OCS Study MMS 95-0047 Harpacticoida (Crustacea: Copepoda) from the California Continental Shelf Final Report TI-m*LIBRARY OF THE UNIV!!.RSITY OF TEXAS AT AUSTIN U.S. Department of the Interior ··~Minerals Management Service ID~Pacific OCS Region OCS Study MMS 95-0047 Harpacticoida (Crustacea: Copepoda) from the California Continental Shelf Final Report Edited by: Dr. Paul Montagna Submitted by: Marine Science Institute .. ,.. The University of Texas at Austin Port Aransas, TX 78373-1267 THI:*LIBRARY For: OF THE UNIVERSITY OF TEXAS U.S. Department of the Interior AT Minerals Management Service AUSTIN Pacific OCS Region 770 Paseo Camarillo Camarillo, CA 93010-6064 Under Purchase Order No. 13244 Minerals Management Service Pacific OCS Region October 1995 DISCLAIMER This report has been reviewed by the Pacific Outer Continental Shelf Region, Minerals Management Service, U.S. Department of the Interior and approved for publication. The opinions, findings, conclusions or recommendations expressed in this report are those ofthe authors, and do not necessarily reflect the views ofthe Minerals Management Service. Mention oftrade names or commercial products does not constitute endorsement or recommendation for use. This report has not been edited for conformity with Minerals Management Service editorial standards. TAXONOMIC DISCLAIMER This report does not constitute a peer-reviewed publication for the purposes of taxonomic literature. The species described herein do not constitute recognized taxonomic status or new taxa as stipulated in the International Code ofZoological Nomenclature, Article 8b . .""'filililiiiiiili~ll 3000955019 0 5917 3000955019 PROJECT ORGANIZATION PROGRAM MANAGER Dr. Paul Montagna University ofTexas at Austin Marine Science Institute P.O. Box 1267 Port Aransas, Texas 78373 PROJECT TECHNICAL OFFICER Mr. Frank J. Manago U.S. Department of the Interior Minerals Management Service, Pacific OCS Region 770 Paseo Camarillo Camarillo, California 93010-6064 CONTRIBUTING AUTHORS Mr. Robert Burgess University ofTexas at Austin Marine Science Institute P.O. Box 1267 Port Aransas, Texas 78373 Dr. Frank Fiers Koninklijk Belgisch lnstituut voor Natuurwetenschappen Vautierstraat 29 B-1040 Brussels BELGIUM ACCESS NUMBER: 13244 TECHNICAL SUMMARY STUDY TITLE: Harpacticoida (Crustacea; Copepoda) from the California Continental Shelf. REPORT TITLE: Harpacticoida (Crustacea; Copepoda) from the California Continental Shelf. CONTRACT NUMBER(S): MMS Purchase Order: 13244, MMS:l4-12-0001-13244. SPONSORING OCS REGION: Pacific. APPLICABLE PLANNING AREA(S): Southern California. FISCAL YEAR(S) OF PROJECT FUNDING: 1990, 1991, 1992, 1993, 1994, 1995 COMPLETION DATE OF REPORT: September 1995. COST(S): FY 1990: $141; FY 1991: $0; FY 1992: $0; FY1993; $744; FY1994; $0; FY1995: 2385; CUMULATIVE PROJECT COST: $24,470. PROJECT MANAGER(S): Paul Montagna AFFILIATION: University ofTexas at Austin, Marine Science Institute ADDRESS: P.O. Box 1267, Port Aransas, Texas, 78373 PRINCIPAL INVESTIGATOR(S)*: Paul Montagna, Robert Burgess, Frank Fiers KEY WORDS: Central California, Santa Maria Basin, infauna, benthos, meiofauna, Harpacticoida BACKGROUND: The California Harpacticoida Study was a small multi-year, marine program that was conducted to provide the U.S. Department of the Interior, Minerals Management Service with information necessary to assess the potential environmental impacts of leasing tracts of submarine lands for commercial oil production and to properly manage marine resources if recovery of petroleum occurs. The study resulted from data produced in the meiofauna studies of the California Monitoring Program (CAMP) from the Outer Continental Shelf (OCS). Harpacticoid copepods were found to be very numerous and important, but most were species new to science. The report consists of a series of detailed chapters describing the harpacticoids which were found in the CAMP project. The chapters are authored by two participating investigators, who are trained systematists. OBJECTIVES: The objective of the Report is to provide a detailed description of the new harpacticoids found in the Santa Maria Basin. The Report summarizes systematic accounts that are or will be published in peer-reviewed journals. DESCRIPTION: All specimens were derived from samples archived from the CAMP project. Harpacticoids were dissected, and drawings were made of each anatomical feature of the species that has taxonomic significance. Systematic accounts of the species were constructed based on a ACCESS NUMBER: 13244 comparison of other species found in the taxonomic literature. SIGNIFICANT CONCLUSIONS: Over 115 new species were found in the Santa Maria Basin during the CAMP program. Complete systematic accounts are given for 6 of the species, and drawings are provided for 9 others. A pictorial key is provided for the identification ofthe harpacticoids. STUDY RESULTS: The California OCS has a rich taxonomic diversity ofharpacticoid copepods. The harpacticoid community is an important link with higher trophic levels, and is particularly sensitive to pollution. Now that the harpacticoid community is more well known, they can be incorporated into the design of future benthic monitoring programs in the OCS. STUDY PRODUCT(S): Montagna, P. A. 1995. Harpacticoida (Crustacea; Copepoda) from the California Continental Shelf. A final report by the University of Texas, Marine Science Institute for the U.S. Department ofthe Interior, Bureau of Land Management Pacific OCS Office, Camarillo, CA. UTMSI Technical Report No. TR/95-0002. Contract No. 14-12-0001-13244. 135 pp. Fiers, F. 1991. Three new harpacticoid copepods from the Santa Maria Basin off the California Pacific coast (Copepoda, Harpacticoida). Beaufortia, 42:13-47. 1HE MARINE SCIENCE LIBRARY Merine Science Institute 1M University of Texas at Austin P. 0. Box 1267 Port Aranua, Texas 78373-1287 *P.I. 's affiliation may be different than listed for Program Manager(s). ' 0 ' 0 1------­ 1~ I I I I I I ~ " '\. ' '\. ' ' ' '\. '\. CALIFORNIA "\,. ' 0 50 100 STATUTE MILES I I I I I 0 50 100 190 200 IULOllElUIS ·. .... KEY MAft ,..o NORTHERN CALIFORNIA !MIA MJKr iJt,eni1aM 'r*.. ·vrnu errr .0 ~'1 200011! .aeana 4 tto'I \ Central California Planning Area for the Harpacticoida study. 50272-HU REPORT DOCUMENTATION , 1. 1tŁP"ORT NO. ,~ PAGE MMS 95-0047 4. Title and Subtitle October 1995 Continental Shelf Harpacticoida (Crustacea:Copepoda) from the California 7. Author(s) Paul A. Montagna (editor), Robert Burgess, Franky Fiers 9. Performlns Ors•nizatlon N•me •nd Address 10. Proiect/THk/Wor11 Unit No. The University of Texas at Austin _______________.. Marine Science Institute 11. ContractCC) or Grant(G) No. 750 Channelview Drive ~> Purchase Order Port Aransas, TX 78373 (G> No. 13244 12. Sponsorins Orsanization Name •nd Addreu 13. Type of Report & Period Covered Minerals Management Service, U.S. Department of Interior Final ReportPacific OCS Region, 770 Paseo Camarillo 14. Camarillo, CA 93010 9/90 to 10/95 15. Supplementary NotH 16. Abstract (Umit: 200 words) Specimens of new Harpacticoida species were obtained during the California Outer Continental Shelf (OCS), Phase II, Monitoring Program (CAMP) between November 1986 and May 1989. The CAMP project was a multidisciplinary study to detect and evaluate the long-term biological impacts of continental shelf oil drilling and production. The study was centered around a proposed platform site named Julius, which was never put into service. Samples were collected in the Santa Maria Basin on a regional scale (10­20 km). Harpacticoids are the second most abundant meiofaunal taxa in the Santa Maria Basin. Harpacticoids have been intensively studied in the Atlantic OCS. However, Pacific studies are limited to collections made in shallow water. There are a great number of undescribed species in the CAMP samples taken from the Santa Maria Basin. The present study is rather limited in scope and only touches on some of the dominate species found. It contains full taxonomic descriptions of six species, a pictorial key of 18 dominant species, and drawings of 42 other unknown species. 17. Document Analysis a . Descriptors Central California, Santa Maria Basin, infauna, benthos, meiofauna, Harpacticoida b. ldentff"teB/Open-Ended Term• Taxonomic Atlas, Keys to species, Crustacea, Harpacticoida c. COSATI Fleld/G:oup 19. Security CIHs (This Report) 21. No. of Pa•HIL Av•llablllty Statement Unlimited 1-------------t-------·-­ 20. Security Clua (Thia P••e) Z2. Price 336 UnC"h,c;ifiprf OP"TIONAL FORM 272 (4-77) (SH ANSl-Zl9.ll) SH lnatructlona on R.v•r•• (formerly HTIS-l5) 0.p•rtment of Commerce TABLE OF CONTENTS ACKNOWLEDGMENT ....................................................... xv CHAPTER 1: A NEW SPECIES OF CERVIN/A (COPEPODA: HARPACTICOIDA) FROM THE CALIFORNIA CONTINENTAL SHELF (by Robert Burgess) ............. 1 Abstract ...................................................2 Introduction ................................................ 3 Materials and Methods ........................................ 3 Taxonomic Account .......................................... 3 Discussion ................................................ 1O Key ..................................................... 11 Acknowledgments .......................................... 13 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 CHAPTER 2: A NEW SPECIES OF TETRAGON/CfPS (COPEPODA: HARPACTICOIDA) FROM THE CALIFORNIA CONTINENTAL SHELF(by Robert Burgess) ....... 22 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 3 Introduction ............................................... 24 Materials and Methods ....................................... 24 Taxonomic Account ......................................... 24 Discussion ................................................ 27 Acknowledgments .......................................... 29 References ................................................ 35 CHAPTER 3: A NEW GENUS AND NEW SPECIES OF CLETODIDAE (COPEPODA: HARPACTICOIDA) FROM THE CALIFORNIA CONTINENTAL SHELF (by Robert Burgess) ................................................. 36 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 7 Introduction ............................................... 38 Materials and Methods ....................................... 38 Taxonomic Account ......................................... 38 Discussion ................................................. 42 Acknowledgments .......................................... 45 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2 CHAPTER 4: THREE NEW HARPACTICOID COPEPODS FROM THE SANTA MARIA BASIN OFF THE CALIFORNIAN PACIFIC COAST (COPEPODA, HARPACTICIODA) (by Frank Fiers) ............................... 53 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Introduction ............................................... 55 Materials and Methods ....................................... 55 viii Systematic Descriptions ....................................... 55 Acknowledgements .......................................... 72 References . ~ ............................................. 100 CHAPTER 5: PICTORIAL KEY OF DOMINANT HARPACTICOID SPECIES (by Robert Burgess) ................................................ 102 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128 APPENDIX: DRAWINGS OF UNDESCRIBED SPECIES (by Frank Fiers) .......... 130 ix LIST OF FIGURES Figure 1.1.: Cervinia mediocauda n.sp. S? and o. . ......................... 15 Figure 1.2.: Cervinia mediocauda n.sp. S? mouth parts. . ..................... 15 Figure 1.3.: Cervinia mediocauda n.sp. ~ legs............................. 15 Figure 1.4.: Cervinia mediocauda n.sp.o mouth parts. . ..................... 15 Figure 1.5.: Cervinia mediocauda n.sp.o legs............................. 15 Figure 2.1.: Tetragonicips pacifica n. sp. S? •••••••••••••••••••••••••••••• 31 Figure 2.2.: Tetragonicips pacifica n. sp. S? •••••••••••••••••••••••••••••• 31 Figure 2.3.: Tetragonicips pacifica n. sp. S? •••••••••••••••••••••••••••••• 31 Figure 3.1.: Serratithorus geei ~. ..................................... 47 Figure 3.2.: Serratithorus geei ~ mouth parts. . ........................... 47 Figure 3.3.: Serratithorus geei S? periopods............................... 47 Figure 3.4.: Serratithorus geei o...................................... 47 Figure 4.1.: Zosime pacifica n. sp. female. . ............................. 79 Figure 4.2.: Zosime pacifica n. sp. female. . ............................. 79 Figure 4.3.: Zosime pacifica n. sp., female. . ............................. 79 Figure 4.4.: Zosime pacifica n. sp., male. . .............................. 79 Figure 4.5.: Zosime pacifica n. sp., copepodid I. .......................... 79 Figure 4.6.: Zosime pacifica n. sp., copepodid II. .......................... 79 Figure 4.7.: Zosime pacifica n. sp., copepodid Ill. ......................... 79 Figure 4.8.: Zosime pacifica n. sp., copepodid Ill. ......................... 79 Figure 4. 9.: Zosime pacifica n. sp. . .................................... 79 Figure 4.10.: Cletodes macrura n. sp. . ................................. 79 Figure 4.11.: Cletodes macrura n. sp. . ................................. 79 Figure 4.12.: Cletodes macrura n. sp., copepodid I and II legs................. 79 Figure 4.13.: Cletodes macrura n. sp., copepodid I and II. ................... 79 Figure 4.14.: Cletodes macrura n. sp., copepodid Ill and IV ................... 79 Figure 4.15.: Cletodes macrura n. sp., copepodid IV and V. . ................. 79 Figure 4.16.:Cletodes tuberculatus n. sp................................. 79 Figure 4.17.: Cletodes tuberculatus n. sp., mouthparts and o and S? abdomens. . ... 79 Figure 4.18.: Cletodes tuberculatus n. sp., legs. . .......................... 79 Figure 4.19.: Cletodes tuberculatus n. sp., copepodid I and II. ................. 79 Figure 4.20.: Cletodes tuberculatus n. sp., copepodid 111, IV and V.............. 79 Figure 6.1. Ceratonotus pectinatus. . ................................. 131 Figure 6.2. Ceratonotus n. sp. 1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 Figure 6.3. Ceratonotus n. sp. 2. . ................................... 135 Figure 6.4. Ceratonotus n. sp. 2. . ................................... 137 Figure 6.5. Amphiascoides n. sp. . .................................. 139 Figure 6.6. Amphiascoides n. sp. . .................................. 141 Figure 6.7. Amphiascoides n. sp..................................... 143 Figure 6.8. Amphiascoides n. sp. . ................................... 145 Figure 6.9. Species C n. sp......................................... 147 Figure 6.10. Species C n. sp........................................ 149 x Figure 6.11. Species C n. sp........................................ 151 Figure 6.12. Species PADA n. sp..................................... 153 Figure 6.13. Species PADA n. sp..................................... 155 Figure 6.14. Species PADA n. sp..................................... 157 Figure 6.15. Species PADA n. sp..................................... 159 Figure 6.16. Species DANA n. sp. . .................................. 161 Figure 6.17. Species DANA n. sp. . .................................. 163 Figure 6.18. Species DANA n. sp. . .................................. 165 Figure 6.19. Species DANA n. sp. . .................................. 167 Figure 6.20. Species PADB n. sp..................................... 169 Figure 6.21. Species PADB n. sp..................................... 171 Figure 6.22. Species PADB n. sp..................................... 173 Figure 6.23. Species PSLO n. sp. . ................................... 175 Figure 6.24. Species PSLO n. sp. . ................................... 177 Figure 6.25. Species STED n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 Figure 6.26. Species STED n. sp. . ................................... 181 Figure 6.27. Species STED n. sp..................................... 183 Figure 6.28. Species STED n. sp. . ................................... 185 Figure 6.29. Species MM n. sp. . .................................... 187 Figure 6.30. Species MM n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189 Figure 6.31. Species MM n. sp...................................... 191 Figure 6.32. Species STEC n. sp. . ................................... 193 Figure 6.33. Species STEB n. sp. . .................................... 195 Figure 6.34. Species STEB n. sp. . ................................... 197 Figure 6.35. Species STEB n. sp. . ................................... 199 Figure 6.36. Species J n. sp. . ...................................... 201 Figure 6.37. Species HEST n. sp. . ................................... 203 Figure 6.38. Species HEST n. sp. . ................................... 205 Figure 6.39. Species HEST n. sp. . ................................... 207 Figure 6.40. Species HEST n. sp. . ................................... 209 Figure 6.41. Species NOCO n. sp. . .................................. 211 Figure 6.42. Species CLET L n. sp. . .................................. 213 Figure 6.43. Species CLET L n. sp. . .................................. 215 Figure 6.44. Species CLET L n. sp. . .................................. 217 Figure 6.45. Species CLM N. sp. . ................................... 219 Figure 6.46. Species CLM n. sp...................................... 221 Figure 6.47. Species CLET MILL n. sp. . ............................... 223 Figure 6.48. Species CLET MILL n. sp. . ............................... 225 Figure 6.49. Species CLET D n. sp. . ................................. 227 Figure 6.50. Species I n. sp. . ...................................... 229 Figure 6.51. Species I n. sp. . ...................................... 231 Figure 6.52. Species I n. sp. . ...................................... 233 Figure 6.53. Species I n. sp. . ...................................... 235 Figure 6.54. Species VV n. sp....................................... 237 xi Figure 6.55. Species VV n. sp....................................... 239 Figure 6.56. Species VV n. sp....................................... 241 Figure 6.57. Species Z n. sp........................................ 243 Figure 6.58. Species Z n. sp. . ...................................... 245 Figure 6.59. Species EURY n. sp. . .................................. 247 Figure 6.60. Species EURY n. sp..................................... 249 Figure 6.61. Species CLET A n. sp.................................... 251 Figure 6.62. Species CLETA n. sp.................................... 253 Figure 6.63. Species CLET A n. sp.................................... 255 Figure 6.64. Species CLET R n. sp.................................... 257 Figure 6.65. Species CLET P n. sp.................................... 259 Figure 6.66. Species STLO n. sp. . ................................... 261 Figure 6.67. Species STLO n. sp. . ................................... 263 Figure 6.68. Species STVE n. sp. . ................................... 265 Figure 6.69. Species CERD n. sp..................................... 267 Figure 6.70. Species CERD n. sp..................................... 269 Figure 6.71. Species CERD n. sp..................................... 271 Figure 6.72. Species CERD n. sp..................................... 273 Figure 6.73. Species CERC n. sp..................................... 275 Figure 6.74. Species CERC n. sp..................................... 277 Figure 6.75. Species CERC n. sp..................................... 279 Figure 6.76. Species CERC n. sp. . ................................... 281 Figure 6.77. Species MINO n. sp. . .................................. 283 Figure 6.78. Species TISBE n. sp. . ................................... 285 Figure 6.79. Species TISBE n. sp. . ................................... 287 Figure 6.80. Species TISBE n. sp. . ................................... 289 Figure 6.81. Species TISBE n. sp..................................... 291 Figure 6.82. Species ZOS B n. sp. . .................................. 293 Figure 6.83. Species ZOS B n. sp. . .................................. 295 Figure 6.84. Species UU n. sp. . .................................... 297 Figure 6.85. Species UU n. sp. . .................................... 299 Figure 6.86. Species UU n. sp. . .................................... 301 Figure 6.87. Species UU n. sp. . .................................... 303 Figure 6.88. Species IDYA n. sp. . ................................... 305 Figure 6.89. Species IDYA n. sp. . ................................... 307 Figure 6. 90. Species IDYA n. sp. . ................................... 309 Figure 6.91. Species IDYA n. sp..................................... 311 Figure 6.92. Species TAPA n. sp..................................... 313 Figure 6.93. Species TAPA n. sp..................................... 315 Figure 6. 94. Species F n. sp. . ........... . .......................... 317 Figure 6. 95. Species F n. sp. . ...................................... 319 Figure 6.96. Species TACB n. sp..................................... 321 Figure 6.97. Species TACB n. sp..................................... 323 Figure 6. 98. Species 00 n. sp. . .................................... 325 Figure 6.99. Species 00 n. sp. . .................................... 327 Figure 6.100. Species 00 n. sp. . ................................... 329 Figure 6.101. Species TYPA n. sp. . .................................. 331 Figure 6.102. Species PSMA n. sp .................................... 333 Figure 6.103. Species PSMA n. sp.................................... 335 xiii LIST OF TABLES Table 1.1.: Morphological characters of C.synarthra and C.mediocauda n.sp. . ..... 14 Table 2.1.: Setal formula for the pereiopods of Tetragonicips pacifica n.sp......... 30 Table 3.1.: Setal formula for Serrithorus geei. . ........................... 46 Table 4.1.: Chaetotaxy of Zosime Pacifica n. sp. . ......................... 73 Table 4.2.: Chaetotaxy of the copepodids of Zosime pacifica n. sp.............. 74 Table 4.3: Chaetotaxy of Cletodus macrura n. sp. . ........................ 75 Table 4.4.: Chaetotaxy of the copepodids of Cletodes macrura n. sp............. 76 Table 4.5.: Chaetotaxy of Cletodes tuberculatus n. sp. . ..................... 77 Table 4.6: Chaetotaxy of the copepodids of Cletodes tuberculatus n. sp. . ........ 78 Table 5.1. Harpacticoid species covered in pictorial key. . .................. 127 xiv ACKNOWLEDGMENT This study was funded by the Pacific Outer Continental Shelf Region of the U.S. Department of the Interior, Minerals Managemtnt Service, Washington, D.C., under Purchase Order No. 13244. xv CHAPTER 1: A NEW SPECIES OF CERVIN/A (COPEPODA: HARPACTICOIDA) FROM THE CALIFORNIA CONTINENTAL SHELF (by Robert Burgess)1 Additional key words: Harpacticoid Taxonomy, Key to the genus, Sexual dimorphism, 1Marine Science Institute, The University of Texas at Austin, Port Aransas, Texas 78373-1267 1 Abstract Cervinia mediocauda n. sp., is a new species of Cerviniidae collected from 15 kilometers off the central California coast, at 154 meters of depth, in the Santa Maria basin. The female and male are described and figured. The following characters on the female make it unique in the genus: the presence of a four segmented exopod on the mandible with 5 associated setae, two sets of two apically mounted setae on the mandibular basis, a maxilliped formula of 1:2:1 for setae associated with the claws on the syncoxa, and moderate length caudal rami. This species appears to be most closely related to C. synarthra. Sexual variation between male and female are pronounced. Introduction Specimens of the new species were obtained during the California Outer Continental Shelf Phase II Monitoring Program (CAMP) (Hyland et. al., 1990), between November 1986 and May 1989. This project was a large, multidisciplinary study to detect and evaluate the long term biological impacts of continental shelf oil drilling and production. The study was centered around a proposed platform site named Julius (referred to hear after as PJ), which was never put into service. Samples were collected in a radial pattern around PJ (0.4 to 2 km rings), and also on a regional scale (10-20 km) on al I sides of the proposed site (Hyland et. al. 1990). A map showing the sample sites, and explanation of the general features of the study area can be found in Montagna 1991. Specimens of Cervinia mediocauda were obtained from both the macrofaunal and meiofaunal samples collected for this study. In addition to a species description, a discussion of sexual dimorphism, a table of some important morphological characters, and a key to the genera are presented. Materials and Methods Holotypes were drawn whole, then dissected. The individual appendages were permanently mounted in lactophenol ringed with BIOSEAL. Paratypes not dissected were preserved in 70% ethanol with glycerol. The figures were drawn with the aid of a drawing tube on a phase contrast microscope. The descriptive terminology is predominately adapted from; Huys et al (in press), except for hyaline frills (Moore 1976), and general body shape (Coull 1977). The following abbreviations are used throughout the text, tables, and figures: R= rostrum, A1 == Antennule, 2 A == Antenna, Md.= mandible, Mxr. == maxillule, Mx.=maxilla, Mxp.= maxilliped, P1-P6 = pereiopods 1-6, Exp.= exopod. Taxonomic Account Family Cerviniidae Sars, 1903 Genus Cervinia Norman, 1878 Cervinia mediocauda n.sp. Cervinia sp. A; Hyland et. al., 1990 Material examined: 42 Females, 2 Males. Holotype: Adult female (dissected), and an adult male (dissected), both from station Platform Julius (PJ). Two female paratypes were dissected and mounted on two slides. All other paratypes were examined whole. All female material was from site PJ. The male paratype was from site R4. Site PJ. was located 15 kilometers off Point Sal in the Santa Maria basin on California continental shelf, and site R4 was located 25 km south of PJ. 3 Description of the female: Based on a nonovigerous female. Body. (Fig. 1.1.) Body shape is fusiform (Coull 1977), which is the typical Cervinia body shape. The length of the body, as measured from the base of the rostrum to the end of the caudal rami, is 0.98 mm. The segment bearing the P, is distinct. Body tapers approximately 50% through the prosome, with the widest point near the anterior edge of the cephalosome. There is almost no taper in the urosome. Somatic hyaline frill is plain and entire (Moore 1976). Urosomal segments have a single row of fine spinules along the posterior edge. The penultimate somite of the urosome is covered with several lines of fine spinules. The genital field situated on the anterior half of the genital segment. Genital field (Fig. 1.1.) Located medially along the fusion ridge of the genital segment. On the integument there is both an anterior, and a posterior sclerotized ridge. There is paired seminal receptacles. Most of the internal structures of the genitalia were obscured because of the large amount of sand that the animals ingest. Caudal Rami (Fig. 1.1.) The length/width ratio is 5.8. The rami are divergent, and covered with fine spinules. The rami are longer than the last 2 abdominal segments. The anterolateral setae is reduced. The anterolateral accessory setae is located within the proximal 60% of the rami. The dorsal setae is well developed, and triarticulate. The inner terminal setae is longer than outer terminal setae. Both of the terminal setae are bipinnate and have very long outer setules. Rostrum. (Fig. 1.2.) Rounded and fused to cephalosome. Rostral tip strongly protruding. Antennule. (Fig. 1.2.) Seven segmented, aesthetasc on third segment (165 µm long). All long setae are bipinnate, and most of the short setae are not plumose. Antenna. (Fig. 1.2.) Long allobasis with 2 bipinnate setae on the abexopodal margin. Exopod has 4 segments. First exopodal segment has 2 setae, the next 2 segments have 1 setae each. The penultimate exopodal segment has 3 setae, with a transverse row of small spinules. The endopod consists of one segment, that is covered with 6 rows of small spinules, armed with 3 lateral pennate setae, as well as seven bipinnate terminal setae. Mandible. (Fig. 1.2.) Gnathobase is large and heavily sclerotized. The pars incisiva is bidentate. The lacina mobilis is tridentate. The gnathobase also has 8 additional spines and 1 bipinnate setae on the dorsal corner. Basis has 2 sets of 2 bipinnate apical setae. The endopod is one segmented, with 3 strong bipinnate inner setae, as wel I as 6 terminal setae. Exopod has 4 segments: The joint between first and the second segment is rather indistinct or possibly partly fussed. There is one bipinnate setae associated with each of the first three exopodal segments. The penultimate endopodal segment has 2 4 setae, one being bipinnate. Maxillule. (Fig. 1.2.) Praecoxa has a large arthrite armed with 9 distal spines. It also has 2 setae on the anterior surface (one of which has a blunt tip), and 2 setae on the posterior surface. The coxa-basis bears 5 setae. The coxal epipodite is represented by one unipinnate setae. The endopod has 12 setae. The exopod is reduced, and bears 2 bipinnate setae. Maxilla. (Fig. 1.2.) The syncoxa bears 4 endites. The second endite is reduced, consisting of only a slight protrusion of the syncoxa bearing one setae. Other endites are more pronounced, and bear 3 setae each. The syncoxa is also armed with 2 rows of setules. The basis is armed with a large claw and 3 setae. The endopod is 4 segmented and is armed with 7 setae. Maxilliped. (Fig. 1.2.) The shape of the maxilliped is not subchelate, but instead has a well developed exopod. The syncoxa has 3 claws, the proximal claw has 1 associated setae, the second claw has 2 associated setae, and the distal claw has 1 associated setae. The inner margin of syncoxa has 3 rows of short spinules, and outer margin has 2 rows of long setules. The basis is armed with a large claw and 1 associated setae. The exopod is 2 segmented. The proximal segment has 2 small setae, while the distal segment has 2 terminal claws and 2 lateral setae. P1 (Fig. 1.3.) The intercoxal plate is armed with 5 long setules. The praecoxa has a row of long setules and a small outer bipinnate spine. The coxa has 3 rows of minute spinules, and one long row of setules. The basis is armed with one row of spinules, and one row of setules. The inner spine on the basis is well developed, bipinnate, and reaches beyond the first endopodal segment. While the outer spine is also bipinnate, it is less developed. The exopod and endopod are 3 segmented. All segments, on both rami, have a row of outer margin setules. The endopod is very ornate with each segment having at least 2 rows on interior setules or spinules. The setal"formula is listed in Table 1.1. P2 (Fig. 1.3.) The intercoxal plate is armed with 2 rows of setules. The distal row has 4 stout setules, and the proximal row has 2 fine setules. The coxa is not armed with any spinules or setules. The basis is armed with 3 rows of spinules and one short row of large setules under the exopod, and a small outer spine. The endopod is 2 segmented, and the exopod is 3 segmented. The first endopodal segment has a large inner thorn, a smaller outer thorn, 3 rows of distal setules, and 2 distal rows of spinules. The second segment has 2 distal thorns, and it has 1 row of outer margin setules. The proximal exopodal segment has an outer spine, and is also armed with 3 rows of spinules. The middle segment has an inner setae a row of spinules on the outer margin. Setal formula is Iisted in Table 1. 1. P3 (Fig. 1.3.) lntercoxal plate has 2 sets of setules, the distal set contains one stout 5 setule, and a proximal set consists of 2 finer setules. The coxa is armed with 2 short rows of short spinules. The basis has 3 rows of spinules, and a small outer spine. The endopod is 2 segmented, and the exopod is 3 segmented. The proximal endopodal segment has a small thorn, 2 distal rows of spinules, and one row of outer margin setules. The only ornamentation on the distal endopodal segment is outer margin setules. The proximal exopodal segment has 2 rows of spinules, and one row of fine setules. The middle segment has a row of outer margin spinules. The distal exopodal segment has one short row of outer margin spinules. Setal formula is listed in Table 1.1. P4 (Fig. 1.3.) The intercoxal plate is armed with 2 distal setules. The coxa is armed with one row of spinules. The basis has one row of inner margin spinules, and one bipinnate outer spine. The endopod is 2 segmented, and the exopod is 3 segmented. The proximal endopodal segment is armed with one distal row of spinules, and one row of outer margin setules. The distal segment has setules associated with all the inner setae, and it also has a row of outer margin setules. The proximal exopodal segment has 3 rows of spinules that wrap around the outer margin. The second exopodal segment has 1 row of outer margin spinules. The distal segment has one short row of outer margin setules. Setal formulas are listed in Table 1.1. P5 (Fig. 1.3.) The baseoendopod is small and unarmed except for basal seta. The exopod has 3 setae, with the innermost setae being un ipinnate. P6 (Fig. 1.3.) The leg is located just posterior of the genital field, and it is laterally displaced. Exopod has one terminal bipinnate setae, one outer bipinnate spine, and an inner thorn. Description of the male: Cerviniids have pronounced sexual dimorphism(Montagna, 1981). Most of the parts of a male are modified from the female. However, despite this dimorphism, males and females of the same species exhibit many taxonomically significant similarities (Montagna, 1981 ). Al I known males have 3 segmented endopods even though the female may only have 2 segments, highlights the extent of sexual dimorphism found within this genus (Huys et. al., in press). Since each appendage has so many slight variations from the female, comparisons would either grossly overstate the differences between the male and female or gloss over small differences, Similarities between the male and female will be noted as to aid in description of the male, and the differences from the female will be pointed out. Body. (Fig. 1.1.) Body shape is fusiform (Coull, 1977). Length of the body, as measured from the base of the rostrum to the end of the caudal rami, is 0.97 mm. The segment bearing the Pl is distinct. Body tapers approximately 30% through the prosome, which is less than the female's 50%. The widest point of the body is near the posterior edge of the cephalosome. Somatic hyaline frill is plain and entire (Moore 1976). The last prosomal segment has a single row of spinules along the posterior margin. The urosomal segments have many interior rows of fine spinules, and a row of larger spinules at the posterior edge of each segment. The second urosome segment carries paired latterly displaced spermatophores. Of course, urosome 3 is not fused to urosome 2, as in the female. Caudal Rami. (Fig. 1.1.) The length/width ratio is 6.1, which is slightly shorter than the females 5.8. All other aspects of the rami were the same a the female. The inner and outer terminal setae were missing from both the holotype and the paratype. Rostrum. (Fig. 1.4.) The rostrum is much larger than the female, triangular, and has a heavily sclerotized ridge running down the center and along the borders. Like the female, the rostrum has a strongly protruding rostral tip, and is fused to cephalosome. Antennule. (Fig. 1.4.) There are eight segments, unlike the females seven segments. It is only weakly haplocer. Large aesthetascs are found on second, third, and fourth segments (183 µm, 194 µm, and 164 µm long respectively). All setae are bipinnate except one on segment 4 and the terminal setae on segment 8. Antenna. (Fig. 1.4.) The appendage on the male is approximately 1/3 smaller than the female's. However, the features are much the same as what is found on the female. There is only 2 minor differences: the setae on first exopodal segment are much shorter than female and they are not plumose, and second there is only 4 instead of 6 rows of small spinules on the endopod. Mandible. (Fig. 1.4.) Th is appendage is almost half the size of the female appendage. The pars incisiva is much more pointed, but like the female it is also bidentate. The lacina mob ii is, Iike the female, is tridentate. The gnathobase has 1 bi pinnate setae on the distal corner, and 4 additional spines, whereas the female has 8 spines. The basis, as in the female, has 2 apical rows of 2 bipinnate setae. The endopod is like the female, except for having 1 row of setules instead of 3 rows. The exopod has 4 segments, however the first one is distinct from the second. Also the male has several rows of large spinules on the proximal exopodal segment, which the female lacks. The setal formula of the exopod is exactly Iike female. Maxillule. (Fig. 1.4.) The male appendage is almost half the size of the female's. The shape of the coxa-basis, and the exopod are only slightly different than female. The setal formula is the same as the female for the all the parts, excluding the arthrite. The distal portion of the arthrite was damaged in dissection and therefore is unknown, but like the female it also has 2 setae on the anterior surface. 7 Maxilla. (Fig. 1.4.) The male's appendage is less than half the siz.e of the female's, but the shape is the same except the basis protrudes more. Syncoxa bears 4 endites as in female. Setal arrangement on the syncoxa is the same as female. The male syncoxa lacks the 2 rows of setu les that are found on the female. SetaI arrangement on the basis is the same as the female. The endopod is 4 segmented, I ike female, but has 10 setae. Maxilliped. (Fig. 1.4.) Approximately 1/3 smaller, but shaped like the female's. The syncoxa has 3 claws. However, no setae are associated with the claws, unlike the female part . The inner margin of syncoxa has 3 rows of long spinules, like the female, but they are arranged differently. The outer margin only has 1 line of long setules, unlike the female who has 2 rows. The basis is armed with a large claw, I ike the female, however there is no associated setae. The exopod is shaped the same and the setal formula is the same as the female. P1 (Fig. 1.5.) Approximately the same size leg as in the female. It also has the same number and arrangement of setae as the female. The intercoxal plate has the same number of rows of setules as the female. The coxal armature is the same as the female. While the basis is armed with one row of spinules and one row of setules as in the female, the arrangement on the basis is different. The inner spine, on the basis, is well developed and bipinnate, however it is not as developed as the female. All endopodal and exopodal segments have large spinules, unlike the female who's spinules are comparatively short and fine. The endopodal segments have less pronounced spines and thorns than the female. But they have rows of inner spinules on each segment, as does the female. The exopod is more ornate, than the female, having large outer margin spinules on all the segments, and 2 rows on the interior of exopod 1. Setal formula is listed in Table 1.1. P2 (Fig. 1.5.) As in the female, the intercoxal plate is armed with 2 rows of setules, however, the distal row has 9 stout setules not 4, the proximal row has 3 fine spinules not 2. The male coxa is armed with 5 rows spinules, while the female is unarmed. The male basis armed the same as the female, but the spinules are much larger. The endopod has 3 segments, unlike female, therefore the setal formula is different. The first endopodal segment has the same setal and thorn arrangement. However, the distal thorns are less pronounced, and lacks a row of inner margin setules. The second segment has the 2 distal thorns, and 2 rows of spinules. The terminal endopodal segment has 6 terminal thorns, and 3 rows of spinules. The setal formula of the exopod is the same for both sexes. The proximal exopodal segment of the male is different than the female in having 2 rows of interior spinules, a row of inner margin setules, and the outer margin spinules are much larger than the female's. The middle exopodal segment differs from the female only by a few spinules located on the interior of the segment. The terminal segment is the same as the female, except the outer most terminal setae is transformed into a spine, and there is a row of small outer margin spinules. Setal formula is listed in Table 1.1. 8 P3 (Fig. 1.5.) lntercoxal plate armed with 2 sets of setules, as in the female. However, the distal row has 8 fine setules, the proximal row has 3 stout setules, instead of 1 and 2 respectively. Coxal ·armature is the same as the female. The basis has 3 rows of spinules, as in the female, but the arrangement on the segment is different. Also, there is a small spine below the outer spine. The endopod is 3 segmented, therefore the setal formula is different. The proximal segment has the same setal and thorn arrangement. However, the distal thorns are less pronounced, it lacks a row of setules near the inner distal thorn, and has an extra row of spinules located proximally to the inner spine. The middle segment has two rows of setules. The distal segment has 2 distal thorns, and a row of outer margin setules. Setal formula for the male exopod is the same as the female. The proximal exopodal segment has an row of interior spinules, which the female lacks. The middle exopodal segment has a row of interior spinules, which the female lacks. The terminal exopodal segment is like the female, except the outer most terminal setae of the female is transformed into a spine on the male. Setal formula is listed in Table 1.1. P4 (Fig. 1.5.) This leg is approximately 20 percent larger in the male, than in the female. lntercoxal plate is armed with 2 row of setules, unlike the female which only has one row. The distal row has 6 setules, instead of the female's 2, and the proximal row has 3 stout setules. The coxa has 2 more rows of spinules than the female. The basal armature is similar to the female, however it also has one spinule under the outer spine. The endopod is 3 segmented, and therefore has a different setal formula than the female. The proximal endopodal segment's inner thorn is more pronounced than on the female. The second segment has a row of outer margin setules, and an inner thorn. Th~ distal segment ends in 8 thorns. The setal formula for the exopod is the same as in the female. The proximal exopodal segment has an inner setae which is much longer than the female's, and it is also missing a row of spinules that is found on the female. The second exopodal segment also has an inner setae which is greatly enlarged as compared to the female, and it also has an extra row of spinules. The only difference between the male and female on the terminal exopodal segment is that the outer most terminal setae on the female is transformed into a spine on the male. Setal formulas are listed in Table 1.1. P5 (Fig. 1.5.) This leg is exactly the same as the female, except for having a short row of outer margin spinules. P6 (Fig. 1.5.) This leg's armature is like the female. However, it is not as well defined as in the female. Variations. Variation in setation of the appendages of the female were not observed in any of the dissected paratypes. Some slight variation in the number of the intercoxal setules was observed. Variation in the female caudal rami length width ratio was so slight, that the observed variation was mostly due to vertical pitch of the rami during observation of the specimen. Only one male was dissected. Etymology. The name mediocauda ( L. 'medius' = middle + L. 'cauda' ==tail) refer to the 9 moderate length of the caudal rami. Discussion According to the latest key in Montagna 1981, Cervinia. mediocauda n. sp. Is in the Cervinia synarthra group. There are 2 known species in this group, Cervinia synarthra (Sars, 1903) and Cervinia pi/osa (Lang, 1948). Both of which have the same setal formula for the pereiopods. C. pi/osa has a long proximal setae on the caudal rami, and C. synarthra has a short setae. C. mediocauda has a short proximal setae and therefore resembles C. synarthra in this way. The setal formula of the mouthparts and the number of segments on the rami of the mandible is the most reliable way of separating the species in the genus Cervinia (Montagna, 1981 ). While other families use the setal formula of the pereiopods as a reliable way to separate species of harpacticoids this doesn't hold true in Cervinia. All three of the species in C. synarthra group have the same setal formula on the pereiopods. In Cervinia, this similarity of the setal formula of the pereiopods is not limited to the C. synarthra group, four out of the remaining seven species in the genera have also group together to share a common pereiopodal setal formula (Montagna 1981 ). In addition, one of the remaining three can not grouped by pereiopodal formula is due to the fact that the pereiopodal setal formula is unknown. C. mediocauda is unique in the genus in the following ways: the mandibular basis is armed 2 sets of 2 bipinnate setae, the mandibular exopod is 4 segmented with 5 associated setae, and on the maxillipedal endites the claws have an accessory setae formula of 1-2-1, respectively. The new species also differs from C. synarthra in the in the fact that its approximately 1/3 smaller, and has shorter caudal rami. The new male has caudal rami that place it into the genus Cervinia. C. magna is the only male from this genus that has been described (Montagna 1981 ). The C. mediocauda male has a 3 segmented endopod unlike C. magma's 2 segments. The setal formula of the P 1 and the exopods of P2 through P.t are the same in the male as the female C. mediocauda. The P5 and P6 of the male and female are shaped exactly the same and possess the same setal formula. The setal number and arrangement on mandibular basis are the same between the sexes, and the mandibular exopod has the same number of segments, while being different than any other known species. I found no other species of Cerviniids in my samples, and the 2 males found are the same. The caudal rami length\width ratio is relatively constant between male and female in C. magna (Montagna 1981), and this appears to be true in C. mediocauda, were it varies between 5.8 for the female to 6.1 for the male. Sexual dimorphism is pronounced in this family(Montagna 1981). Most parts have differences in the number of setal/spinule rows or their arrangement. However, the taxonomically important parts between the male and female are very similar. Some parts are characteristically modified in Cervinia males, such as the rostrum, antennule, and also all males have 3 segmented endopods even if the female only has 2 segments(Huys et al, in press). In C. mediocauda the mouthparts are reduced 30% to 50% in size from the female. 10 Montagna (1981) had concluded that setation of the mandible and the maxilliped are taxonomically important and conservative between the sexes. The mandibular palp was unchanged between the sexes, and unique in the genus. However, sexual dimorphism has resulted in the Mxp. being greatly reduced in size and it also lacks any accessory seate on the syncoxal endites. The caudal rami UW ratio does not change greatly, varying only from S.8 for the female to 6.1 for the male. The PS and P6 have exactly the same shape and setal formula in either sex of C. mediocauda, as was noted for the male and female PS of C. magna (Montagna 1981 ). The setal formula for the exopod and endopod of the P1, and exopods of P2 through P4 were also conservative between the male and female, but as I pointed out earlier, a third of the genus also have this same setal formula. Key to the species in the genus Cervin ia Based on females. Key format adapted from Montagna 1981. C.brevipes was omitted since the setal formula for the walking legs is unknown. 1. P2-P4 with 2 segmented end.-"synarthra" group . . . . . . . . . . . . . . . . . . . . . 2 P2-P4 with 3 segmented end.-"bradyi" group . . . . . . . . . . . . . . . . . . . . . . . 6 2 P1 with 3 segmented end . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 P1 with 2 segmented end. C.magna Smirnov 3 P2-P4 end. terminal segment with 6, 7, 6 setae and spines respectively . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 P2-P4 end. terminal segment with 7, 8, 7 setae and spines respectively ............................. C.langi Montagna 4 CR with short lateral proximal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 CR with long lateral proximal setae ..................... C.pilosa Lang 5 Maxilliped syncoxa claws with 0.1.0 associated setae, mandibular exp. 2 segmented .................. C.synarthra Sars Maxilliped syncoxa claws with a 1.2.1 associated setae, mandibular exp. 4 segmented . . . . . . . . . . . . . . . C.mediocauda n.sp. 6 CR longer than last urosome segment (UW >S) . . . . . . . . . . . . . . . . . . . . 7 CR as long as the last urosome segment (UW -=4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C.bradyi Norman 7 Basis of Mxp. Twice as long as end. segments; P5 exp. with 3 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Basis of the Mxp. As long as the 3 enp. segments; P 5 exp .with 2 setae .......................... C.tenuiseta Brodskaya 11 8 CR as long as last 4 urosome segments (UW = 12) C.tenuicauda Brodskaya CR as long as-last 2 urosome segments (UW =8) ................................... C.unisetosa Montagna Acknowledgments I am much indebted to Dr. Paul Montagna, at the University of Texas Marine Science Institute, not only for the specimens and for use of the facilities, but also for his professional and technical support. I am grateful to Rick Kalke for his taxonomic prowess and his moral support in my times of need. I am also grateful to Mark Lanzotti for his grammatical suggestions on earlier drafts. 13 Table 1.1.: Morphological characters of C.synarthra and C.mediocauda n.sp. Information for C.synarthra is from Montagna 1979 and Sars 1903. Both species have a P1 exopod formula of 1.1.123, and the females have a P2-P4 exopodal formula of 1.1.223. However, as illustrated below, the species differ in the following charaters: the endopodal setal formulas, the length width ratio of the caudal rami, the setal formula on the mandibular basis (Md. basis) and exopod(Md. exp.), and also in the setation of the syncoxal endites of the maxilliped(Mxp. setae). The male has a reduced Mxp with no accessory setae on the endites. Species Endopodal setal formula CR Md. Mxp. P1 P2 P3 P4 Ps L/W Basis Exp. setae C. synarthra Sars S? 1.1.22 1 1.321 1.421 1.321 3 7.3 3 2 I 7 1.1.1 C. mediocauda n.sp. S? 1.1.22 1 1.321 1.421 1.321 3 5.8 4 4/5 1.2.1 C. mediocauda n.sp o 1.1.22 1 1.2.22 1 1.2.32 1 1.2.22 1 3 6.1 4 4/5 0.0.0 14 Figure Legends Figure 1.1.: Cervinia mediocauda n.sp. ~ and o. Figure 1.2.: Cervinia mediocauda n.sp. ~ mouth parts. Figure 1.3.: Cervinia mediocauda n.sp. ~ legs. Figure 1.4.: Cervinia mediocauda n.sp.o mouth parts. Figure 1.5.: Cervinia mediocauda n.sp.o legs. 15 Femal.e Mal.e Whole Body Drawings .2mm / ·1\ ~~ .. l --·,~r---_) ' ~ -~ " ~ 1~!\ --............................. I ."2'. -­ !, .· , ' . /' j GF Mal.e Femal.e VENTRAL CR SOµm SOµm SOµm .1 mm Mx Md, Mxl, Mx, Mxp SOµm /fl n'u j I \\ j v; p6 Ps P, SOµm SOµm Ai & R Md \ \ I I \! I Mxp Md, Mxp Mxl, Mx SOµm SOµm SOµm SOµm l.O References Coull B. C. (1977) Marine flora and fauna of the northeastern United States. Copepoda: Harpacticoida. NOAA Tech. Report. NMFS Circ 399.48pp Huys R., Gee J.M., Moore C.G., and Hamond R. (In press) Marine and brackish water harpacticoid copepods Part 1. Synopses of the British Fauna (Ed. R.S.K. Barnes & J.H. Crothers) No. 51 Field Studies Council Press UK, pp 363 (approx). Hyland J., D. Hardin, I. Crecelius, D. Drake, P. Montagna, and M. Steinhauer (1990) Monitoring long-term effects of offshore oil and gas development along the southern California outer shelf and slope: background environmental conditions in the Santa Maria Basin. Oil and Chemical Pollution, 6, 195-240. Lang, K. (1948). Mongraphie der Hrpacticiden, /&II. Hakan Ohlsson, Lund. 1682 pp. Lang, K. (1965). Copepoda Harpacticoida from the California Pacific coast. Kungl. Svenska Vetensk. Hand I., Fjarde, 10(2): 1-560. Montagna P.(1979) Cervinia langi n.sp. and Pseudocervinia magna (Copepoda: Harpacticoida) from the Beaufort Sea(Alaska, USA.). Trans. Amer. Micros. Soc., 98(1):77-88. Montagna P.(1981) A new species and a new genus of Cerviniidae (Copepoda: Harpacticoida) from the Beaufort Sea, with a revision of the family. Proc. of Biol. Soc. of Washington, 93(4): 1204-1219. Montagna P. (1991) Meiobenthic communities of the Santa Maria Basin on the California shelf. Continental Shelf Research, 11(11):1355-1378. Moore C. G. (1976) The form and significance of the hyaline frill in harpacticoid copepod taxonomy. J. Nat. Hist Sars, G.O. (1903) An account of the Crustacea of Norway. Copepoda Harpacticoida. Bergen Museum, Norway. 499 pp. 21 CHAPTER 2: A NEW SPECIES OF TETRAGONICEPS (COPEPODA: HARPACTICOIDA) FROM THE CALIFORNIA CONTINENTAL SHELF(by Robert Burgess)1 Key words: Taxonomy, Tetragonicepitidae. 1Marine Science Institute, The University of Texas at Austin, Port Aransas, Texas 78373-1267 Abstract Tetragoniceps pacifica n.sp., is a new species of Tetragonicipitidae collected 15 kilometers off the central California coast, at 154 meters of depth in the Santa Maria basin. The female is described and figured. The following characters on the female make it unique in the genus: a setal formula of 1.021 on endopods of P2 through P4, the P2 exopod having a setal formula of 1.1.123, ~ exopod having a setal formula of 1.1.222, and the P4 exopod having the formula of 1.1.123. This species appears to be most closely related to T. scotti. Introduction Specimens of the new species were obtained during the project named The California Outer Continental Shelf Phase II Monitoring Program (CAMP) (Hyland et al., 1990), between November 1986 and May 1989. This was a large, multidisciplinary study designed to detect and evaluate the long term biological impacts of continental shelf oil drilling and production. The study was centered around a proposed platform named Julius (hereafter referred to as PJ), which was never put into service. Samples were collected in a radial pattern around PJ (0.4-2km), and also on a regional scale (10-20 km) on all sides of the platform (Hyland et al., 1990).A map showing the sample sites, and an explanation of the general features of the study area can be found in Montagna 1991. Specimens were obtained from both the macrofaunal and meiofaunal components of this study. Materials and Methods The holotype was drawn whole, then dissected. The appendages were permanently mounted in lactophenol ringed with BIOSEAL. The paratype was not dissected, and is preserved in 70% ethanol with glycerol. The figures were drawn with the aid of a drawing tube on a phase contrast microscope. The descriptive terminology is predominately adopted from; Huys et al. (in press), except for hyaline frills (Moore, 1976), and general body shape (Coull, 1977). The following abbreviations are used throughout the text, tables, figures: R== rostrum, A1 == Antennule, Ai== Antenna, Md== mandible, Mxl== maxillule, Mx== maxilla, Mxp== maxilliped, P1-P6 == pereiopods 1-6, Exp== exopod, End== endopod, GF-= genital field, CR== caudal rami Hyl Frill== Hyaline Fri II. Taxonomic Account Family Tetragonicipitidae Lang, 1948 Genus Tetragoniceps Brady, 1880 Tetragoniceps pacifica n.sp. Material examined: 2 Females. Holotype: Adult female (dissected),from station Platform Julius (PJ). The paratype was examined whole. All material was from site PJ., located 15 kilometers off Point Sal in the Santa Maria basin, on California continental shelf. Description of the female: Based on a nonovigerous female. Body. (Fig. 2.1.) There is almost no taper to the body, therefore the shape is cylindrical (Coull, 1977). Length from the base of the rostrum to the end of the caudal rami is 0.72 mm. The segment bearing the P1 is distinct. Somatic hyaline frill is plain and entire (Moore, 1976). Genital field situated in the proximal half of the genital segment. The anal operculum has a row of fine spinules. Genital field (Fig. 2.1.) Located medially, on the anterior half of the genital segment. It has paired genital pores, each located near the P 6• The copulatory pore is pronounced, and located near the fusion ridge between the somites. There seems to be a continual formation of paired seminal receptacles, with the most developed being the anterior most pair which are connected to the genital pores. Caudal Rami (Fig. 2. 1.) The length/width ratio is 2.6. The anterolateral accessory setae is very reduced. The anterolateral setae is large and located within the proximal 25% of the rami, and is located on a large dorsal protrusion. The posterior lateral setae is also large and is located in line with a transverse row of spinules. The dorsal setae is well developed and is triarticulate at the base. The outer terminal setae is greatly reduced, being only about 10% as long as the inner terminal setae. Rostrum. (Fig. 2.2.) Rounded, defined at base. Two sensilla are located on the outer margin. Antennule. (Fig. 2.2.) Nine segmented. There is an aesthetasc on fourth segment (78 µm long). All setae plain, having no associated setules or spinules. Antenna. (Fig. 2.2.) It has a long basis. The exopod is one segmented, with 2 bipinnate setae, and 1 unipinnate setae. The endopod is 2 segmented. The first endopodal segment is unarmed. The distal endopodal has the following ornamentation: a thorn and a small spine on the exopodal margin, two small lateral nonpinnate spines and two larger unipinnate spines on the abexopodal margin, 3 nonpinnate terminal setae, 2 sets of 2 geniculate, subterminal setae which are dorsally inserted. Mandible. (Fig. 2.2.) Gnathobase is large and heavily sclerotized. The. pars incisiva is bidentate. The lacinia mobilis is quadradentate, with the setae closest to the pars incisiva being bidentate itself. The gnathobase also has 4 additional spines and 1 unipinnate setae on the dorsal corner. Basis has 3 apical bipinnate setae and 1 small spine. The endopod is made from 2 fused segments, has a single lateral plain setae on the inner edge of the fusion point, and 4 plain terminal setae. The exopod is 2 segmented. The first exopodal segment has 2 lateral setae, and the second segment has 2 terminal setae. Maxillule. (Fig. 2.2.) Praecoxa has a large arthrite which is armed as follows: 7 distal spines, 2 terminal bipinnate setae, one nonpinnate terminal setae, 2 bipinnate setae on the anterior surface, and 2 bipinnate setae on the posterior surface. The coxal lobe of the coxa-basis bears 4 terminal setae. The basal lobe bears 3 nonpinnate setae and 2 larger bipinnate setae. The coxal epipodite is represented by one bipinnate setae. The endopod has 4 setae, and a row of long outer margin setules. The exopod has 2 setae, and a row of long inner margin setules. Maxilla. (Fig. 2.2.) Syncoxa bears 4 endites, and a row of spinules. The first endite has one large bipinnate spine, and one small setae. Second endite only has one setae. The third has one bipinnate setae and two nonpinnate setae. The most distal endite has one unipinnate, one bipinnate and one nonpinnate setae. The allobasis is armed with a claw and 4 accessory setae. The endopod is 4 segmented, segments 1 through 3 have a single associated lateral setae, and the fourth segment has 2 terminal setae. Maxilliped. (Fig. 2.2.) The shape is subchelate(Huys et al., in press). Syncoxa has 3 pinnate setae, and 3 rows of spinules. The basis is armed along the inner margin as follows: a setae, a small spine, and several setules. The exopod has 1 terminal claw, 1 terminal pinnate setae, and 1 subterminal nonpinnate setae. P1 (Fig. 2.3.) The coxa has 2 rows of minute spinules. The basis is armed with 3 rows of spinules, and 2 spines. The inner spine is well developed and bipinnate, the outer spine is also bipinnate, but is less developed. Exopod is 3 segmented, the endopod is 2 segmented. All exopodal and endopodal segments have a row of outer margin setules. The first endopodal segment is elongate reaching beyond exopod 3. It is armed with an inner comb of long setules. The second endopodal segment has only an comb of short outer margin setules and two terminal unipinnate setae. There are no inner setae on the first 2 segments of the exopod. Exopod 1 through exopod 3 have margin setules, but only exopod 2 has a comb of inner setules. Exopod 3 has 2 geniculate unipinnate terminal setae. Setal formula is listed in Table 2.1. Hyaline Frill (Fig 2.1.) On legs P2 thru P4 on exopod 1 and exopod 2, at the distal end of these segments there is a row of semi-incised broad subulate appendicular hyaline frill (Moore, 1976). P2 (Fig. 2.3.) Coxa is armed with 2 rows of long setules. Basis is armed with: one row of long setules, one row of short spinules under the endopod, and an nonpinnate outer setae. The endopod is 2 segmented, and the exopod is 3 segmented. The first endopodal segment has both inner and outer margin setules, and an outer thorn. The distal endopodal segment has 2 distal thorns, and both an inner and a outer row of marginal spinules. The proximal exopodal segment has a large outer thorn, a row of inner setules, and a row of outer margin spinules. The middle segment has a large outer thorn, a row of spinules on the outer margin, and a row setules on the inner margin. Exopod 3 has a terminal thorn, and also a row of outer margin setae. Setal formula is listed in Table 2.1. p (Fig. 2.3.) The coxa is armed with 2 short rows of long spinules. The basis has 1 short row of spinules, and a thorn on either side of the endopod. The endopod is 2 segmented, and the exopod is 3 segmented. The proximal endopodal segment has a row of outer spinules, a small inner thorn, and a small outer thorn. The distal segment has 2 distal thorns and a row of outer margin setules. The proximal exopodal segment has a large outer thorn, a row of outer margin spinules, and a row of inner margin setules. Exopod 2 has a large outer thorn, a row of outer margin spinules, and a row of inner margin setules. The distal exopodal segment is armed with 2 outer thorns. Setal formula is listed in Table 2.1. P4 (Fig. 2.3.) The coxa is armed with one row of long spinules. The basis has one outer setae, and 2 small thorns surrounding the endopod. The endopod is 2 segmented and the exopod is 3 segmented. The proximal endopodal segment has a small outer thorn. The distal segment has 2 distal thorns, a row of inner setules, and a row of outer setules, The exopod is almost 3 times as long as the endopod, and conspicuously stout. The proximal exopodal segment has a large outer thorn, a row of outer spinules, and a row of inner setules. The second exopodal segment has a large thorn, a row of inner margin setules. The distal exopodal segment has 2 small outer spines, and inner and outer marginal spinules. Setal formulas are listed in Table 2.1. (Fig. 2.3.) The Baseoendopod is very pronounced and is armed not only with a basal P5 seta but also 2 unipinnate spines, 3 setae, 2 rows of setules, and a pore or scar located distally to the distal spine. The exopod very pronounced and elongate. It has 6 setae. It also has 2 pores or scars located on the inner and outer margin, both of which are located on the anterior third of the appendage. Variations. Variation the female were not obvious in between the holotype and the un­dissected paratype. No variation of the PS was found between the two specimens. Etymology. The name pacifica refers to Pacific Ocean, where it was found. Discussion Tetragoniceps pacifica keys to Tetragoniceps according to the latest key (Coull 1973). There are ten known species in the genus. Tetragoniceps. pacifica is most closely related to T. scotti (Sars, 1911 ). However, T. pacifica has the following characteristics that are not only different than T. scotti, but also unique in the genus: it has P2 exopod setal formula of 1.1.123, a P3 exopod setal formula of 1.1.222, a P2 through P4 endopod setal formula of 1.021,and 6 setae on the baseoendopod of the P 5• I have come to the conclusion that the "holes" on the P 5 are pores. Both the appendages from the holotype, and paratype, were examined. All appendages were exactly the same as far a location of the pores, and none had any evidence of a setal attachment. No appendage showed any sign of physical damage. Also the pores on the exopod occur in the upper third of the segment, a position where they would be relatively protected from physical damage if they were a setal attachment points. Acknowledgments I am much indebted to Dr. Paul Montagna, at the University of Texas Marine Science Institute, not only for the specimens and for use of the facilities, but also for his professional and technical support. I am grateful to Rick Kalke, for his taxonomic prowess and also his technical expertise. Table 2.1.: Setal formula for the pereiopods of Tetragonicips pacifica n.sp. LEG EXOPOD ENDOPOD P1 0.0.013 1.020 P2 1.1.123 1.021 P3 1.1.222 1.021 P4 1.1.322 1.021 Figure Legend Figure 2.1.: Tetragonicips pacifica n. sp. ~ Figure 2.2.: Tetragonicips pacifica n. sp. ~ Figure 2.3.: Tetragonicips pacifica n. sp. ~ ~ABITUS GF . 2mm 50 µm Hyl Frill 20 µm 50 µm 32. ~ f: ~ A2 Mxp Mx Md & Mxp Mxl & Mx 50 µm 50 µm 50 µm 50 µm References Coull B. C. (1973) Harpacticoid copepods of the family Tetragonicipitidae (Lang); a review and revision, with keys to the genera and species. Proceedings of the Biological Society of Washington D.C., 86 (2):9-24. Coull B. C. (1977) Marine flora and fauna of the northeastern United States. Copepoda: Harpacticoida. NOM Tech. Report. NMFS Circ 399.4Bpp Huys R., Gee J.M., Moore C.G.,and Hamond R. (In press) Marine and Brackish water harpacticoid copepods Part 1. Synopses of the British Fauna (Ed. R.S.K. Barnes & J.H. Crothers) No. 51. Field Studies Council Press UK, pp363 approx. Hyland J., D. Hardin, I. Crecelius, D. Drake, P. Montagna, and M. Steinhauer (1990) Monitoring long-term effects of offshore oil and gas development along the southern California outer shelf and slope: background environmental conditions in the Santa Maria Basin. Oil and Chemical Pollution, 6, 195-240. Lang, K. (1948). Mongraphie der Hrpadiciden, 1&11. Hakan Ohlsson, Lund. 1682 pp. Lang, K. (1965). Copepoda Harpadicoida from the California Pacific coast. Kung/. Svenska Vetensk. Hand/., Fjarde, 10(2): 1-560. Moore C. G. (1976) The form and significance of the hyaline frill in harpadicoid copepoda taxonomy. J. Nat. Hist Sars, G.O. (1911) An account of the Crustacea of Norway. Copepoda Harpacticoida. Bergen Museum, Norway. 499 pp. CHAPTER 3: A NEW GENUS AND NEW SPECIES OF CLETODIDAE (COPEPODA: HARPACTICOIDA) FROM THE CALIFORNIA CONTINENTAL SHELF (by Robert Burgess)1 Key words: Taxonomy, Serratithorus N. Gen., Cletodidae. 1Marine Science Institute, The University of Texas at Austin, Port Aransas, Texas 78373-1267 Abstract Serratithorus geei is anew genera and species of Cletodidae collected 15 kilometers off the central California coast, at 154 meters of depth in the Santa Maria basin. The following characters on the female make it unique in the family: a large fused dorso-ventrally curved rostrum, Ai with an exopod with three setae and two setae on the abexopodal margin, two dorsal rows of large spines, and no inner seta on the P, exopod 2. The copepodite Enhydrosoma nicobarica described by Sewell (1940), has a close affinity with the new species, and should be transferred to the new genus. This genus appears to be most closely related to the genus Enhydrosoma based on the exopod of the Ai, and its having five setae on the distal segment of the exopod of P3 andP4• Introduction Specimens of the new species were obtained during the project named The California Outer Continental Shelf Phase II Monitoring Program (CAMP) (Hyland et al., 1990), between November 1986 and May 1989. This was a large, multidisciplinary study designed to detect and evaluate the long term biological impacts of continental shelf oil drilling and production. The study was centered around a proposed platform named Julius (hereafter referred to as PJ), which never was put into service. Samples were collected in a radial pattern around PJ (0.4-2km), and also on a regional scale (10-20 km) on all sides of the platform (Hyland et al., 1990). A map showing the location and an explanation of the general features of the study area can be found in Montagna 1991. Specimens were obtained from both the macrofaunal and meiofaunal components of this study. Materials and Methods The holotype was drawn whole, then dissected. The appendages were permanently mounted in lactophenol ringed with BIOSEAL. The paratypes were not dissected, and are preserved in 70% ethanol with glycerol. The figures were drawn with the aid of a drawing tube on a phase contrast microscope. The descriptive terminology is predominately adopted from: Huys et al. (in press), except hyaline frills (Moore, 1976), and general body shape (Coull, 1977). The following abbreviations are used throughout the text, tables, figures: R= rostrum, A1 == Antennule, Ai== Antenna, Md= mandible, Mx=maxillule, Mxl= maxillule, Mxp= maxilliped, P1-P6 -= pereiopods 1-6, Exp= exopod, End-= endopod, GF= genital field, CR= caudal rami Taxonomic Account Family Cletodidae Scott Genus Serratithorus, n.gen Serratithorus geei n.sp. Material examined: 9 Females, 10 Males. Holotype: Adult female (dissected) and male (dissected), are from station R4. Two paratypes were dissected, the rest were examined whole. All material was from site R4, located 15 kilometers off the Californian coast in the Santa Maria basin. Description of the female: Based on a nonovigerous female. Body. (Fig. 3.1.) There is almost no taper to the body, therefore the shape is cylindrical (Coull, 1977). Length from the base of the rostrum to the end of the caudal rami is 0.88 mm. The cephalothorax has a pronounced dorsal protrusion of the cephalic shield, which almost reaches the distal end of the first free prosomite. The shield has four sensilla on the posterior margin. The somatic hyaline frill is plain and entire (Moore, 1976). The free somites except the last 2 abdominal segments have 2 dorsal rows of large thorns, and a row of ventral thorns on each side. There are also 2 other rows of small sensilla on either side. The anal operculum has a sensilla on either side. Genital field (Fig. 3.1.) Located medially, on the anterior half of the genital segment. There are paired genital pores. The P 6 is very reduced being represented only by one setae. The copulatory pore is pronounced, and located near the fusion ridge between the somites. A subcuticular ridge marks the line of fusion. Caudal Rami (Fig. 3.1.) The length/width ratio is 4.4. The anterolateral accessory setae is located on the ventral surface, and is very reduced. The anterolateral setae is large and located near the proximal edge of the rami. The posterior lateral setae is large and is on the outer margin within the anterior 40 percent of the rami. The dorsal setae is well developed, triarticulate at the base, and is located on the anterior 20 percent. The outer terminal setae is greatly reduced, being only about 10% as long as the inner terminal setae. Rostrum. (Fig. 3.2.) Fused at base. Triangular with the point being recurved dorsally. Two sensilla are located on the outer margin, and 2 small spinules on the tip. Antennule. (Fig. 3.2.) Five segmented, with an aesthetasc on third segment (53 µm long). Setae on the first 2 segments and some on the penultimate segment are bipinnate. The peduncle is very elongate and protrudes beyond the anterior edge of the integument of the cephalothorax. Antenna. (Fig. 3.2.) It has an allobasis with 2 bipinnate setae on the abexopodal margin. The exopod has 1 lateral bipinnate setae, 1 terminal bipinnate setae, 1 small plain setae, and a row of subterminal spinules. The terminal bipinnate setae has stout outer spinules while the inner setules are much finer and more numerous. The endopod has the following ornamentation: a row of small subterminal spinules starting on the exopodal margin, two rows of large spinules on the abexopodal margin, 2 nonpinnate subterminal setae on the abexopodal distal third of the exopod, 2 geniculate terminal setae, 2 terminal nonpinnate setae, on bipinnate setae, and one very small nonpinnate setae. Mandible. (Fig. 3.2.) Syncoxa bears one row of large spinules. Gnathobase is large and heavily sclerotinized. The pars incisiva is unidentate. The lacinia mobilis seems to be fused to the gnathobase. All teeth are multicuspid. There is one unipinnate setae on the distal corner. The endopod and exopod are fused to the basis but can still can be recognized. The basis is armed with 3 bipinnate setae, and each vestigial rami has one bipinnate setae. 39 Maxillule. (Fig. 3.2.) Praecoxa has a large arthrite which is armed as follows: 4 distal spines, 1 terminal bipinnate setae, one nonpinnate anterior setae. The coxa bears one large claw. The endopod and exopod are fused to the basis. The basal lobe is partially separated dorso-ventrally into anterior and posterior lobes or nabs. On the anterior lobe there are 1 bipinnate, one plain terminal setae, and a row of spinules around the outer margin of the anterior lobe. On the posterior lobe this is also 1 bipinnate and one plain setae. The endopod is fused to the basis, and is armed with 2 plain setae, and one bipinnate setae. The exopod is also fused to the basis, and is armed with one bipinnate and one plain setae. Maxilla. (Fig. 3.2.) Syncoxa bears 2 endites, and a row of spinules. The first endite has one bipinnate spine, and one small setae. Second endite has one bipinnate setae, and one plain setae. The allobasis is armed with a claw and 2 accessory setae. The basal endite has one terminal claw with 2 accessory setae. The endopod is reduced, fused to the basis and represented by 2 bipinnate setae. Maxilliped. (Fig. 3.2.) The shape is subchelate (Huys et al., in press). Syncoxa has a short row of spinules. The basis is armed along the inner margin a row of long stout spinules, and along the outer margin with a row of spinules. The exopod has 1 pinnate terminal claw, and 1 terminal nonpinnate setae. P1 (Fig. 3.3.) The coxa has a row of inner and outer margin setules and a row of minute setules located medially. The basis has 2 rows of spinules and 2 bipinnate ~pines. The inner spine has very large spinules, the outer spine's are less developed. The exopod is 3 segmented, segment 1 and 2 are 1.5 times long as broad but the third segment is 2.5 long as broad. Exopod 1 through exopod 3 have outer margin spinules and a row of interior spinules. The proximal exopodal segment has no inner margin setules. The second segment has a row of inner margin setules. The distal exopodal segment is armed with two outer spines and two long bipinnate setae with a small plume of setules at each tip. The first endopodal segment is almost a broad as wide, and the distal segment is 3 times longer than broad. The first endopodal segment is unarmed except for an inner row of long setules and row of outer spinules. The second endopodal segment has both inner margin setules and outer margin spinules. It is armed with 1 small inner subterminal terminal nonpinnate setae, 1 terminal bipinnate spine, and 1 long tripinnate setae with a small plume of setules at the tip. There is no inner setae on the exopod. Setal formula is listed in Table 3.1. Hyaline Frill (Fig 3.1.) On legs P2 thru P4 at the distal end of exopod 1 and exopod 2, there is a row of fully-incised fine subulate appendicular hyaline frill along the inner half of the anterior surface (Moore, 1976). P2 (Fig. 3.3.) Coxa is armed with 2 rows of long setules and one row of minute setules. The basis is armed with: one row of long setules around the basal setae, and one row of long spinules under the endopod, and a nonpinnate basal setae. The endopod is 2 , ... segmented, and the exopod is 3 segmented. The first endopodal segment has both inner margin setules, and outer spinules, but has no setae or spines. The distal endopodal segment has a row of long inner margin setules, 2 rows of spinules which start on the outer margin and stop in the interior of the segment. Exopod 1 through exopod 3 have outer margin spinules and at least one row of interior spinules. The proximal exopodal segment has no inner margin setules. The middle segment and the distal exopodal segments have inner margin setules. The distal exopodal segment has three short rows of interior spinules, and is armed with two outer bipinnate spines and two long bipinnate setae with a plume of setules on the tips of these setae. Setal formula is listed in Table 3.1. P3 (Fig. 3.3.) The coxa is armed with one row of long spinules. The basis has 1 short row of spinules under the endopod, and a row around the basal setae. The endopod is 2 segmented, and the exopod is 3 segmented. The proximal endopodal segment has no other ornamentation than a row of outer margin spinules. The distal segment integumental ornamentation is 2 small spinules, a row of inner margin setules, a row of outer margin spinules. It also has one outer bipinnate spine and 2 terminal bipinnate setae. All exopodal segments have outer margin spinules and at least one row of interior spinules. The proximal exopodal segment has no inner margin setules. Exopod 2 has row of long inner margin setules. The distal exopodal segment has 2 rows of is armed with 2 outer thorns. Setal formula is listed in Table 3.1. P4 (Fig. 3.3.) The praecoxa is armed with one long proximal row of spinules. The coxa is armed with 2 rows of long spinules, and one row of minute spinules. The basis has one row of spinules beneath the endopod, and one row of spinules next to the basal setae. The endopod is 2 segmented and the exopod is 3 segmented. The proximal endopodal segment is unarmed except a row of outer margin setules. The distal segment integument has a row of inner setules, and small subterminal spinule. All exopodal segments have outer margin spinules and at least one row of interior spinules. The proximal exopodal segment has no inner margin setules. The second exopodal segment has a row of inner margin setules. The distal exopodal segment has 2 short row of interior spinules and a row of inner margin setules. Setal formulas are listed in Table 3.1. (Fig. 3.3.) The baseoendopod is very pronounced and is armed not only with a basal P5 seta but also 3 bipinnate setae. Two of these setae have strong spinules, and the other setae has only small setules. The baseoendopod also has 3 groups of setules on the inner margin, 2 groups of spinules on the outer margin, and a terminal row of lobate spinules. The exopod very pronounced and is as long as the baseoendopod. It has 3 bipinnate setae, all having strong spinules. It also has 2 groups of setules on the outer margin, 2 large spinules on the inner margin, and a subterminal row of fine setules. Description of the Male: Body. The shape is similar to the female. Second and third somites not fused. Antennule. (Fig. 3.4.) The shape is sub-chriocer, with 6 segments. Segment 1 has 3 row of spinules and one bipinnate setae on the distal outer margin. Segment 2 has 2 setae on the distal outer surface (one plain, one unipinnate), four setae on the anterior surface (2 pinnate, 2 plain), and one unipinnate setae on the posterior surface. Segment 3 has 8 plain setae along the anterior edge, and one unipinnate setae on the posterior surface. Segment 4 is swollen, with 6 plain setae along the anterior edge, 2 unipinnate setae along the anterior edge, one long plain setae on the posterior surface, one large aesthetasc (42 µm), and 5 small dorsal aesthetascs (12 µm to 17 µm). Segment 5 along the anterior margin has a strong spine and a plain setae. Segment 6 has the following segmentation: 2 plain setae along the anterior margin, 3 terminal plain setae, one terminal aesthetascs( 21 µm), 3 plain setae on the dorsal face (1 plain, one inserted on a projection of the integument, and one is very fine), 3 setae on the posterior face (all inserted on projections of the integument). P3• The shape is similar to the female except that the distal endopodal outer margin spinules are slightly thicker and longer. P5 (Fig. 3.4.) The baseoendopod is reduced as compared to the female. It is missing the proximal inner setae, 2 groups of spinules along the outer margin, and 3 groups of setae along the inner margin. However, it has retained the row of subterminal lobate spinules. Exopod similar to the female in the following respects: general shape, 2 sets of setules along the outer margin, a row of subterminal setules, and 2 distal large bipinnate setae. However, it lacks the proximal outer bipinnate setae found in the female. Caudal Rami. Similar to the female in shape and size. Variations. Variation in the female were not obvious in between the holotype and the 2 dissected paratypes. The antenna I exopod always had 3 setae in males and females. There was little sexual dimorphism of the body or the P3. Etymology. The generic name Serratithorus (L. 'Serrat' = saw shaped and L. 'thorus =thorn) refers to the dorsal rows of large socles, which are quite pronounced in the type specimen of the genera. The species name refers to Dr. Mike Gee who was instrumental in furthering my interest in harpacticoids. Discussion Remarks. The new species shares many taxonomically important characters with Cletodes and its sister group Enhydrosoma as they are discribed in Gee 1994,. Although the new species exhibits characteristics that are shared by the two genera, it also has characters that are used to distinguish between the two genera. It also has some characters that are not found in either genera. Serratithorus geei shares the following genera distinguishing characteristics with Cletodes: 2 bipinnate setae on the abexopodal margin of the allobasis of the A2, 5 setae on the mandibular palp, and a small endopodal lobe on the PS. It differs from Cletodes in the following important respects: 3 setae on the exopod of the A2, only one setae on the anterior surface of the arthrite, the distal segments of P3 and P4 both have 5 setae, and the caudal rami setae Ill is implanted medially on the outer margin. Serratithorus geei shares the following genera distinguishing characteristics with Enhydrosoma: S setae on the distal exopodal segment of P3 and P4, and seta Ill on the caudal rami is inserted medially on the outer margin of the ramus and seta VII is implanted near the base of the ramus as in some of the Ł. bucholtzi group (Gee 1994). S. geei also has an A2 exopodal segment which is shaped like a lowercase y. It differs from Enhydrosoma in the following respects according to the revision in Gee 1994: the lack of a bifid structure on the rostrum, 3 setae on the exopod of A2, 2 bipinnate setae on the abexopodal margin of the allobasis of the A2, 5 setae on the mandibular palp (most Enhydrosoma have 3), and the endopodal lobe is undistinguishable on the baseoendopod. Serratithorus New Genus A new genus is proposed to encompass the type species Serratithorus geei. The genus has several features which make it unique within the family Cletodidae: a large fused rostrum which is triangular shaped with a dorso-ventrally recurved tip which lacks a bifid structure. There are two dorsal rows of large socles along the free somites, the A2 has 3 setae on the exopod in both the male and the female, and has 2 bipinnate setae on the abexopodal margin, the lack of an inner setae on the exopod-2 of P1-P4 and also the endopodal lobe is indistinguishable from the baseoendopod. ' There are several features which are prominent, and quite unusual, in the type species. But, considering the variation seen in other genera of Cletodidae, I hesitate to propose that they occur across the genera. The type species has a PS on the female which bears 3 exopodal setae, and 3 setae on the baseoendopod. The male PS is reduced in size and only has 2 exopodal setae, and 2 setae on the baseoendopod. The cephalic shield in both sexes of the type species is very pronounced posteriorly to form a plate with associated socles along the posterior margin. A specimen described by Sewell (1940) as Enhydrosoma nicobarica shares several characteristics with Serratithorus geei. It was probably was placed in Enhydrosoma, because the distal segment P3 and P4 have S setae. At the time, this was the dominate characteristic separating Enhydrosoma and Cletodes. It also has an A2 with 3 setae on the exopod of the A2, as do some other Enhydrosoma copepod ites (Gee personal conversation). However, it also has a pointed recurved rostrum which is not bifid, and 2 setae on the abexopodal margin of the allobasis of the A2. These characters are not found in Enhydrosoma (Gee, 1994), and are included in the discription of the new genus Serratithorus. On this basis, I think that E. nicobarica should be moved to the new genus as Serratithorus nicobarica. The description of this species is somewhat confusing. Sewell's materials section lists the animal found as being a single female, in the description he mentions that the animal is not an adult and his drawings are of a S'th stage copepodite. The setation of PS of the animal closely resembles the male of S. geei. It differs from 5. geei in that it is missing a small inner seta on the Pl endopod 2, the setae of the endopods of P2-P4 are much shorter, the shape of the appendicular segments are different, and in the text Sewell states that some of the exopodal setae on P2 and P3 are modified to spines. The cephalic shield is not as pronounced as the type species. However, during a study in the Gulf of Mexico called GOOMEX, also for the Department of the Interior, I found unknown cletodid which displayed Serratithorus genera characters and also had a less pronounced cephalic shield. Acknowledgments I am much indebted to Dr. Paul Montagna, at the University of Texas Marine Science Institute, not only for the specimens and for use of the facilities, but also for his professional and technical support. I am grateful to Dr. Mike Gee, for his taxonomic prowess in this area. Table 3.1.: Setal formula for Serrithorus geei. LEG EXOPOD ENDOPOD P1 0.0.022 0.111 P2 0.0.022 0.020 P3 0.0.122 0.021 P4 0.0.122 0.021 Figure Legend Figure 3.1.: Serratithorus geei S?. Figure 3.2.: Serratithorus geei S? mouth parts. Figure 3.3.: Serratithorus geei S? periopods. Figure 3.4.: Serratithorus geei o. 47 ~ J DORSAL HABITUS 0 · 2rnrn CR GF CR 50 µm 50 µm 50 µm Mxl- Mx Mxp Md Mxl Mx Mxp 20 µm 50 µm sc­ segments . w/o setae 20 µm GF Ps 50 µm References Coull B. C. (1977) Marine flora and fauna of the northeastern United States. Copepoda: Harpacticoida. NOAA Tech. Report. NMFS Circ 399.4Bpp Gee M. (1994) Towards a revision of Enhydrosoma Boeck, 1872 (Harpacticoida: Cletodidae Sensu Par); A re-examination of the type species, E. Curticauda Boeck, 1972, and the establishment of Kollerua Gen. Nov. Sarsia, 79:83-107. Bergen. ISSN 0036-4827. Huys R., Gee J.M., Moore C.G.,and Hamond R. (In press) Marine and Brackish water harpacticoid copepods Part 1. Synopses of the British Fauna (Ed. R.S.K. Barnes & J.H. Crothers) No. 51. Field Studies Council Press UK, pp363 approx. Hyland J., D. Hardin, I. Crecelius, D. Drake, P. Montagna, and M. Steinhauer (1990) Monitoring long-term effects of offshore oil and gas development along the southern California outer shelf and slope: background environmental conditions in the Santa Maria Basin. Oil and Chemical Pollution, 6, 195-240. Lang, K. (1948). Mongraphie der Hrpacticiden, 1&11. Hakan Ohlsson, Lund. 1682 pp. Lang, K. (1965). Copepoda Harpacticoida from the California Pacific coast. Kung/. Svenska Vetensk. Hand/., Fjarde, 10(2): 1-560. Moore C. G. (1976) The form and significance of the hyaline frill in harpacticoid copepod taxonomy. J. Nat. Hist Sewell, R. B. S. (1940) Copepoda, Harpacticoida. Scientific Reports of the John Murray Expedition 1933-1934. British Musem (Nat. Hist.) 7 (2) :117-382. CHAPTER 4: THREE NEW HARPACTICOID COPEPODS FROM THE SANTA MARIA BASIN OFF THE CALIFORNIAN PACIFIC COAST (COPEPODA, HARPACTICIODA) (by Frank Fiers)1 1Koninkliik Beleisch lnstituur voor Natuurwetenschappen, Recent Invertebrate Section, Vautierstraat 29, B-1040 Brussels, Belgium Abstract Three new harpadicoid copepods from the Santa Maria Basin off the south-central Californian coast are described: Zosime Pacifica n. sp., C/etodes macrura n. sp. C/etodes tuberculatus n. sp. The three species are the most dominant harpadicoids found during a sampling program studying the possible effects of oil and gas developments in this area. The description of the copepodids of each species is included and a key to the species of the genus Cletodus is given. 54 Introduction In 1986 a sampling program began on the outer continental shelf in the Santa Maria Basin off the south-central coast of California, U.S.A. This is a pristine environment which is currently subject to future oil and gas development. The purpose of the sampling program was to determine if there were long-term impacts due to materials discharged during platform development and production (Hyland et. al., 1990). One aspect of the multidisciplinary program is to determine if there are reproductive or life history effects on harpacticoid copepod communities. Harpacticolds are usually more sensitive to hydrocarbon exposure than the dominant nematodes (Fricke et al., 1981; Bodin and Boucher, 1983; Hennig et al., 1983). During the course of this study we found over 100 species (Montagna, in preparation), most of them new to science. Three of the species were chosen for detailed life history analyses, and these are also new. These species were the dominant harpacticoids in the community (Montagna, in preparation). The details of the population dynamics are published elsewhere (Webb and Montagna, in preparation). The purpose of this paper is to describe these three new species. Materials and Methods The copepods described below were gathered during eight cruises to the Santa Maria Basin, off the Californian coast, between November 1986 and May 1989. Exact localities and sampling methods are described in detail in Hyland et al., 1990. Most of the specimens of the herein described copepod species are used for detailed life history analyses but a representative sampleof them isreserved as type material and deposited inthe collections of the Zoologisch Museum Amsterdam (ZMA Coll. no. Co.), the Invertebrate collection of the Koninklijk Belgisch lnstituut voor Natuurwetenschappen (Brussels: COP) and in the reference collection of Dr. P. Montagna (M.S.I, Port Aransas). Dissected specimens are mounted in lactophenol and the preserved ones are stored in 75 % ethanol. Drawings were made with the aid of a camera lucida on a microscope equiped with phase contrast. Terminology and abbreviations are used according to Lang (1965), and setal formulae of the copepodids are listed following the method of Humes and Ho (1969). Systematic Descriptions Family Tisbidae Stebbing 1910 Genus Zosime Boeck, 1872 Zosime Pacifica n. sp. Type-region. -Pacific Ocean, Santa Maria Basin, California (U.S.A.) between Punta San Luis and Purisma Point, at a depth ranging from 50 to 565 m. Type-material. -Holotype: one dissected female, mounted on three slides from station PJ 23­2 (ZMA coll. no. Co. 102.855); allotype: one dissected male, mounted on three slides from station PJ 7-1: (ZMA Coll. no. Co. 102.856); paratypes: dissected, 5 females (ZMA Coll. no. Co.102.857, COP 3113-3116; 1 male (ZMA Coll. no. Co.102.858); 2 Cop I (COP 3117­3118); 3 Cop 11 (COP 3119-3121 ); 1 Cop Ill (COP 3122); 2 Cop IV, females (COP 3123­3124); 1 Cop IV, male (COP 3177); 3 Cop V (males, COP 3125-3127); preserved: 27 females, 9 males, 15 Cop 1, 10 Cop 11, I Cop 1, 10 Cop 11, 16 Cop 111, 7 Cop IV (females), 4 Cop V (females) (ZMA Coll. nom 102.859 -102.862 and COP 3128-3137, M.5.1.). Etymology. -The specific name refers to the Pacific Ocean, type area of the present species. Description. Adults Female:Habitus (Fig.4.1. a, b) fusiform compressed in dorsal view. Length, including rostrum and furcal rami, 420 µm. Largest width near the posterior edge of the cephalothorax and the second thoracic segment (160 µm). Cephalothorax large, almost one third of the entire body length, smoothly bending anterior in dorsal view. Lateral margins of thoracic segments bending smoothly posteriad. Body constricted at the anterior margin of the first genital segment. Genital segments and second abdominal segment with posteriorly directed lateral extensions. Penultimate and anal segments with slightly rounded lateral margins. lntegumental structures. Entire surface of cephalothorax and pleurotergites covered with an irregular pattern of minute spinules (not illustrated). Posterior margin of cephalothorax smooth dorsally. Posterolateral margins and posteroventral edge of cephalothorax, as well as the thoracic pleurotergites furnished with irregularly formed blunt extensions (Fig. 4.1 e, d). Ventral and anterolateral margins of cephalothorax smooth. Median region of posterior margin of third and fourth thoracic segments with distinctly smaller and more rounded extensions. Posterior margin of first genital segment smooth, of the second one and abdominal ones set with sharper lips. Ventral surface of genital and abdominal segments smooth; posteroventral margin of second genital and second abdominal segments showing a wide hyaline frill, minutely incised. Posterior margin of third abdominal segment with, dorsally, a large hyaline frill, incised, covering the entire anal segment and, ventrally, set with long and slender spinules. Genital field situated close to the posteroventral margin of the first genital segment; clasping organ convex and bearing two long setae on each side. Furcal rami (Fig. 4.1. c) with nearly parallel margins and about twice as long as wide (45 µm x 21 µm). Lateral and dorsal setae implanted in the distal fourth, the latter on a small socle and articulating on two basal parts. Ventral seta implanted close to the distal margin. Inner apical seta small, less than half the length of the ramus. Outer and inner apical setae robust, the former about half as long as the latter. Both set with a dense pattern of small spinules around the stem. Surface of the rami spinulose. Outer margin with a few transversal rows of distinctly longer spinules. Rostrum (Fig. 4.2. a) strongly tapering anteriad. Rostral tip ventrally directed and strongly sclerotized. Rostral tip slightly protruding. Antennule (Fig. 4.2. b) six-segmented. First segment, bearing a single seta, set with strong spinules along the anterior margin. Aesthetascs implanted on the fourth (85 µ long) and sixth (39 µ long) segments. All setae setulose or spinulose, except for some small setae on the penultimate and ultimate segments. Antenna (Fig. 4.2. c, d) with large coxa. Basis cylindrical, bearing a seta along the inner side and a three-segmented exopodite. Surface of the basis having an oblique row of long and slender spinules. Exopodite articulating with the basis near the distal outer edge of the latter. Endopodite two-segmented. Proximal segment with a single, armed seta. Distal endopodal segment with three lateral setae (two of them strongly armed) and six distal ones. First and second exopodal segments with one seta, third exopodal segment small, about half as long as the preceding one, and third segment nearly twice as long as the first one. Mandible (Fig. 4.2. e) having a robust and strongly sclerotized gnathobasis. Biting edge with four rigid teeth, one seta and some spinules. Mandible palp with distinctly one segmented exopodite and endopodite, bearing three and four feathered setae respectively. Basis bearing three feathered setae, apically. Prae-coxa of maxillule (Fig. 4.2. f) large and triangular. Arthrite with two slender lateral setae and with eight strongly armed spines arising from distal margin. Coxa bearing two epipodal setae and four inner setae. Basis bearing six setae implanted on the inner extension. Exopodite and endopodite present, cylindrical and bearing three and six setae respectively. Maxilla (Fig. 4.2. h) having three endites. Proximal endite V-shaped, each branch bearing three setae. Median and distal endites cylindrical having three distal setae each. Basis hook-shaped, bearing a robust armed setae along with four slender and smooth ones. Two of the latter arising close to the articulation with the exopodite. Exopodal segment cylindrical, bearing five setae: three apical and two lateral ones. Maxilliped (Fig. 4.2. g) with a triangular coxa, having a tuft of slender spinules. Basis slightly protruded having a robust armed spine and some small spinules. Endopodite one segmented, furnished with two apical feathered setae and two smooth lateral ones. Prae-coxa and coxa of Pl (Fig. 4.3. a) furnished with rows of long spinules as in the other legs. Basis with an outer and an inner armed seta, both reaching beyond the first segments of the rami. Exopodite three-segmented, endopodite two-segmented. Exopodal and endopodal spines set with much longer spinules along one side of the stem. Inner setae of the rami setulose. Chaetotaxy of Pl in Table 4.1. P2 -P4 (Fig. 4.3. b, c, d, respectively) with rows of long spinules on protopodite surfaces. Outer seta of the basis striated and set with minute setules. Exopodites and endopodites three-segmented. Outer exopodal and endopodal spines symmetrically armed. Endopodites reaching towards the middle of the third exopodal segments of P2 and P3 and just beyond the articulation between the second and third exopodal segment in P4. All setae with a normal setulose appearance except for the dwarfed and smooth outer distal seta of the third endopodal segment of P4. Chaetotaxy of the legs in Table 4.1. PS (Fig. 4.2. i) exopodite fused with baseoendopodite, bearing three and two setae respectively. Outer baseoendopodal seta implanted on a long cylindrical extension. One smal I seta, implanted on a small elevation, arising near the outer proximal edge of the exopodal lobe. Male: habitus (Fig. 4.4. a) resembling the female but showing a more slender fifth thoracic and abdomen. Ventrolateral margins of the sixth segment strongly extended laterally. Antennule (Fig. 4.4. b) sub-chirocer, seven segmented. First segment with rigid spinules. Aesthetasc implanted on the fifth segment, distinctly wider than that of the female and about 95µm long. Penultimate and ultimate segment each with a blunt conical process. Apex of ultimate segment extended, forming a straight and sharp claw. Mouthparts and natatorial legs as in the female except for a distal endopodal segment of P2 bearing only one apical seta and an apical, curved, smooth hook-shaped processus (Fig. 4.4. c). P 5 (Fig. 4.4. d) fused with the supporting segment and with chaetotaxy as in the female but bearing considerably smaller setae; endopodal lobe not protruded, exopodite articulating and less large than in the female. P6 (Fig. 4.4. e) symmetrical. Both sides distinctly protruded and bearing two long spinulose setae and an inner dwarfed smooth one. Copepodids Copepodid I Habitus (Fig. 4.5. a, d) fusiform compressed in dorsal view. Body composed of five somites. Lateral margins of the cephalothorax nearly parallel. Length, including rostrum and furcal rami, 225 µm. Cephalothorax almost half as the entire body length. Second and fourth thoracic segments with rounded lateral margins, third thoracic segment with parallel margins. Anal segment with a convex anal operculum. Posterior margins of all the somites smooth. Ventral surface and posterolateral edges of the anal segment with long spinules (Fig. 4.5. c). Rostrum triangular, steeply bending anteriad. Rostral tip rigid and straight. Furcal rami twice as long as wide. Two lateral setae and dorsal seta implanted near the middle, one lateral seta arising from the distal outer edge and one seta implanted on the ventral surface. Principal setae fused over a short distance. Outer principal one armed with long spinules but striated near the tip. Inner principal seta set with minute spinules over the entire length. Antennule (Fig. 4.5. h) three-segmented, bearing an aesthetasc on the second and third segments. All setae spinulose except for some slender ones on the ultimate segment. Antenna as in the adult but less sclerotized. Exopodal setae smaller and less strongly armed (Fig. 4.5. f}. Mandible, maxillule, maxilla and maxilliped as in the adult. Pl (Fig. 4.5. b) and P2 (Fig. 4.5. g) protopodites with distinct prae-coxa, coxa and basis, all furnished with spinules. Bases with an outer spine. Pl without inner spine on the basis. Rami one segmented. Chaetotaxy in Table 4.2. P3 (Fig. 4.5. e) represented as an ovate plate, bearing three setulose setae. Cope pod id 11 Habitus (Fig. 4.6. a) 250 µm long. Body with six somites. Cephalothorax with curved lateral margins and bending strongly anteriad. Integument and posterior margins of the somites smooth except for the spinules arising along the posterior edges and median ventral surface of the anal segment. Rostrum as in the preceding stage. Furcal rami twice as long as wide. Two lateral setae and the dorsal one implanted near the middle and one ventral seta arising near the distal edge. Outer principal seta shorter than the ramus. Inner principal and inner apical setae as in the adult. Antennule (Fig. 4.6. e) four-segmented, bearing an aesthetasc on the second and ultimate segments. Appearance of the setae as in Cop 1. Antenna and mouthparts as in the adult. Pl (Fig. 4.6. b) and P2 (Fig. 4.6. f) with protopodites as in the adult. Rami two­segmented, chaetotaxy in Table 4.2. Basis of Pl with an inner spine, reaching almost to the apical edge of the endopodite. P3 (Fig. 4.6. g) with smooth protopodal parts and one-segmented rami. Chaetotaxy in Table II. P4 (Fig. 4.6. d) represented as a small ovate socle, bearing three setulose setae. Cope pod id 111 Habitus (Fig. 4.7. a) resembling the adult facies closely except for the smooth integument and posterior margins of the thoracic segments. Body composed of seven som ites. Length 360µm. Posterior margins of the fifth and sixth segments set with irregularly rounded extensions. Anal operculum convex and furcal rami as in the adult. Antennule (Fig. 4.8. e) four­segmented. Aesthetascs arising from the second and ultimate segments. Antenna with a three segmented exopodite with an adult appearance (Fig. 4.8. d). Other mouthparts as in the adult. Pl-P3 (Fig. 4.7. b, c, e, respectively) with distinct protopodal components and two segmented rami. Shape and armature of the exopodal spines as in the adult. Chaetotaxy in Table 4.2. P4 (Fig. 4.7. f) with distinct coxa and basis. Exopodite and endopodite one­segmented with chaetotaxy as listed in Table 4.2. PS (Fig. 4.7. d) represented as a slight elevation, bearing three setae, the outermost feathered and longer than the two inner smooth ones. Cope pod id IV (female) Body facies as in the adult, composed of eight segments (Fig. 4.9. a). Length 380 µm. Antennule (Fig. 4.8. f) five-segmented with aesthetascs on the second and ultimate segment. Mouthparts as in the preceding stage. P 1-P4 (Fig. 4.8. a, b, c) with two-segmented rami. lntegumental structures more dense than in copepodid Ill. P3 almost identical with P2, differs only by the slightly shorter endopodite. Chaetotaxy in Table 4.2. PS (Fig. 4.9. b) with a distinct inner baseoendopodal lobe (with two setae), a distinct exopodal lobe (with three setae) and a small socle, bearing one seta, near the outer proximal edge of the exopodal lobe. Baseoendopodite still fused with the supporting segment. P6 (Fig. 4.9. g) represented as an oblong ovate plate, bearing one long spinulose seta and a small rounded process near the outer distal margin. Copepodid IV male. Habitus, length, mouthparts and natatorial legs as in the female. Antennule differing from that of the female by a small rounded process on the apicalmost edge of the last segment (Fig. 4.8. g). PS (Fig. 4.9. c) with a less differentiated exopodal lobe, bearing three setae and much smaller endopodal setae. P6 (Fig. 4.9. h) as in the female but without an invagination between the opposite legs. Cope pod id V (female) Body with nine segments. General appearance more slender than in the adult (fig. 4.9. f). Free genital segments. Length 400µm. Antennule, mouthparts and legs as in the adult. PS (Fig. 4.9. d) almost identical with that of the adult but still fused with the supporting segment and showing shorter setae. P6 (Fig. 4.9. i) represented as a slightly elevated socle along the posteroventral margin of the segment, bearing three minute setae. Cope pod id V (male) Habitus and length as in the female copepodid V. Antennule (Fig. 4.8. h) six segmented with the aesthetasc on the second segment. Ultimate segment slightly prolonged, forming a rather thick but sharp extension. Mouthparts and natatorial legs as in the adult. P3 endopodite as in the female copepodid V, not dimorphic. PS (Fig. 4.9. e) as in the preceding stage except for the larger exopodal lobe and the slightly longer endopodal setae. P6 (Fig. 4.9. j) represented as a small elevation on the outer ventral edge of the segment and bearing three setae. Outer and median seta spinulose, inner one smooth and short. Discussion. -Zosime pacifica n.sp. is unique among the species of the genus because of the remarkably reduced apical seta on the endopodite of the P4. Although several species of the genus Zosime are described without illustrations of the natatorial legs, a comparable reduction of a seta has never been mentioned. Moreover, species discrimination within the genus is mainly based on the chaetotaxy of the legs. It seems obvious that if reduced setae were present they would have been observed. Within the genus, Z. pacifica n. sp. resembles most closely Z. valida G.O. Sars, 1919. Both species possess a PS with only two inner baseoendopodal setae which discriminates them from all other members of the genus. However, Z. pacifica n. sp. is easily distinguishable from its congener by the above mentioned reduced seta on the P4 and by the chaetotaxy of the Pl endopodite, bearing four setae/spines on the second segment instead of three as in Z. valida. Family Cletodidae T. Scott, 1904 Genus Cletodes Brady, 1872 Including the two herein described species, the genus Cletodes comprises 21 different species. Since the compilation of the most recent key (Hamond, 1973) four species, Cl. reductus Moore, 1978 Cl. dorae Por, 1979, Cl. selosus Marinov & Apostolov, 198S and Cl. endopodita (Schriever, 1984) were added. A revised key, discriminating females as well as males, is given below. Cl. brucel T&A Scott, 1901 and Cletodes sp. (Krishnaswamy, 19S7) are omitted. The male characteristics of Cl. millerorum Hamond, 1973 (untransformed endopodite P3, baseoendopodite PS without setae, exopodite PS with three setae) are included, based on specimens found in the present collection. The description of the male will be given in detail elsewhere. In the present key the baseoendopodite of the female PS of Cl. yotabis is considered to have three setae instead of four as originally stated by Por (1967) and listed by Hamond (1973: Table 38, p. 480), later on in his revision of the genus. The position of the inner distal seta and the statement that this seta is reduced (Por, op cit.) clearly indicates that this structure represents a hyaline tubular pore identical with those discussed by Bodin (1970) for Cl. limicola Brady, 1872 and Cl. tenuipes T. Scott, 1896. Key to the species 1 -P3 exo 2 without, P4 exo 2 with an innerseta .......................... 2 -P3 exo2 and P4 exo 2 without inner seta ............................. S -P3 exo2 and P4 exo 2 with an innerseta ............................. 9 2 -Furcal rami rather short, about as long as the anal segment.. . . . . . . . Cl. setosus Marinov & Apostolov, 198S -Furcal rami much longer, at least twice as long as the anal segments . . . . . . . . . 3 3 -P2 end 2 with one apical seta only; exo PS three times as long as wide in the female, 2.S times in the male . . . . . . . . . Cl. tuberculatus n. sp. -P2 end 2 with two setae, inner one slender; exo PS with different lengths. . . . . . 4 4 -Furcal rami strongly tapering posteriad; female and male exo PS twice as long as wide ................................. Cl. reductus Moore 1978 -Furcal rami cyl in_drical; female and male exo PS 3-3.S times as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. smirnovi Bodin 1970 S -Baseo PS of females and exo PS of males with three setae/spines; P3 end of males two segmented without apophysis ................................... 6 -Baseo PS of females with one, exo PS of males with four setae/spines; P3 end of males three-segmented with apophysis ................................. 7 6 -Furcal rami elongated ovate (UW: 3/1 ); male end P3 with a large spine and seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. dorae Por 1 9 79 -Furcal rami broadly ovate (UW: 2/1 ); male end P3 with a small and a long seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. hartmannae Lang 196S 7 -P2 exo 2 with an inner seta . . . . . . . . . . . . . . . . . . . . . . . Cl. pusillus Sars 1920 -P2 exo 2 without inner seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8 -P4 end 2 with two setae; P4 end 1 small,less high than wide .............................. Cl. endopodita Schriever 1984 -P4 end 2 with one seta; P4 end 1 as long as wide . . . . Cl. tenuipes T. Scott 1896 9 -P2 end 2 and P4 end 2 with two setae . . . . . . . . . . . . . Cl. longifurca Lang 1948 -P2 end 2 and P4 end 2 with four setae . . . . . . . . . Cl. latirostris Drzycimski 1967 -P2 end 2 with two setae, P4 end 2 with three setae .................... 10 -P2 end 2 with two setae, P4 end 2 with four setae ..................... 13 1O -Female PS exo with four setae, male unknown . . Cl. carthaginiensis Monard 193S -Female PS exo with five setae, male known . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11 -Female baseo PS with two setae; furcal rami nearly cylindrical; male P3 end transformed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. reyssi Sayer 1 964 -These characteristics not combined ................................ 12 12 -Furcal rami ovate, at the most three times as long as wide; male PS exo with three setae ................................... Cl. limicola Brady 1872 -Furcal rami nearly cylindrical, slightly tapering posteriad, at least six times as long as wide; male PS exo with two setae (lateral one absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. longicaudatus Boeck 1872 1 3 -Furcal rami not ovate in dorsal view, at least five times as long as wide; female P3 end 2 with four setae; male end P3 transformed ........................... 14 -Furcal rami ovate in dorsal view about three times as long as wide; female P3 end 2 with three setae; male end P3 not transformed ...................... 1 S 14 -Outer subdistal spine of P3 and P4 end 2 implanted near the distal edge; Pl end reaching to the middle of Pl exo 3; male PS with 2 baseoendopodal smooth setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. macrura n. sp. -Outer subdistal spine of P3 and P4 end 2 implanted far below distal edge; Pl end reaching to the distal margin of Pl exo 3; male baseoendopodite PS without setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. yotabis Por 1 96 7 1 S -Inner seta of female PS baseoendopodite setulose; furcal rami equally in both sexes ................................... Cl. spinulipis Por 1967 -Inner seta of female PS baseoendopodite stout, spinulose and rather short; furcal rami differing in both sexes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16 -Largest width of the female furcal ramisituated in the anterior half . . . . . . . . . . 17 -Largest width of the female furcal rami situated in the posterior half (male unknown) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cl. dissimillis Willey 193S 1 7 -Proximal outer spine of P4 exo 3 much shorter than subdistal one; female furcal rami as long as the anal segment . . . . . . . . . . . . . . Cl. pseudodissimilis Coull 1971 -Proximal outer spine of P4 exo 3 only slightly shorter than subdistal one; female furcal rami longer than the anal segment . . . . . . . . . . . Cl. millerorum Hamond 1973 Cletodes macrura n. sp. Type-region. ·_ Pacific Ocean, Santa Maria Basin, California (U.S.A.) between Punta San Luis and Purisma Point, at a depth ranging from 50 to 565 m. Type material. -Holotype: one dissected female, mounted on three slides from station PJ 7-2 (ZMA Coll. no. Co. 102.863); allotype: one dissected male, mounted on two slides from station PJ 7-2; (ZMA Coll. no. Co.102.864); paratypes: dissected, 2 females (M.S.I.); 1 male (M.S.I.); 2 Cop I (COP 3169, 3173); 1 Cop 11 (COP 3174); 1 Cop Ill (COP 317S); 1 Cop IV (COP 3171 ); 2 Cop V (COP 3170, 3172); preserved: 12 females, 17 males, 9 Cop 11, 7 Cop 111, 6 Cop IV (females), I Cop V (female) (ZMA Coll. no. Co.102.86S 102.866, COP 31S7-3168 and M.S.I.). Etymology. -The specific name macrura, a conjunction of the Greek words makros (long) and aura (tail), refers to the long furcal rami. Description. Adults Female: Habitus (Fig. 4.1 O. a, b) fusiform, slightly tapering posteriad. Length, including rostrum and furcal rami, SSO µm. Head rather small, about one fifth of the entire body length. Cephalothorax, in dorsal view, with rounded lateral margins and a distinct constriction in the posterior half. Body segments constricted in the anterior half and widening with smoothly curved lateral margins posteriad. Fifth thoracic and genital segments markedly extended laterally. Pre-anal and anal segment cylindrical. Anal operculum straight, lateral edges shaped by conical structures bearing the sensillae. Integument of thoracic and abdominal pleurotergites clothed with an irregular pattern of minute spinules, that of the head furnished with an irregular pattern of fine lines. Posterior margins of the thoracic pleurotergites smooth. Pleural region of the genital segments and anal segments set with some spinules. Posteroventral margin of the second and third abdominal segments spinulose (Fig. 11 a). Ventral surface of the genital segments smooth. Gen ital field strongly sclerotized, bearing a single seta on each side. Rostrum (Fig. 4.11. c) with a slightly curved rostral tip, set with fragile hairs. Furcal rami 5.5 times as long as wide, tapering towards the distal edge. Inner margin slightly undulating. Dorsal seta implanted in the posterior half. Two lateral setae arising almost medially and one subdistally. Outer principal seta small, as long as the inner apical one and fused with the principal one. Surface of the rami spinulose. Antennule (Fig. 4.11. h) five-segmented, showing long spinules on the first segment. Most setae on the first to fourth segments setulose. Ultimate segment with strongly armed spines and some smooth setae. Aesthetascs implanted on the third and fifth segments. Antenna (Fig. 4.11. j) with allobasis, having a transversal ridge beyond the articulation with the exopodite and one subdistal seta. Exopodite small, slightly longer than wide, bearing a single seta. Endopodite with three lateral and six distal spines/setae. Mandible (Fig. 4.11. k) with a large, strongly sclerotized gnathobasis, bearing two setae and three strong teeth. Palp with vestigial endopodite represented as a single seta. Exopodal lobe distinct but fused, bearing four setae. Apical seta of the palp distinctly stronger than the other ones. Arthrite of the maxillule (Fig. 4.11. i) with two setae and five unarmed curved spines. Coxa with two slender setae and basis with six apical and five lateral setae. Maxilla (Fig. 4.11. f) large and compact. Syncoxa strongly sclerotized, having three endites. Proximal endite represented as a slender seta, median and distal endites each with two armed and one smooth seta. Basis with three setae and typically hook-shaped. Unarmed. Maxilliped (Fig. 4.11. g) small, less high than the maxilla. Basis with a single seta. Endopodal segment set with strong spinules along the inner margin. Claw unarmed and bearing a long seta. Pl-P4 (Fig. 4.12. b, c, e, g, respectively) with three-segmented exopodites and two segmented endopodites. Coxae and bases with spinules. Bases of P2-P4 laterally prolonged. Basis of the Pl with an inner seta reaching far beyond the endopodite. Third exopodal segment of the P4 with an invagination along the inner margin, probably representing a pore orifice. Chaetotaxy of the legs in Table 4.3. PS (Fig. 4.12. d) Baseoendopodite with a long cylindrical outer extension, bearing the outer seta. Endopodal lobe of the bascoendopodite protruded, having three setae. Hyaline pore not observed. Exopodite with nearly parallel margins, six times as long as wide and bearing five setae. Male: Habitus (Fig. 4.10. c) resembling the female closely but more slender and slightly more tapering posteriorly. Length 480 pm. lntegumental structures as in the female except for an additional row of spinules along the posteroventral margin of the first abdominal segment (Fig. 4. 11. b). Furcal rami as in the female only slightly more slender. Antennulae (Fig. 4.11. d, e) six-segmented, sub-chirocer. First and second segments as in the female. Ultimate segments long, nearly cylindrical. Fourth segment with a curved row of long spinules on the dorsally directed surface. P3 (Fig. 4.12. a) with a three-segmented endopodite, earring a long, 5-shaped apophysis on the second segment and only two setae on the terminal segment. PS (Fig. 4.12. f) with a small but distinct endopodal lobe on the baseoendopodite, bearing two smooth setae. Inner seta rigid, outer one slender. Exopodite 3.5 times as long as wide and having four setae. P6 (Fig. 4. 11. b) represented as an oblong plate, without setae. Variability. -The holotype specimen lacks the continuous row of spinules along the dorsal border of the pre-anal segment. Copepodids Copepod id I _ Habitus (Fig. 4.13. a) tapering strongly posteriad. Body composed of five segments. Length of cephalothorax more than one third of the entire body length. Largest width near the posterior margin of the head. Anal segment constricted anteriorly, having a convex and setulose anal operculum. Length, including rostrum and furcal rami, 155 pm. Dorsal integument of the segments spinulose. Anal segment with two transversal rows of long rigid spinules ventrally (Fig. 4.13. b, c). Furcal rami (Fig. 4.13. a, b, c) three times as long as wide, slightly convex in the anterior half of the inner margin. Dorsal seta arising in the middle, close to the inner margin. Two lateral setae implanted in the anterior half and one lateral seta implanted in the posterior third. Second dorsal seta near the distal margin. Principal distal setae fused, the outer one as long as the ramus. Outer distal edge of each ramus showing a well defined tubular pore. Rostrum with smoothly bent lateral margins and a straight anterior tip. Antennule (Fig. 4.13. f) three-segmented, bearing an aesthetasc on the second and third segments. Most setae spinulose. Antenna (Fig. 4.13. g) resembling the adult shape closely but having a larger exopodite, bearing two setae. Mouthparts as in the adult. Pl-P2 (Fig. 4.13. d, e, respectively) with distinct protopodal components. Pl basis without inner seta. Endopodites and exopodites one-segmented, chaetotaxy listed in Table 4.4. P3 (Fig. 4.13. a) represented as a distinct socle, bearing three equal setae. Cope pod id II Habitus (Fig. 4.13. h) tapering. Cephalothorax with strongly folded posterolateral surface. Thoracic segments 1-3 sixth rigid integumental elevations dorsally. Length, 21 O µm. lntegumental structures as in copepodid I. Furcal rami nearly five times as long as wide, slowly tapering posteriad. Dorsal seta implanted in the middle of the rami. Proximal lateral setae as in Cop I but distal lateral seta implanted closer to the edge of the rami. Outer principal seta and inner distal seta small and smooth. Antennule (Fig. 4.13. 1) four-segmented, bearing the aesthetascs on the second and fourth segments. Exopodite of antenna (Fig. 4.13. g) small, having only one feathered sets. P1-P2 (Fig. 4.13. i, j, respectively) with two segmented rami. Basis of Pl with an inner seta, reaching beyond the distal edge of the second endopodal segment. P3 (Fig. 4.13. k) with distinct protopodal components and one-segmented rami. Chaetotaxy of the legs in Table 4.4. P4 (Fig. 4.13. h) represented as three setae on a distinct lobe. Copepodid Ill Body (Fig. 4.14. a) seven-segmented with facies resembling the preceding stage. Length 280 µm. Furcal rami as in copepodid II. Antennule (Fig. 4.14. e) four-segmented with an aesthetasc on the second and ultimate segments. Antennule and mouthparts as in the adult. P1-P3 (Fig. 4.14. b, c, d, respectively) with distinct protopodal components and two segmented rami. Chaetotaxy listed in Table 4.4. P4 (Fig. 4. 14. f) with prae-coxa, coxa and basis. Exopodite and endopodite one­segmented. Chaetotaxy in Table 4.4. PS (Fig. 4.14. a) represented as an elevation bearing three setae. Copepod id IV (female) Habitus (Fig. 4.14. g) resembling the adult body shape closely except for the smaller length (3SO µm) and fewer body segments (eight). Antennule (Fig. 4.1 S. f) five-segmented with the aesthetascs on the third and last segments. General appearance as in the adult. Pl -P4 (Fig. 4.1 S. a, b, c, respectively) with two segmented rami. lntegumental structures and shape of spines resembling closely the adult form. PS (Fig. 4.1 S. e) with a slightly protruded baseoendopodite, still fused with the supporting segment, and bearing two endopodal setae and one outer seta arising from a long cylindrical extension. Exopodal lobe distinct, about twice as long as wide and having five setae. P6 not differentiated. Male copepodid IV not found. Cope pod id V (female) Habitus as in the preceding stage but 480 µm. long and composed of nine segments. Antennule and mouthparts as in the adult. P 1 -P4 with three-segmented exopodites and endopodites smaller than in the adult legs. Chaetotaxy in Table 4.S. PS (Fig. 4.1 S. h) still fused with the body segment. Inner baseoendopodal lobe with three setae. Exopodite oblong, nearly three times as long as wide and bearing five setae. P6 not differentiated. Copepod id V (male) Habitus, length and mouthparts as in the female. Antennule (Fig. 4.1 S. j) four­segmented with aesthetasc on the third and last segments. Last segment long and slightly extended with a rounded process distally. P3 (Fig. 4.1 S. j) with protopodite and exopodite as in the female copepodid V. Endopodite two segmented bearing two apical setae, one inner, minute seta and a large, robust, outer subdistal spine on the second segment. Frontal surface of the second segment showing a sclerotized transversal ridge near the implantation of the outer subdistal spine. PS (Fig. 4.1 S. g) bearing two baseoendopodal setae and four exopodal ones. Exopodite about 2.S times as long as wide and still fused with the bascoendopodite. P6 not differentiated. Discussion. -Cl. macrura n. sp. resembles Cl. yotabis Por, 1967 in many aspects. Chaetotaxy of the legs and shape of the, furcal rami are nearly identical in both species. The here described species differs from the latter by the more conventional implantation of the outer subdistal spines on the endopodites in P3 and P4. Whereas in Cl. yotabis these spines arise near the middle of the outer margin of the segment, opposite the implantation of the inner lateral seta, the subdistal outer endopodal spines in Cl. macrura n. sp. are implanted just below the outer edge of the segment. The males of both species differ markedly in chaetotaxy of their PS. The exopodite of this leg in Cl. macrura bears four setae, while the baseoendopodite has a distinct inner lobe and carries two long smooth setae. In Cl. yotabis the male PS lacks baseoendopodal setae and has only three setae on the exopodite. Other differences between both species are the body length (Cl. macrura: SSO µm, Cl. yotabis: 420-480), the strongly developed maxillule and the smooth anal operculum in Cl. macrura. Cletodes tuberculatus n.sp. Type-region. -Pacific Ocean, Santa Mari Basin, California (U.S.A.) between punta Sa Luis and Purisma Point, at a depth ranging between 50 to 565 m. Type-material. -Holotype: one dissected female, mounted on two slides from station PJ 7-1 (ZMA Coll. no. Co. 102.867); allotype: one dissected male, mounted on two slides from station PJ 7-1: (ZMA Coll. no. Co.102.868); paratypes: dissected, 7 females (ZMA Coll. no. Co.102.869, M.S.I.); 2 males (M.S.I.); 3 Cop I (COP 3141-3142, 3144); 1 Cop 11 (COP 3143); 1 Cop Ill (COP 3138); 1 Cop IV (COP 3140); 1 Cop V (COP 313 9); preserved: 31 females, 13 males, 10 Cop 1, 10 Cop 11, 7 Cop Ill, 1 Cop IV (female), 2 Cop V (females) (ZMA Coll. no. Co.102.870-102.871, COP 3145-3156 and M.S.I.) Etymology. -The specific name refers to the small but distinct humps (Latin, luberculum) along the posterior margins of the segments, bearing the sensillae. Description. Adults Female: Habitus (Fig. 4.16. a, b) cylindrical with largest width near the posterior margin of the cephalothorax (100 µm). Length, including rostrum and furcal rami, 440 µm. Length of cephalothorax about one fifth of body-length. Cephalothorax slightly constricted in the posterior half. Anterior part of cephalothorax strongly tapering towards the rostrum. Thoractic and abdominal segments with curved lateral margins. Pleural regions strongly convex in the thoracic segments, and posterolateral extended in the first and second abdominal segments. Sensillae on posterior margins of the segments arising from a small cylindrical tubercle. Integument of all somites strongly sclerotized, forming distinct curved internal bands and rigid edges. Integument of cephalothorax forming a dense pattern of irregular pits (see Fig. 4.16. a), interrupted by longitudinal smooth bands. Posterior margins of cephalothorax and thoracic segments smooth, except for a few fragile hairs along the posterolateral margin of Th 5. Posterodorsal margins of the genital and abdominal segments smooth, lateral margins spinulose or hairy. Ventrally, posterior margin of genital and second abdominal segments set with spinules. Anal operculum slightly convex and furnished with teeth along the edges. Rostrum (Fig. 4.17. g) strongly tapering anteriad and directed downwards. Rostral tip not protruded. Dorsal integument reticulated. Furcal rami slightly curved, tapering posteriad and about, 5.5 times as long as wide. Inner proximal edge forming a blunt extension. Dorsal seta, articulating on two basal parts, and two lateral setae arising in the anterior third of the rami. Third lateral seta implanted in the posterior third. Outer and inner principal setae fused near their basis. Inner apical seta small, Antennule (Fig. 4.17. b) five-segmented, furnished with rigid spine-like setae. Aesthetascs implanted on the third and ultimate segment. Integument of the segments smooth except for some rows of sharp spinules on the first and second one. Antenna (Fig. 4.17. 1) with allobasis and a one segmented cylindrical exopodite, bearing a single long and feathered seta. Inner seta of the allobasis arising in the apical half of the segment. Endopodal segment furnished with strong spinules and having three lateral and six apical setae/spines in al I. Mandible (Fig. 4.17. i) with slender gnathobasis, bearing three long teeth and a single seta apically. Pars molaris situated in the outer half of the gnathobasis. Mandibular palp having two apical and two lateral setae. Rami obsolete. Maxillulae (Fig. 4.17. j) with four strong spines and two setae on the arthrite. Coxa with two setae. Basis cylindrical, bearing eight lateral setae, representing the vestigial rami and two apical setae. Maxilla (Fig. 4.17. k) small, only slightly higher than the preceding appendage. Two endites, both with three setae. Basis hook-shaped, furnished with two lateral and two apical setae. Maxilliped (Fig. 4.17. h) with a single seta and furnished with several rows of spinules. Inner margin of endopodal segment and claw armed. Pl-P4 (Fig. 4.18. a, b, c, d, respectively) bases slightly extended externally, bearing a feathered outer seta. Pl basis with an inner seta, reaching almost to the apical end of the endopodite. Exopodites three-segmented, endopodites two-segmented. First endopodal small, at the most as high as wide. Second endopodal segments long and slender, all densely clothed with long spinules. Chaetotaxy of the legs in Table 4.S. PS (Fig. 4.18. f} without protruded endopodal lobe. Exopodite 3.S times as long as wide and bearing five setae. Baseeoendopodite with a long outer cylindrical extension bearing the outer setae. Endopodite represented as a single feathered seta. Male: Habitus (Fig. 4.16. c) as in the female but more tapering posteriad and without fused genital segments. Length as in the female. Posteroventral margins of the abdominal segments set with long spinules. Antennule (Fig. 4.17. e) six-segmented, subchirocer. Aesthetasc implanted on the fourth segment. Ultimate segment prolonged, forming a long curved claw. Ventrally directed surface of the fourth segment smooth but dorsally directed surface armed with a comb of rigid spinules (Fig. 4.17. d). P3 (Fig. 4.18. e) . with a three-segmented endopodite, showing a distinct apophysis on the inner distal edge of the median segment. Protodite and exopodite as in the female. PS (Fig. 4.18. g) without endopodal seta. Exopodite slightly longer than double the width and having four setae. P6 (Fig. 4.17. a) strongly asymmetrical. Right leg irregularly ovate, left leg only presented by a chitinized strip. Both legs without setae. Variability. -Some specimens were found showing a misbuilt furcal ramus. As illustrated in Fig. 4.17. f, such rami are short, only three times as long as wide, and having much longer apical setae. Copepodids Remarks: The body facies of the copepodids resemble closely that of the adult, except however for the number of body segments. As such, they are illustrated here only for the first and second copepodids. Copepodid I Body length (Fig. 4.19. a) 175 µm. Body composed of five segments. Largest width near the posterior end of the cephalothorax. Length of the head almost one third of the entire body length. Integument clothed with minute spinules. Ventral surface of the anal segment with a transversal row of long spinules. Anal segment smooth, slightly convex. Furcal rami with an adult appearance but with two dorsal setae (one in the proximal half and one near the distal edge) and a long outer principal seta. Inner apical seta absent. Antennule (Fig. 4.19. k) three-segmented with aesthetascs on the second and third segments. Antenna (Fig. 4.19. c) as in the adult but with a larger exopodite, bearing two setae. Mouthparts as in the adults. P1-P2 (Fig. 4.19. d, e, respectively) with differentiated protopodites and one­segmented rami. Chaetotaxy in Table 4.6. Inner seta of the Pl basis absent. P3 (Fig. 4.19. b) represented as a distinct socle bearing three setae. Copepod id 11 Habitus (Fig. 4.19. f) resembling closely the preceding stage but composed of six segments. Anal operculum set with a transversal row of spinules. Length 215 µm. Furcal rami as in the adult, having one dorsal, three lateral and three apical ones. Antennule (Fig. 4.19. I) four-segmented. Aesthetascs implanted on the second and fourth segments. Antenna (Fig. 4.19. g) with a small exopodal segment, having only one seta. P1-P2 (Fig. 4.19. h, i, respectively) with two-segmented rami. Basis of P1 with an inner seta, as long as the entire endopodite. P3 (Fig. 4.19. j) with distinct protopodal components and one-segmented rami. Chaetotaxy of the legs in Table 4.6. P4 as the P3 in the copepodid II Copepod id Ill Habitus as in the preceding stage but with seven segments. Length 260 µm. Antennule (Fig. 20 g) four-segmented as in the preceding stage but with a longer second segment and more setae on the latter. P1 -P3 (Fig. 4.20. a, b, c, respectively) with two-segmented rami. P4 (Fig. 20 d) with one-segmented rami. PS (Fig. 4.20. h) present, represented as a small socle with three setae. Cope pod id IV (female) Habitus resembling closely the adult facies, with eight segments. Length 300 µm. Antennule (Fig. 4.20. k) five-segmented with aesthetascs on the third and fifth segments. General appearance as in the adult but with smaller segments. Pl-P4 having two-segmented rami. P2-P4 with the same morphology (Fig. 4.20. e, f). Chaetotaxy in Table 4.6. PS (Fig. 4.20. i) with a distinct exopodal lobe, bearing three setae. Baseoendopodite with an outer seta implanted on a long cylindrical extension and with one inner seta. P6 not present. Male copepodid IV not found. Copepodid V (female) Habitus as in the adult but with only nine segments. Length 3SO µm. Antennule, mouthparts, as well as legs 1-4, as in the adult. The latter with three segmented exopodites and two-segmented endopodites. Chaetotaxy in Table 4.6. PS (Fig. 4.20. k) as in the copepodid IV, somewhat larger and bearing five exopodal setae and a single inner baseoendopodal one. P6 not present. Male copepodid V not found. Discussion. -It is apparent that Cl. tuberculatus n. sp. is most closely related to Cl. smirnovi Bodin, 1970 and C. reductus Moore, 1978. These three species share a common chaetotaxy of the exopodites with only one inner seta on the median exopodal segment of the P4. Cl. tuberculatus n. sp. differs from its congeners by the more reduced chaetotaxy of the endopodite P2, bearing only one seta and the shape of the furcal rami, showing a distinct process on the inner proximal edge. Furthermore, the here described species exhibits considerably smaller endopodites in P1 and P2. In Cl. reductus and Cl. smirnovi, the endopodal ramus of Pl reaches almost to the distal margin of the exopodite, whereas the endopodite of the P2 is as long as the two proximal exopodal segments. In Cl. tuberculatus n. sp. the endopodites reach only halfway the third exopodal segment in Pl and only halfway the second exopodal segment in P2. Males of these closely related species are distinguishable mainly by the chaetotaxy and shape of the PS. Whereas Cl. reductus and Cl. smirnovi bear three setae on the rather short exopodal ramus, Cl. tuberculatus n. sp. bears a much longer exopodite furnished with four feathered setae. 71 Acknowledgements I am much indebted to Dr. Paul Montagna (M.S.1, Port Aransas) for the facilities he offered me in studying this material and for the hospitality during my stay at the Marine Science Institute, Port Aransas. I am grateful to Mine V. Arkosi for reading the manuscript. Part of this study was performed as a Postdoctoral Fellow at The University of Texas at Austin, Marine Science Institute, Port Aransas, Texas, USA. Table 4.1.: Chaetotaxy of Zosime Pacifica n. sp. P1 P2 P3 P4 PS~ PSo exo 0-1-123 1-1-123 1-1-223 1-1-223 4 3 end 1-121 1-1-121 1-1-121 1-1-121 2+1 2+1 73 legs Cop I Cop II Cop 111 Cop IV Cop V Table 4.2.: Chaetotaxy of the copepodids of Zosime pacifica n. sp. Pl exo 1, 11,IV 0,1-2,11,11 0,1-2,11,111 0I 1-2I 11, 111 0I I-1I I-1I 11, 111 end 2,1,I 1,0-1, 1, 1 1,0-1,1,1 1,0-1,2,1 1,0-1,2,1 P2 exo 1,11,111 0,1-2,ll,ll 1,1-3,11,111 1I 1-3I11, Ill 1,l-1,l-3,11,111 end 2,2, 1 1,0-1,2,1 1,0-2,2, 1 1,0-2,2,1 1I 0-1 I 0-1 I 2I 1 P3 exo 3 1,11,111 1,1-2,11,11 1,1-3,11,111 1,1-1,1-3, 11,111 end 2,2,1 1,0-1,2, 1 1,0-2,2,1 1,0-1,0-1,2, 1 P4 exo 3 1,11,111 0I1-3I 11 I Ill 1,1-1,1-3, 11,111 end 2,2, 1 1,0-2,2,1 1I 0-1 I 0-1 I 2I 1 PS exo 3 3 3 bas 2 +I* 2 +I* P6 1 3 2 setae on the endopodal lobe + 1 near the basis of the exopodite. Table 4.3: Chaetotaxy of Cletodus macrura n. sp. P1 P2 P3 P4 PSS? P6o exo 0-0-022 0-1-022 0-1-022 0-1-022 5 4 end 0-111 0-020 0-1 21 0-1 21 3 2 75 Table 4.4.: Chaetotaxy of the copepodids of Cletodes macrura n. sp. legs Cop I Cop II Cop 111 Cop IV Cop V Pl exo 0, 11,IV 0I 1-0, I1, 111 0, 1-0,li,ill 0,1-0, 11,111 0,1-0,1-0, 11,11 end 1, 1, I 0,0-1, 1,1 0,0-1, 1, i 0,0-0, 1, I 0,0-0, 1,1 P2 exo 0.11,111 0,1-0, 11,ll 0,I-1 , 11 ,Iii 0, I-I, II, Ill 0,1-0,1-1,11,11 end 0,2,0 0,0-0,2,0 0,0-0,2,0 0,0-0,2,0 0,0-0,2,0 P3 exo 3 0, 11,111 0,1-0, 11,11 0,1-1, 11,111 0,1-0,1-1, 11,11 end 0,2,0 0,0-0,2,1 0,0-0,2,1 0,0-1,2,1 P4 exo 3 0, 11,111 0I I-1 , 11, I I I O,l-0,1-1,11,11 end 0,2,0 0,0-0,2,1 0,0-1,2,1 PS exo 3 5 5/4* bas 2 3/2 P6 *: chaetotaxy in the female/male 76 Table 4.5.: Chaetotaxy of Cletodes tuberculatus n. sp. Pl P2 P3 P4 PS~ P5o exo 0-0-022 0-0-022 0-0-022 0-1-022 5 4 end 0-011 0-010 0-010 0-010 1 0 77 Table 4.6: Chaetotaxy of the copepodids of Cletodes tuberculatus n. sp. legs Cop I Cop II Cop 111 Cop IV Cop V Pl exo 0, 11,IV 0, 1-0, 11, 111 0,1-0, 11,111 0,1-0, 11,111 0,1-0,1-0, 11,11 end 0, 1,1 0,0-0, 1,1 0,0-0, 1,1 0,0-0, 1,1 0,0-0, 1,1 P2 exo 0,11,111 0,1-0, 11,11 0,1-0, 11,111 0,1-0, 11,111 0,1-0,1-0, 11,11 end 0, 1,0 0,0-0,1,0 0,0-0, 1,0 0,0-0, 1,0 0,0-0,1,0 P3 exo 3 0, 11,111 0,1-0, 11,11 0,1-0, 11,111 0,1-0,1-0, 11,11 end 0, 1,0 0,0-0,1,0 0,0-0, 1,0 0,0-0,1,0 P4 exo 3 0, 11,111 0,1-0, 11,111 O,l-0,1-0, 11,11 end 0, 1,0 0,0-0,1,0 0,0-0, 1,0 PS exo 3 3 5 has 1 1 P6 List of Figures. Figure 4.1.: Zosime· pacifica n. sp. female. Figure 4.2.: Zosime pacifica n. sp. female. Figure 4.3.: Zosime pacifica n. sp., female. Figure 4.4.: Zosime pacifica n. sp., male. Figure 4.5.: Zosime pacifica n. sp., copepodid I. Figure 4.6.: Zosime pacifica n. sp., copepodid II. Figure 4.7.: Zosime pacifica n. sp., copepodid Ill. Figure 4.8.: Zosime pacifica n. sp., copepodid Ill. Figure 4. 9.: Zosime pacifica n. sp. Figure 4.10.: Cletodes macrura n. sp. Figure 4.11.: Cletodes macrura n. sp. Figure 4.12.: Cletodes macrura n. sp., copepodid I and II legs. Figure 4.13.: Cletodes macrura n. sp., copepodid I and II. Figure 4.14.: Cletodes macrura n. sp., copepodid Ill and IV. Figure 4.15.: Cletodes macrura n. sp., copepodid IV and V. Figure 4.16.:Cletodes tuberculatus n. sp. Figure 4.1 7.: Cletodes tuberculatus n. sp., mouthparts and o and ~ abdomens. Figure 4.18.: Cletodes tuberculatus n. sp., legs. Figure 4.19.: Cletodes tuberculatus n. sp., copepodid I and II. Figure 4.20.: Cletodes tuberculatus n. sp., copepodid 111, IV and V. 79 Fig~l. Zosime pacifica n. sp .. female: a, habitus, dorsal; b. habirns. lareral; c, righr furcal ramus, dorsal; d, pleural region of third plcurorergite; e, posterovenrral edge of cephalothorc..x. Bo Fig!f.2. Zosime pacifica n. sp., female: a, rostrum, ventral; b, antennule; c, antenna; d, exopodite of the antenna; e, man­:lible; f, maxillule; g, maxilliped; h, maxilla; i. P5; j, abdomen, ventral. 1g:r, . . nnrijica n. sp.' fiemale·. a ' Pl,. b , P2; c, P3,. d, P4. F. u 3 Zosime r­ Fig.H. Zosime pacifica n. sp., male: a. habitus, dorsal; b, antennule; c. endopodite P2; d, PS; e, P6. , Fig4J.5. Zosime pacifica n. sp., copepodid I: a, habitus, dorsal; b, Pl, c, penultimate and anal segment, ventral; d, habitus, lateral; e, P3; f, exopodite of the antenna; g, P2; h, antennule. B'f Fig~.6. Zosime pacifica n. sp., copepodid II: a, habitus, dorsal; b, Pl, c, anal segment, ventral; d , P4; e, antennule; f, P2; g, P3 . 85 a b-f SOµm 25µm Fig.~.7. Zosime pacifaa n. sp., copepodid III: a, habitus, dorsal; b, Pl, c, P3; d, PS; c, P2; f, P4. Fig1.8. Zosime pacifica n. sp., copepodid III: a, Pl; b, P2; c, P4; d, exopodite of antenna; e, antennule; copepodid IV: f, female antennule; g, ultimate segment of male antennule; copepodid V: h, male antennule. Fig~.9. Zosime pacifica n. sp.: a, habitus (Cop IV); b, female P5 (Cop IV); c, male P5 (Cop IV); d, female P5 (Cop IV); e, male P5 (Cop V); f, habitus (Cop V); g, female P6 (Cop IV); h, male P6 (Cop IV); i, female P6 (Cop V); j, male P6 (Cop V). 8? b c Fig.'1.10. Cletodes macrura n. sp.: a, female habitus, dorsal; b, female habitus, lateral; c, male habitus, dorsal. ·-... Fig:f.11 . Cletodus macrura n. sp.: a, female abdomen, ventral; male abdomen, ventral; c, rostrum, ventral; d, male anten­nule, ventral; e, ultimate segments of the male antennule, dorsal; f, maxilla; g, maxilliped; h, female antennule; i, max­illule; j, antenna; k, mandible. -p·? _! -.\ ...... . . ;. ' , -~ i Fig~.12. Cletodes macrura n. sp., copepodid I: a, habitus, dorsal; b, anal segment, lateral; c, anal segment, ventral; d, Pl; e, P2 ; f, antennule; g, exopodite of the antenna; copepodid II: h, habitus, dorsal; i, Pl; j, P2; k, P3; I, antennule; m, exopodite of the antenna. q, 42 ..,-·-·-·-,­ Fig.4.17. Cletodes tuberculatus n. sp.: a, male abdomen, ventral view; b, female antennule; c, female abdomen, ventral view; d, ultimate segments of male antennule, dorsal; e, male antennule; f, abberant furcal ramus; g, rostrum, dorsal; h, maxilliped; i, mandible; j, maxillule; k, maxilla; I, antenna. Fig#,18. Cletodts tuberculatus n. sp.: a, Pl; b, P2; c, P3 (female); d, P4; e, endopodite P3 (male); f, female PS; g, male PS. Fig.'/J9. Cletodes tuberculatus n. sp., copepodid I: a, habitus, dorsal; b, third, fourth and anal segment, ventral; c, exopodite of the antenna; d, Pl; e, P2; copepodid II: f, habitus, dorsal; g, exopodite of the antenna; h, Pl; i, P2; j, P3; k, antennule of Copl; l, antennule of Cop II. FigS.20. Cletodes tuberculatus n. sp., copepodid III: a, Pl; b, P2; c, P3; d, P4; copepodid IV: e, Pl; f, P4; copepodid III: g, antennule; h, PS; copepodid IV: i, PS; j, antennule; copepodid V: k, PS. References BODIN, PH., 1970. Copepodes Harpacticoides marins des environs de La Rochelle. 1 -Especes de la vase intertidale de Chatelaillon. Tethys, 2: 385-436. BODIN, PH., & D. BOUCHER, 1983. Evolution a moyen terme du meiobenthos et des pigments chlorophylliens sur quelques plages polluees par la Maree Noire de l'Amoco Cadiz. Oceanologica ACTA, 6: 321-332. FRICKE, A. H., H. F.-K. HENNIG & M, J. ORREN, 1981. Relationship between oil pollution and psammolittoral meifauna density of two South African beaches Mar. Environ. Res., 5: 59-77. HAMOND, R., 1973. A review of Cletodes (Crustacea: Harpacticoida), with the description of a new species from Queensland. Mem. Qd. Mus., 16: 471-483. HENNIG, H. F.-K., G. A. EAGLE, L., FIELDER, A. H, FRICKE, W. J. GLEDHILL, P. J. GREENWOOD & M. J. ORREN, 1983. Ratio and population density of psammolittoral meiofauna as a perturbation indicator of sandy beaches in South Afrika. Environ. Mntrg. Assmt., 3: 45-60. HUMES, A. G. &J.-S. Ho, 1969. The genus Sunaristes (Copepoda, Harpacticoida) associated with hermit crabs in the western Indian Ocean. Crustaceana, 17: 1-18 HYLAND, J., D. HARDIN, E. CRECELIUS, D. DRAKE, P. MONTAGNA & M. STEIHAUER, 1990. Monitoring longterm effects of offshore oil and gas development along the southern Californian outer continental shelf and slope: background environmental conditions in the Santa Maria Basin. Oil & Chem. Pollut., 6 (in press). LANG, K., 1948. Monographic der Harpacticiden. Hakan Ohlosson, Lund, 2 vol.: 1­1682. LANG, K, 1965. Copepoda Harpacticoidea from the Californian Pacific Coast. Kungl. Svens. Vetensk, Akad., 10: 5-566. MARINOV, T. & A. APOSTOLOV, 1985. Copepodes harpacticoides de l'Ocean Atlantique 1. Especes des cotes du Sahara Espagnol. Cah. Biol. mar., 26: 165­ 180. MONTAGNA, P.A. (in preparation). Meiofauna communities in the Santa Maria Basin off the coast of California, USA., Cont. Shelf Res. MOORE, C. G., 1978. Une forme nouvelle de Cletodes Brady (Copepoda, Harpacticoida) de la cote catalan Francaise. Vie Milieu, 27 (2-A), 1977: 255-262. POR, D. F., 1967. Level bottom Harpacticoida (Crustacea, Copepoda) from Elat (Red Sea); part 1. Israel journ. Zool., 16: 101-165. POR, D. F., 1979. The Copepoda of Di Zahav pool (Gulf of Ela.t, Red Sea). Crustaceana, 37: 13-30. SCHRIEBER, G., 1984. New Harpacticoidea (Crustacea, Copepoda) from the North Atlantic Ocean. IV. Four new species of the families Diosaccidae, Ameiridae and Ancorabolidae. Cr-ustaceana, 47: 52-71. WEBB, D. G. & P. A. MONTAGNA, (in preparation). Reproductive patterns in three species of continental shelf harpacticoid copepods. PubI ished in: BEAUFORTIA,INSTITUTE OF TAXONOMIC ZOOLOGY (ZOOLOGICAL MUSEUM) UNIVERSITY OF AMSTERDAM Vol. 42, no. 2 July 12, 1991 101 CHAPTER 5: PICTORIAL KEY OF DOMINANT HARPACTICOID SPECIES (by Robert Burgess)1 1Marine Science Institute, The University of Texas at Austin, Port Aransas, Texas 78373-1267 102 Between November 1986 and May 1989, the Department of the Interior sponsored a study evaluate the long term biological impacts of continental shelf oil drilling and production. The study was named The California Outer Continental Shelf Phase II Monitoring Program (CAMP) (Hyland et al., 1990), It was centered around a proposed platform named Julius (hereafter referred to as PJ), which was never put into service. A map showing the sample sites, and an explanation of the general features of the study area can be found in Montagna 1991. Th is key was created to be used by people who are interested in identifying harpacticoid copepods, but are not taxonomists. This key is only valid for species found in the Santa Maria basin, which is on the central Californian coast. Generally the key groups animals phylogenetically. However, this does not mean that if an animal cannot be found in the key'then it is closely related to the nearest animal represented in the key. The key tries to use features which are visible without dissection. However some dissection may be necessary. To effectively use the key you must be able to see the following general features: body shape, antennules, first swimming leg, fifth leg, and the caudal rami. You then match features of your unknown animal to the appropriate group. Note it is essential to start at the beginning of the key and then follow the sequence set out in the key. The following abbreviations are used throughout: Gr. =Group, A 1 =Antennule, P1 =First swimming leg, PS =Fifth swimming leg, Cr=Caudal Rami, Exp.= Exopod, End.= Endopod, Benp. == Baseoendopod. For a more complete explanation of Harpacticoid morphological terminology see Lang (1965). 103 Choose the correct body shape and proceed to the corresponding group. G-10 G-20 G-30 G-40 G-50 G-60 IOL.f Group 1 Body Types Figure Descriptions G-1 O Torpedo-shaped body: These are from the family Ectinosomatidae. Rostrum always fused to cephlothorax, and ends in a point or is shovel shaped. G-20 Fusiform compressed body: Has a broadened prosome. There is a distinct articulation between the prosome and the urosome, or in other words the fourth and fifth segments. May or may not have a prominent rostrum. Most of these are from the families Diosaccicidae or Tisbidae. G-30 Cylindrical body: No appreciatable change in width between the prosome and urosome. Segments distinct from one another, often with ornamentation on the posterior edges. These are from the family Cletodidae. G-40 Depressed body: Dorsoventrally flattened, like isopods. These are from the family Pelt id i idae. G-50 Distinct body type: Very unusual body shape. Segments distinct. From the family Laeophontidae. G-60 Fusiform body: Large. Very pronounced second antennas, which are larger than the first antennas. From the family Cervinia. 105 Torpedo shaped body types: G-110 G-120 Microsetella Distind Separation No Distind norvegica Of Prosome Separation and Metasome Group 10 Body Types Figure Descriptions Microstella Norvigica: Rostrum turned under like a hook when viewed laterally. Small size. Caudal Rami setae as long as the body (Boeck). G-110: Has distinct separation of the prosome and the urosome. Rostrum large and fused to the cephlothorax. These are from the genera Bradya. G-120: No distinct break in the body between the prosome and urosome. These are from the genera Ectiniosoma and Pseudobradya. .. G-110 Distinct Joint Between Prosome and Urosome: Bradya Bradya cladiofera typica Group 110 Genus Bradya Figure Descriptions: Bradya typica: Rami square, with no spike off the distal ventral edge. All setae on the PS are fine (Boeck, 1872). Bradya cladiofera: Rami have a large spine originating from the distal ventral edge. The inner setae on the PS Benp. is modified to a thick spine, and the middle setae on the exopod is also modified to a thick spine (Lang 196S, p. 61 ). G-120 No Distinction Between Prosome and Urosome: G-121 G-122 Rami longer Rami almost than wi~e square Group 120 Caudal Rami Shape Figure Descriptions: G-121 Caudal Rami long: Rami have a length to width ratio greater than 1. Most of these animals come from the Genus Pseudobradya. G-122 Caudal Rami almost square: Rami have a length to width ratio of approximately 1. From the genera Halectinosoma and Ectinosoma. 111 .. G-121 Long Caudal Rami: Pseudobradya Pseudobradya pectinifera crassipes • • Group 121 Genus Pseudobradya - Figure Descriptions: - Pseudobradya pectinifera: Caudal rami are lobate in shape. Anal operculum goes past the • end of the anal segment and is the distal edge is serrated (Lang 1965, p. 82). • Pseudobradya crassipes: Caudal rami gradually taper from the base to the distal end. No serrations on the anal operculum (Lang 1965, p 73). .. ... - 113 - G-122 Short Caudal Rami: Ectinosoma Halectinosoma Paranormani kunzi Group 122 Short Caudal Rami. Figure Descriptions: Ectiniosoma paranormani: Shovel shaped rostrum. Eyespots at the base of the Antennules. Pores on all segments. The PS has no setae on the anterior surface of the exopod (Lang 196S, p. 22). Halectinosoma kunzi: Pointed rostrum. All segments have one or more rows of fine hairs. The PS a setae on the anterior surface of the exopod (Lang 196S, p. S2). 11 S Fusiform compressed body shape. Zosime Diagoniceps Stenheli pacifica trifidus latipes Group 20 Body Type Figure Descriptions Zosime pacifica n.sp: Large stout Antenulles. Rostrum small and fussed to the cephlothorax. Strong thorns along the distal edge of each segment (Fiers 1991). Diagoniceps trifidus: Small rounded rostrum. P4 is very large and pronounced. PS is ventrally displaced (Yeatman 1980). Stenheli latipes: Large pronounced rostrum, with a bifid tip. PS is laterlly displaced (Lang 196S, p. 236). 117 • • - Cylindrical body shape. - ... - Reduced Rosrum Small Rounded Pronounced Rostrum Rostrum G-310 G-320 G-330 • '1 e ­ Group 30 Body Type Figure Descriptions ­ G-310 Reduced rostrum: Fused to cephlothorax, looking like a small bulge with 2 hairs near the apex. G-320 Round rostrum: Rostrum articulated at base, small and round. G-330 Pronounced rostrum: Rostrum is fused, large, pointed and pronounced. 119 G-310 Reduced Rostrum Stylicletodes C/etodes Cletodes longicaudatus Macrura turberculatus Group310 Figure discription Stylicletodes longicaudatus: Very long caudal rami, legnth to width ratio is 9.2 I 1. Large pronounced PS (Brady et Robertson 1875, p 196). Cletodes macrura n. sp. Moderate legnth caudal rami, legnth to width ratio is3. 9 I 1. Dorsal setae on the caudal rami is inserted on distal half of the caudal rami, marked with an arrow on the drawing (Fiers 1991). Cletodes turberculatus n. sp.:Moderate legnth caudal rami, legnth to width ratio is 4.9 I 1. Dorsal setae on the caudal rami is inserted on the proximal half of the caudal rami, marked with an arrow on drawing (Fiers 1991). 121 .. Group 320 Figure discription Only one species of this type has been identified, Tetragoniceps pacifica (Burgess unpublished). Group 330 Figure discription Only one species of this type has been identified, Serratithorus geei n. sp. (Burgess unpublished) 123 Group 40 Body Type Figure Descriptions Only one species of this body type has been identified, Paralteutha simile (Monk 1939). Group 50 Body Type Figure Descriptions· Only one species of this body type has been identified, Echinolaeophonte armiger (Gurney in Lang 1965, p. 509). 125 Group 60 Body Type Figure Descriptions Only one species of this body type has been identified, Cervinia mediocauda n.sp (Burgess unpublished). Table 5.1. Harpacticoid species covered in pictorial key. Family Species Cerviniidae Ectinosomatidae Tisbidae Peltidiidae Diosaccidae Cletodidae Tetragonicipitidae Laeophontidae Cervinia mediocauda M icrostella norvigica Bradya typica Bradya cladiofera Pseudobradya crassipes Pseudobradya pectinifera Ectiniosoma paranormani Halectiniosoma kunzi Zosime pacifica Paraalteutha simile Stenhelia latipes Cletodes macrura Cletodes turberculatus Serratithorus geei Stylicletodes longicaudatus Diagoniceps trifidus Tetragoniceps pacifica Echinolaeophonte armiger 127 Acknowledgements I am much indebted to Dr. Paul Montagna, at the University of Texas Marine Science Institute, not only for the specimens and for use of the facilities, but also for his professional and technical support. This study was supported by the US Department of the Interior, Minerals Management Service (Pacific OCS Office, Los Angles, California, ·usA 90017) under contract No. 14-12-0001-30262. References Coull, B. C. (1977) Marine flora and fauna of the northeastern United States. Copepoda: Harpacticoida. NOAA Tech. Report. NMFS Circ 399.48pp Fiers, F. (1991) Three new harpacticoid copepods from the Santa Maria Basin of the Californian Pacific Coast (Copepoda, Harpacticioda). Beaufortia Institute of taxonomic zoology University of Amsterdam, 42 (2): 13-47. Hyland J., D. Hardin, I. Crecelius, D. Drake, P. Montagna, and M. Steinhauer (1990) Monitoring long-term effects of offshore oil and gas development along the southern California outer shelf and slope: background environmental conditions in the Santa Maria Basin. Oil and Chemical Pollution, 6, 195-240. Lang, K. (1948). Mongraphie der Hrpacticiden, 1&11. Hakan Ohlsson, Lund. 1682 pp. Lang, K. (1965). Copepoda Harpacticoida from the California Pacific coast. Kung/. Svenska Vetensk. Hand/., Fjarde, 10(2): 1-560. Monk, C. (1939). Marine hapacticoid copepods from California. Trans. of Amer. Microsc. Soc. 60: 75-103. Sars, G.O. (1911) An account of the Crustacea of Norway. Copepoda Harpacticoida. Bergen Museum, Norway. 499 pp. 129 APPENDIX: DRAWINGS OF UNDESCRIBED SPECIES (by Frank Fiers)1 1Koninkliik Beleisch lnstituur voor Natuurwetenschappen, Recent Invertebrate Section, Vautierstraat 29, B-1040 Brussels, Belgium 130 APPENDIX: DRAWINGS OF UNDESCRIBED SPECIES (by Frank Fiers)1 1Koninkliik Beleisch lnstituur voor Natuurwetenschappen, Recent Invertebrate Section, Vautierstraat 29, B-1040 Brussels, Belgium Figure 6.1. Ceratonotus pectinatus. a: Female Habitis. b: Female abdomen. c: Female antennule. d: Female maxilliped. 132.. r c RA 7(J\/ r T ()~ ()C( hnCJ fu S '1 Figure 6.2. Ceratonotus n. sp. 1. a: Male Habitis. b: Male habitis, lateral. c: Male abdomen. d: Male PS. 25 µm: c Id c ( Ł RfJ TO NC' n v·;, nsp ~ Figure 6.3. Ceratonotus n. sp. 2. a: Male Habitis. b: Male abdomen, ventral. c-h: Setal variability. c d i I I SOµm '/ I b-h I I 25µm e ® f"K,") J 13b I' ,-. /') fl 7 r: /\ · ro -/ · ~ Figure 6.4. Ceratonotus n. sp. 2. a: Antenna. b: P1. c: P2. d: P3. e: PS. f-h: Pereiopodal setal variablity. I;/I .'/ \ / / / 138 /)5;) 3 Figure 6.5. Amphiascoides n. sp. a: Habitus female. b: Habitus male. c: Male CR. d: Female CR. e: Female CR, ventral. A MfA 7 Figure 6.6. Amphiascoides n. sp. a: Antennule female. b: Rostrum and Antenna. c: Labrum. d: Maxilliped. e: Gnathobase of the mandible. f: Maxilla. g: PS female. h: PS male. tJif pf} I AMPA 1 Figure 6.7. Amphiascoides n. sp. a: Pl. b: P2. c: Ornamentation variation. d: Ornamentation variation. e: Basis and coxal variation. f: Abdomen Male, ventral. 143 AM PA 3 Figure 6.8. Amphiascoides n. sp. a: P3 Female. b: P3 Male. c: P4. AN PA 't Figure 6. 9. Species C n. sp. a: Habitus Female. b: Abdomen Female. c: Abdomen Male. d: Habitus Male. e: CR, dorsal view. f: CR, ventral view. g: CR, lateral view. l4B . C I (_ 1 Figure 6.10. Species C n. sp. a: Pl. b: Inner coxal seta. c: P2. d: P3. e: P4. f: Baseoendopod and P 5 of the male. g: Modified seta. h: Baseoendopod and P 5 of the female. 149 ISO Sp c 2. Figure 6.11. Species A n. sp. a: Habitus Male. b: Abdomen Male. c: Anten n na. d: P2 basis of the male. e: P 2 endopod of the male. f: PS of the male. g: Modified seta. h: Baseoendopod and PS of the female. 1S1 152.. Figure 6.12. Species PADA n. sp. a: Habitus Female. b: Abdomen Female. c: PS female. d: Genital segment. e: Maxilliped. J5a+ Figure 6.13. Species PADA n. sp. a: Habitus Male. b: Caudal rami setae. c: Abdomen Male. d: Endopod P2. e: Endopod P3. f: P3. Figure 6.14. Species PADA n. sp. a: Antennule of the female. b: Antennule of the male. c: Rostrum. d: Antenna. e: Exopodite of the Antenna. f: Modified setae on the Maxilla. g: Mandible. h: Maxillule. i: Maxilla. j: Baseoendopod and P 5 of the female. 25µm 25µm Figure 6.15. Species PADA n. sp. a: P1. b: P2. c: P3. d: P4. Figure 6.16. Species DANA n. sp. a: Habitus female. b: Female abdomen, ventral view. c: Male abdomen, ventral view. 7:;Jl1 NA 1 Figure 6.17. Species DANA n. sp. a: Antennule Female. b: Antennule female, dorsal view. c: Antennule female, ventral view. d: Antenna. e: Exopodite of the Antenna. f: Mandible. g: Maxilliped. h: Maxilla. h':Syncoxal endite. I: Maxillule. j: Arthrite of the maxillule. Figure 6.18. Species DANA n. sp. a: PS. b: P4. c: P3. d: P2. e: P1. ,., ' 165 Figure 6.19. Species DANA n. sp. a: Abdomen male. b: P2 endopod, male. b': Endopod 3, male P2. c: P3 exopod, male. d: P1 endopod, male. e: P3 exopod, male. f: P2 exopod, male. g: P1 exopod, male. h: Antennule, male. i: PS male. Figure 6.20. Species PADB n. sp. a: Habitus female. b: Abdomen male, ventral view. c: Abdomen female, ventral view. Figure 6.21. Species PADB n. sp. a: Pl female. b: P2 female. c: P2 endopod, male. d: P3 basis and endopod of the female. e: P3 exopod, female. f: P4 basis and endopod of the female. g: P4 exopod, female. \ '7 2. Figure 6.22. Species PADB n. sp. a: Antennule Female. b: Antennule Male. c: Mandible. d: Antenna. e: Rostrum. f: Maxilla. g: P 5 female. h: PS male. 25µm I '7~ Figure 6.23. Species PSLO n. sp. a: Habitus Female. b: Abdomen Female. c: Genital field. d: Maxillule. e: Arthrite of the maxillule. f: Antenna of the female. g: Mandible. h: P 5 female. 175 ?Slo -:1­ Figure 6.24. Species PSLO n. sp. a: Pl. b: P2. c: P3. d: P4. e: Modified seta on P3 endopod 3. 177 110 ?st-o 2 Figure 6.25. Species STED n. sp. a: Habitus female. b: Caudal rami. c: Habitus male. d: P1 endopod, male. e: Antennule Male. f: Rostrum. -1­ 5/C:O Figure 6.26. Species STED n. sp. a: Female abdomen, ventral view. b: Antennule female. c: Maxilliped. d: Maxilla. e: Mandible. f: Antenna. g: Maxillule. 181 Figure 6.27. Species STED n. sp. a: P2. b: P3. c: P4. 1'2A­ S7ED3 Figure 6.28. Species STED n. sp. a: Male abdomen, ventral view. b: P1 female. c: Male abdomen, lateral view. d: PS male. e: P 5 female. f: lntercoxal sclerite. 185 s TED L1 Figure 6.29. Species MM n. sp. a: Habitus Female. b: Terminal seta of the caudal rami. c: Female abdomen, ventral view. d: Antennule female. e: Antennule male. 187 c:::; p /"7 ;\'( 1 Figure 6.30. Species MM n. sp. a: P2. b: P3 endopod of the male. c: P3. d: P4. 189 2 Figure 6.31. Species MM n. sp. a: Male abdomen, dorsal view. b: Male abdomen, ventral view. c: PS male varient. d: PS male. e: Baseoendopod of the female. f: PS female. g: P1. 191 Figure 6.32. Species STEC n. sp. a: Habitus Female. b: Abdomen Female. c: Rostrum. 193 SOµm c Figure 6.33. Species STEB n. sp. a: Habitus female. b: Antennule male. c: Exopod of the antenna. d: Antenna. e: Antennule of the female. f: Female abdomen, ventral view. 195 i'h Figure 6.34. Species STEB n. sp. a: P4. b: PS female. c: PS male. d: Abdomen female. e: Abdomen male. 197 .. Figure 6.35. Species STEB n. sp. a: P2 endopod of the male. b: Pl. c: P2 female. d: P4. e: P3 endopod of the male. 199 Figure 6.36. Species J n. sp. a: Habitus Female. b: Rostrum. c: Abdomen female. d: Habitus male. 201 '2.t2­ 5PJ ~ Figure 6.37. Species HEST n. sp. a: Habitus Female. b: Habitus male. c: Maxilliped. d: Maxilla. e: Mandible. f!~S T-f Figure 6.38. Species HEST n. sp. a: Female abdomen. b: Male abdomen, lateral view. c: Male abdomen, ventral view. d: P 5 female. ftc=-S7 2­ Figure 6.39. Species HEST n. sp. a: Antennule Female. b: Maxillule. c: Antennule Male. d: Antenna. e: Male P 1 coxa. f: P1. 1-1c=ST] Figure 6.40. Species HEST n. sp. a: P4. b: P3. c: P2 endopod of the male. d: P2. 209 c a -d 25µm Figure 6.41. Species NOCO n. sp. a: Habitus female. b: Endopod of the P2. c: Endopod of the P3. d: Habitus male. e: PS. 211 )./OC o 1 Figure 6.42. Species CLET L n. sp. a: Habitus female. b: Abdomen male. c: Abdomen female. d: Habitus male. 213 Ł. I 'i Figure 6.43. Species CLET L n. sp. a: Antennule of the female. b: Antenna. c: Mandible. d: Maxilliped. e: Maxillule. f: Maxilla. g: Antennule of the male. h: Rostrum. 215 21b Figure 6.44. Species CLET L n. sp. a: Pl. b: P2. c: P 5 of the female. d: P4. e: PS male. f: P2 male. 217 Clt;Tl-3 Figure 6.45. Species CLM n. sp. a: Caudal ram i. b: Habitus Female, lateral view. c: Abdomen female. d: Abdomen male. e: Habitus female, dorsal view. 219 ,. 25µm ..,. Figure 6.46. Species CLM n. sp. a: P1. b: P2. c: Male abdomen. d: P 5 female. e: P3. f: PS male. g: P4. 221 2. 22. {_ /'1 2 c Figure 6.47. Species CLET MILL n. sp. a: Female habitus, dorsal view. b: Female habitus, lateral view. c: Male habitus, dorsal view. 223 2 2..4 c l E T-/'11 L l . -d. Figure 6.48. Species CLET MILL n. sp. a: Abdomen male. b: Antennule Female. c: Antenna. d: PS male. e: P2 endopod of the male. f: PS female. g: Last 3 segments of the male antennule. h: Male antennule. 22S f 22.~ b -h ( .. t,.c--I 1~1Ll. 2-. Figure 6.49. Species CLET D n. sp. a: Female habitus, dorsal view. b: Female habitus, lateral view. c: Habitus male. d: Genital segment. e: Maxilliped. 227 228 CL ET]) Figure 6.50. Species I n. sp. a: Habitus Female. b: Habitus Male. 229 Figure 6.S 1. Species I n. sp. a: Female abdomen, lateral view. b: Antennule Female. c: Antenna. d: Mandible. e: Rami of the mandible. f: Maxillule. g: Maxilla. h: Maxilliped. i: gen ital segment. j: PS female. k: PS Male. I: Male abdomen, ventral view. 25µm SOµm 25µm Figure 6.52. Species I n. sp. a: P1. b: P2. c: P3. d: P4. e-f: Varation of P2 endopod. '5-P1 3 Figure 6.53. Species I n. sp. a: Male antennule. b-d: Male P2 variation. srT4 Figure 6.54. Species W n. sp. a: Habitus female. b: Female abdomen, ventral view. c: Female abdomen, lateral view. Figure 6.55. Species VV n. sp. a: Antennule Female. b: Antenna. c: Mandible. d: Maxilliped. e: Rostrum. srv v 2­ Figure 6.56. Species W n. sp. a: P1. b: P2. c: P3. d: P4. 2 42. 5-P LIV 3 Figure 6.57. Species Z n. sp. a: Habitus Female. b: Abdomen Female. c: PS female. 243 c a 25µm SOµm Figure 6.58. Species Zn. sp. a: P4. b: P3. c: P2. d: P1. c d Figure 6.59. Species EURY n. sp. a: Habitus Female. b: Abdomen Female. c: Antenna. d: Antennule female. e: PS. 247 _____.i--­ e /-/ i /) v A Figure 6.60. Species EURY n. sp. a: P1. b: P2. c: P3. d: P4. '2.. so F / J()V ',7 Figure 6.61. Species CLET A n. sp. a: Habitus female. b: Abdomen female. c: Habitus male. d-e: Variation of caudal rami. 251 ' . -;-A /! -( --1 Figure 6.62. Species CLETA n. sp. a: Abdomen male. b: Antennule female. c: Antennule male. d: Antenna. e: Mandible. f: Maxillule. g: Maxilla. h: Maxilliped. 253 h 25µm 1. 54 Ct FT/} ~ c. (___ Figure 6.63. Species CLET A n. sp. a: P1. b: P2. c: P3. d: PS male. e: Male PS baseoendopodal variation. f: PS female. g: P2 male. d e a -g 25µm .3 Figure 6.64. Species CLET R n. sp. a: Habitus male. b: Male abdomen, ventral view c: Male abdomen, lateral view d: PS male. e: Antennule male. 257 2. 56 CL ~T i<. Figure 6.65. Species CLET P n. sp. a: Female habitus, dorsal view. b: Female habitus, lateral view. c: Habitus male. CLET? ~ Figure 6.66. Species STLO n. sp. a: Habitus female. b: Terminal seta of caudal rami. c: Habitus female lateral. d: Habitus male. e: P2 endopod male. f: P6. g-h: Variation of rostrum. Figure 6.67. Species STLO n. sp. a: Abdomen male. b: Habitus male lateral. c: Caudal ram i. d: Gen ital segment. e: Antennule female. f: P6. g: Antennule male. 263 5-rto 2.. Figure 6.68. Species STVE n. sp. a: Female habitus, dorsal view. b: Female habitus, lateral view. c: Terminal seta of the caudal rami. d: Genital segment. e: P6. 265 e 25µm 2 bto Figure 6.69. Species CERD n. sp. a: Habitus female. b: Female abdomen, ventral view. c: Female abdomen, lateral view. d: Maxilla. ( c--,~ u ,1_ Figure 6.70. Species CERD n. sp. a: Antennule Female. b: Antenna. c: Exopod of the Antenna. d: Maxilliped. 269 (c1-ftfll°t 1 ,,,,~\\\\'''},,,1111 r;~r-i-t-'-l-1-'-'·'-'-'-"'''''"' r.; 1 I I • 1 ' 110 '' I~ I 11 ',1' ,'11 I ,,1 ,1 II II• 1•,1,,\\ 1 I I • r I I 11' ft f I 0 'I I f f t \a\ t , I I,, I 1 t I ; ', .' '. ' : ';, ', ::: ,',• : I ' ' I ' ', I ' ' ,' 1 I~ : I; 111 I ~ ~.. .,,.,.,,,,, ...........,,, ,,~ ~ 1, t I t I I I I I / I I I I r I I t I\ '\ \ ' \ \ " ~ ~ \ 1 I ' ,,,,I r•tl.J.,._,_1.r.tLL\.I.\.\.\.\.\,;~~" \ .~,,,,,,,··f.J......~" \ ;~'' ::..:::---­ //, . e 25µm a -d 25µm Figure 6.97. Species TACB n. sp. a: Male abdomen, dorsal view. b: Male abdomen, lateral view. c: Male abdomen, ventral view. d: Male P2 endopod. e: Female P2. f: P3 g: P4 h: Female PS 323 32'#­ 2 Figure 6.98. Species 00 n. sp. a: Female Habitus. b: Male antennule. c: Antenna. d: Exopod of the antenna. e: Female antennule. f: Male PS. 325 Figure 6.99. Species 00 n. sp. a: Female abdomen, dorsal view. b: Female abdomen, ventral view. c: Male abdomen, dorsal view. d: Male abdomen, ventral view. 327 328 <;foo 2­ Figure 6.100. Species 00 n. sp. a: P1. b: PS female. c: P2. d: P3. 329 Figure 6.101. Species TYPA n. sp. a: Habitus female. b: Female abdomen, ventral view. c: Female abdomen, lateral view. d: Male P2 endopod. e-g: P1 basis and precoxa variation. h: Male abdomen, ventral view. 331 a -c ,h --·p;; -d. I y => ;· .• Figure 6.102. Species PSMA n. sp. a: Female habitus, dorsal view. b: Female habitus, lateral view. c: Rostrum. d: Female abdomen. 25µm ~I \~~ ~ 75Jf/.) 4. --. Figure 6.103. Species PSMA n. sp. a: Pl. b: P2. c: P3. d: P4. e: PS. f: lnnercoxal sclerite. 335 c a -f 25 µm 33b Ps rn A ? '-­ The Department of the Interior Mission As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural resources. This includes fostering sound use of our land and water resources; protecting our fish, wildlife, and biological diversity; preserving the environmental and cultural values of our national parks and historical places; and providing for the enjoyment of life through outdoor recreation. The Department assesses our energy and mineral resources and works to ensure that their development is in the best interests of all our people by encouraging stewardship and citizen participation in their care. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration. The Minerals Management Service Mission As a bureau of the Department of the Interior, the Minerals Management Service's (MMS) primary responsibilities are to manage the mineral resources located on the Nation's Outer Continental Shelf (OCS), collect revenue from the Federal OCS and onshore Federal and Indian lands, and distribute those revenues. Moreover, in working to meet its responsibilities, the Offshore Minerals Management Program administers the OCS competitive leasing program and oversees the safe and environmentally sound exploration and production of our Nation's offshore natural gas, oil and other mineral resources. The MMS Royalty Management Program meets its responsibilities by ensuring the efficient, timely and accurate collection and disbursement of revenue from mineral leasing and production due to Indian tribes and allottees, States and the U.S. Treasury. The MMS strives to fulfill its responsibilities through the general guiding principles of: (1) being responsive to the public's concerns and interests by maintaining a dialogue with all potentially affected parties and (2) carrying out its programs with an emphasis on working to enhance the quality of life for all Americans by lending MMS assistance and expertise to economic development and environmental protection.