually even damaged specimens without swimming legs can be identified. Males of most Spinocalanus species are left- handed, but males of S. polaris and S. similis are right-handed; the handedness is reflected not only in P5 but also in Al. Known males in other genera of Spinocalanidae are left-handed. Within the Spinocalanidae, Spinocalanus and Monacilla appear to form one subgroup, while Mimocalanus and Teneriforma comprise a second subgroup. However, most appendages of Monacilla and Teneriforma species are still undescribed. Males are not yet known for most of a complex of species related to Spinocalanus spinosus: S. aspinosus, S. hoplites, S. oligospinosus, S. spinosus, and S. usitatus. Undoubtedly, knowledge of these males will help clarify the relationships of the females, which are very similar to one another. Males are also unknown for S. abruptus and S. hir- tus; Monacilla tenera and M. gracilis; Mimocalanus cultrifer, M. inflatus, M. major, and M. ovalis. The female of Monacilla sp. is unknown. Males of S. terranovae, Monacilla sp., and Mimocalanus heronae are each known only from one briefly described specimen. More specimens of S. abruptus, S. hirtus, Monacilla gracilis, Mimocalanus inflatus, M. major, and M. ovalis should be examined to confirm or clarify the status of these species. ACKNOWLEDGMENTS T. Saunders English, Department of Oceanography, University of Washington, provided the opportunity, through the Office of Naval Research (Contract N00014-67-A-0103-0005 Project NR 083 012), for me to participate in his initial field work on Fletcher's Ice Island in 1966, and he also arranged for me to receive the plankton samples from 1967-68. In connection with her own research with T. Saunders English, Gayle A. Heron often noticed rare specimens of Spinocalanidae and without exception furnished them for this study. She also helped to for- mulate the problems in the systematics of this group, and remained a valuable source of ideas throughout the investigation. Thomas E. Bowman, Smithsonian Institution, provided much necessary and valuable assistance during the research. I wish to thank him and Fenner A. Chace, Jr., for critical reviews of the manuscript. I owe a principal and collective debt to the several persons and institutions who graciously and generous- ly furnished valuable specimens for study: G. M. Bennell, British Museum of Natural History (BM); Janet M. Bradford, New Zealand Oceanographic Institute (NZOI); Marit E. Christiansen, Zoological Museum, University of Oslo (OSLO); A. Fleminger, Scripps Institution of Oceanography (SIO); H. A. Fehlmann, Smithsonian Oceanographic Sorting Center (SOSC); Julio Vidal, formerly with the Arctic Program, Office of Naval Research, University of Southern California (USC); the United States National Museum (USNM); the University of Washington (UW); George D. Grice, Woods Hole Oceanographic Institution (WHOI); and Tai Soo Park, Texas A&M University Marine Laboratory. I also wish to acknowledge the assistance of K. Moore in the final preparation of the figures. This report was adapted from a dissertation sub- mitted to The George Washington University in par- tial satisfaction of the requirements for the degree of Doctor of Philosophy. LITERATURE CITED BELLOC, G. 1960. Catalogue des types de Copepodes du Musee oceano- graphique de Monaco. Bull. Inst. Oceanogr. (Monaco) 1176:1-24. BERNSTEIN, T. 1934. 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Rebman Ltd., Lond., p. 25-44. 1911. Die marinen Copepoden der Deutschen Siidpolar-Ex- pedition 1901-1903. II. Die pelagischen Copepoden der West- winddrift und des Siidlichen Eismeers, mit Beschreibung mehrerer neuer Arten aus dem Atlantischen Ozean. Dtsch. Sudpolar-Exped. 1901-1903, Zool. 12:181-380. 88 ft U.S. GOVERNMENT PRINTING OFFICE: 1975-698-698 /I76 REGION 10 .1?" Collecting and processing data on fish eggs and larvae in the California Current region By David Kramer, Mary J Kalin, Elizabeth G Steven*, James K Thrailkill, and Jtmn K Zweifcl November 1972, iv + 38 pp., 38 Bgi,, 2 tables Fur nale by the Buperintendenl "I Document*, U 8 Government Printing Office, Washington, DC. 171 Ocean fishery management: Discussions and research By Adam A Bokolotki (editor) 1 17 paper*, 24 authors.) 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