A Review of the Genus Eunice - Smithsonian Institution Libraries

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 7. Eunice marovoi 213 8. Eunice mexicana 219 9. Eunice mucronata 226 10. Eunice muUicylindri 227 11. Eunice nicidioformis 238 12. Eunice ornata 246 13. Eunice polybranchia 273 14. Eunice pycnobranchiata 283 15. Eunice torresiensis 322 16. Eunice validobranchiata 335 17. Eunice violaceomaculata 336 18. Eunice vittata 339 Comparison among Species 19. Overview of group A-l 345 20. Group A-l with articulated peristomial cirri 346 21. Group A-l without articulated peristomial cirri 347 22. Overview of groups A-2 and A-4 348 23. Group A-2 348 24. Overview of group B-l 349 25. Group B-l with antcnnal articulations 350 26. Group B-l without antennal articulations 350 27. Overview of group B-2 351 28. Group B-2 without antennal articulations 353 29. Group B-2 with cylindrical antcnnal articulations and all nolopodial cirri articulated 354 30. Group B-2 with cylindrical antcnnal articulations and notopodial cirri articulated anteriorly only 355 31. Group B-2 with cylindrical antcnnal articulations but without notopodial articulations 356 32. Group B-2 with drop-shaped or moniliform antcnnal articulations 357 33. Overview of groups B-3 and B-4 358 34. Groups B-3 and B-4 with articulated antennae 359 35. Groups B-3 and B-4 with articulated antennae and pectinate branchiae .... 360 36. Groups B-3 and B-4 with articulated antennae and palmate or simple branchiae 360 37. Groups B-3 and B-4 without antennal articulations 361 38. Group B-4 without antennal articulations; with branchiae and with median antennae longer than other antennae 362 39. Group B-4 without antennal articulations, with branchiae and all antennae equally long 363 40. Abranchiate species 364 41. Overview of group C-l 365 42. Group C-l with cylindrical antennal articulations 366 43. Group C-l with distally drop-shaped or moniliform articulations 367 44. Group C-l with all articulations drop-shaped or moniliform 367 45. Group C-l without antennal articulations 368 46. Overview of group C-2 369 47. Group C-2 with all notopodial cirri articulated 370 48. Group C-2 with only anterior notopodial cirri articulated 371 49. Group C-2 without notopodial articulations 371 50. Overview of Group D 372 51. Group D 372 52. Overview of species with unusual kinds of setae 373 53. Species with unusual kinds of setae 373
A Review of the Genus Eunice (Polychaeta: Eunicidae) Based upon Type Material Kristian Fauchald Introduction Eunicean polychaetes are uniquely defined by the presence of laterally arranged maxillae and paired ventral mandibles mounted on an eversible, muscular U-shaped structure (Ehlcrs, 1868; Dales, 1962; Clark, 1964; Fauchald, 1974a). Traditionally the group was considered as a single family, Eunicidae, with a series of subfamilies (Fauvel, 1923; Day, 1967). Lately most authors recognize the group as a distinct order, Eunicida (e.g., Fauchald, 1977; Pettibone, 1982), with a series of families. The family Eunicidae Berthold (1827) is the nominal family. Other families include Onuphidae Kinberg (1865), Lumbrineridae Malmgren (1867), Harunaniellidae Imajima (1977), Iphitimidae Fauchald (1970), Arabellidae Hartman (1944), Lysaretidae Kinberg (1865), Oenoniidae Kinberg (1865), and Dorvilleidae Chamberlin (1919a). The parasitic families Histriobdellidae Vaillant (1890) and Ichthyotomidae Eisig (1906) and the mesopsammic family Dinophilidae Remane (1932) are usually considered as members of the order, but are sometimes relegated to a satellite status. Relationships between the Eunicidae and the other families are outlined below in the section on cladistics. Formal definitions of the families can be found in Fauchald (1977), Pettibone (1982), and Colbath (1989). Eunicoid polychaetes are known from fossilized jaws present in Ordovician fossil beds (Kielan-Jaworowska, 1966). Members of the order are present in all marine environments, most are free-living; parasitic taxa are found in part on other polychaetes, in part on fish or crustaceans. In some families, all or nearly all species are tubicolous; in other families most species lack tubes. Eunicids sensu stricto are present in all marine benthic environments. They are especially common in tropical shallow seas, in coral reef rubble, and in mangrove Kristian Fauchald, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington DC. 20560. swamps. Some species form complex tubes (e.g., Hanley, 1986:215); most tubicolous species form less-complicated structures (Ehlers, 1887), but many eunicids lack tubes completely. Most species live in cracks and crevices in mixed rubble, rock, and sand environments in shallow water. Many species drill in dead coral rubble (Hutchings, 1981) and are sufficiently common to be of considerable ecological importance. The Atlantic and Pacific palolo-worms are known for their brief, lunar periods of spawning (e.g., Mayer, 1900; Hofmann, 1974). The Atlantic and Pacific palolos belong to different genera, Eunice and Palolo, respectively. The habit of having brief, intense spawning periods may be more common in the family than currently appreciated. The first described eunicid species was originally placed in Nereis. Nereis norvegica Linnaeus (1767) was described from an ahermatypic coral reef in Norway. Nereis aphroditois was described from a hermatypic coral reef in Sri Lanka (Ceylon) by Pallas (1788). Other species were added by O.F. Miiller from Norway (Muller, 1776, 1779). Cuvier (1817:524) named a new genus, Eunice, to contain these and related taxa (a more detailed discussion of nomenclatural problems is given below). Audouin and Milne Edwards (1832), Kinberg (1865), Quatrefages (1866), Malmgren (1867), Ehlers (1868), and Grube (1870b, 1878a,b) reviewed early work and added numerous new species. Mclntosh (1885) and Chamberlin (1919a) added new taxa based on materials from the Challenger and Albatross cruises, respectively, many from deep-water habitats. Treadwell (1921, 1922) added many new species from coral reefs in the Caribbean Sea and the Pacific Ocean, respectively. He also attempted to expand the set of features used to characterize species to include color patterns; he was not followed in this attempt by any later worker. The live color is, however, very characteristic for several species and may be useful as a character in a well-studied geographical area. Fauvel (1917, 1919) reviewed many of the tropical species, especially from the Indian Ocean, and established numer-
Page 1 and 2: * A Review of the Genus Eunice (Pol
Page 3 and 4: S M I T H S O N I A N C O N T R I B
Page 5 and 6: Contents Page Introduction 1 Nomenc
Page 7 and 8: NUMBER 523 Eunice brevis (Ehlers, 1
Page 9 and 10: NUMBER 523 vii Eunice longicirris G
Page 11: NUMBER 523 ix Eunice stanleyi, new
Page 15 and 16: NUMBER 523 (1820) are exemplary as
Page 17 and 18: NUMBER 523 included, but not specie
Page 19 and 20: NUMBER 523 subsequent workers. They
Page 21 and 22: NUMBER 523 illustrations. Seial len
Page 23 and 24: NUMBER 523 11 uniformly yellowish g
Page 25 and 26: NUMBER 523 13 to the frontal jaws (
Page 27 and 28: NUMBER 523 15 be longer than the ot
Page 29 and 30: NUMBER 523 17 rounded lobe about as
Page 31 and 32: NUMBER 523 19 Both outgroup species
Page 33 and 34: NUMBER 523 21 By "near posterior en
Page 35 and 36: NUMBER 523 23 69. dentition of appe
Page 37 and 38: NUMBER 523 25 Eunice djiboutiensis
Page 39 and 40: NUMBER 523 27 pauper a Grube, 1878b
Page 41 and 42: NUMBER 523 29 length of antennae in
Page 43 and 44: NUMBER 523 31 peristomium rubra 32(
Page 45 and 46: NUMBER 523 33 sulcus shallow tentac
Page 47 and 48: NUMBER 523 35 Peristomial cirri dig
Page 49 and 50: NUMBER 523 37 peristomial cirri dig
Page 51 and 52: NUMBER 523 39 Branchiae palmate [Fi
Page 53 and 54: NUMBER 523 41 .1 .05 1 5 d b c FIGU
Page 55 and 56: NUMBER 523 43 .02 .1 c, d FIGURE 4.
Page 57 and 58: NUMBER 523 45 and 49. In addition t
Page 59 and 60: NUMBER 523 47 4. Eunice afra Peters
Page 61 and 62: NUMBER 523 49 median antennae simil
Page 63 and 64:
NUMBER 523 51 afuerensis and E. pel
Page 65 and 66:
NUMBER 523 53 reaching beyond anter
Page 67 and 68:
NUMBER 523 55 .02 .1 1 b. c d FIGUR
Page 69 and 70:
NUMBER 523 57 FIGURE 10.—Eunice a
Page 71 and 72:
NUMBER 523 59 TABLE 2.—Variable a
Page 73 and 74:
NUMBER 523 61 .02 .5 .5 .1 2 e. f,
Page 75 and 76:
NUMBER 523 63 .02 1 .1 f, y, h C FI
Page 77 and 78:
NUMBER 523 65 midline. Pre- and pos
Page 79 and 80:
NUMBER 523 67 Subacicular hooks (Fi
Page 81 and 82:
NUMBER 523 69 REMARKS.—Eunice arm
Page 83 and 84:
NUMBER 523 71 REMARKS.—Eunice art
Page 85 and 86:
NUMBER 523 73 FIGURE 17.—Eunice a
Page 87 and 88:
NUMBER 523 75 • 02 , d a FIGURE 1
Page 89 and 90:
NUMBER 523 77 Proximal teeth larger
Page 91 and 92:
NUMBER 523 79 FIGURE 20.—Eunice b
Page 93 and 94:
NUMBER 523 81 been unable to see th
Page 95 and 96:
NUMBER 523 83 in 120-150 m depth. T
Page 97 and 98:
Page 99 and 100:
Page 101 and 102:
NUMBER 523 89 FlOURE 24.—Eunice b
Page 103 and 104:
NUMBER 523 91 compound falcigcrs (F
Page 105 and 106:
NUMBER 523 93 EXPECTED STATE OF UNK
Page 107 and 108:
Page 109 and 110:
NUMBER 523 97 FIGURE 29.—Eunice c
Page 111 and 112:
NUMBER 523 99 TABLE 3.—Variable a
Page 113 and 114:
NUMBER 523 101 characters, best exp
Page 115 and 116:
NUMBER 523 103 diagnostic." As indi
Page 117 and 118:
NUMBER 523 105 FIGURE 31.—Eunice
Page 119 and 120:
NUMBER 523 107 very short. Shafts s
Page 121 and 122:
NUMBER 523 109 Presetal lobes low,
Page 123 and 124:
NUMBER 523 111 which are similar to
Page 125 and 126:
NUMBER 523 113 45. Eunice complanat
Page 127 and 128:
NUMBER 523 FIGURE 35.—Eunice cong
Page 129 and 130:
NUMBER 523 117 REMARKS.—Eunice co
Page 131 and 132:
NUMBER 523 119 50. Eunice curticirr
Page 133 and 134:
NUMBER 523 121 Anterior neuropodial
Page 135 and 136:
NUMBER 523 123 a wide triangular ba
Page 137 and 138:
Page 139 and 140:
NUMBER 523 127 55. Eunice edwardsi
Page 141 and 142:
NUMBER 523 129 reduced, but still v
Page 143 and 144:
NUMBER 523 131 1 2 .5 1 1 a h d h,
Page 145 and 146:
NUMBER 523 133 cirri did not reach
Page 147 and 148:
Page 149 and 150:
NUMBER 523 137 J .02 .1 1 a c, e d
Page 151 and 152:
NUMBER 523 139 .0 2 1 2 1 .1 .5 1 h
Page 153 and 154:
NUMBER 523 141 peristomium. Prostom
Page 155 and 156:
NUMBER 523 143 shorter than filamen
Page 157 and 158:
Page 159 and 160:
NUMBER 523 147 TABLE 5.—Variable
Page 161 and 162:
Page 163 and 164:
NUMBER 523 151 .02 1 1 .1 f. g, h b
Page 165 and 166:
NUMBER 523 153 complete specimen).
Page 167 and 168:
NUMBER 523 155 branchiae is rare in
Page 169 and 170:
NUMBER 523 157 Limbate and pectinat
Page 171 and 172:
NUMBER 523 159 indistinct in both a
Page 173 and 174:
NUMBER 523 161 79. Eunice grubei Gr
Page 175 and 176:
NUMBER 523 163 Prostomium (Figure 5
Page 177 and 178:
NUMBER 523 165 24, present in all s
Page 179 and 180:
Page 181 and 182:
NUMBER 523 169 85. Eunice hawaiensi
Page 183 and 184:
NUMBER 523 171 105 setigers; total
Page 185 and 186:
NUMBER 523 173 dorsally and especia
Page 187 and 188:
Page 189 and 190:
NUMBER 523 177 Filaments slender. N
Page 191 and 192:
NUMBER 523 179 tween Mx III and lef
Page 193 and 194:
Page 195 and 196:
NUMBER 523 183 transverse folds. Al
Page 197 and 198:
NUMBER 523 185 CHARACTERS USED IN P
Page 199 and 200:
NUMBER 523 187 • 02 .5 ] .1 2 .5
Page 201 and 202:
NUMBER 523 189 02 FIGURE 62.—Euni
Page 203 and 204:
NUMBER 523 191 .02 1 .1 e. t, q h.
Page 205 and 206:
Page 207 and 208:
NUMBER 523 195 FlOURE 65.—Eunice
Page 209 and 210:
NUMBER 523 197 Peristomial cirri to
Page 211 and 212:
Page 213 and 214:
NUMBER 523 201 near posterior end,
Page 215 and 216:
Page 217 and 218:
Page 219 and 220:
NUMBER 523 207 falcigers agree well
Page 221 and 222:
NUMBER 523 209 cirri, not reduced i
Page 223 and 224:
NUMBER 523 211 Limbate setae margin
Page 225 and 226:
Page 227 and 228:
Page 229 and 230:
NUMBER 523 217 EXPECTED STATES OF U
Page 231 and 232:
Page 233 and 234:
NUMBER 523 221 articulations in A-1
Page 235 and 236:
NUMBER 523 223 J32 J b, c, d a FIGU
Page 237 and 238:
NUMBER 523 225 1 10 b, d, e, g. h,
Page 239 and 240:
NUMBER 523 227 126. Eunice multicyl
Page 241 and 242:
Page 243 and 244:
Page 245 and 246:
NUMBER 523 233 tion between rings d
Page 247 and 248:
NUMBER 523 235 ASSUMED STATES FOR P
Page 249 and 250:
NUMBER 523 237 visible ventrally; a
Page 251 and 252:
Page 253 and 254:
NUMBER 523 241 proximally cylindric
Page 255 and 256:
NUMBER 523 243 digitiform. Anterior
Page 257 and 258:
NUMBER 523 245 Eunice oliga papeete
Page 259 and 260:
NUMBER 523 247 h. i FIGURE 82.—Eu
Page 261 and 262:
NUMBER 523 249 DESCRIPTION.—Body
Page 263 and 264:
Page 265 and 266:
NUMBER 523 253 the type, there is l
Page 267 and 268:
NUMBER 523 255 directed laterally,
Page 269 and 270:
NUMBER 523 257 2 1 .1 b, c, », h d
Page 271 and 272:
NUMBER 523 259 RGURE 86.—Eunice p
Page 273 and 274:
NUMBER 523 261 colored, tridentate
Page 275 and 276:
NUMBER 523 263 papeetensis has expa
Page 277 and 278:
Page 279 and 280:
NUMBER 523 267 These two species ar
Page 281 and 282:
NUMBER 523 269 MATERIAL EXAMINED.
Page 283 and 284:
NUMBER 523 271 evenly spaced, simil
Page 285 and 286:
Page 287 and 288:
Page 289 and 290:
NUMBER 523 277 Peristomial cirri to
Page 291 and 292:
NUMBER 523 279 18+0, 12+12, and 1+1
Page 293 and 294:
NUMBER 523 281 than peristomium, as
Page 295 and 296:
Page 297 and 298:
NUMBER 523 285 167. Eunice reducta
Page 299 and 300:
NUMBER 523 287 articulations. Grube
Page 301 and 302:
Page 303 and 304:
NUMBER 523 291 antennae, without ar
Page 305 and 306:
Page 307 and 308:
NUMBER 523 295 Jaws not examined. B
Page 309 and 310:
NUMBER 523 297 posiscial lobes low,
Page 311 and 312:
NUMBER 523 299 ASSUMED STATES FOR P
Page 313 and 314:
Page 315 and 316:
NUMBER 523 303 smaller specimens. M
Page 317 and 318:
NUMBER 523 305 thereafter, always s
Page 319 and 320:
NUMBER 523 307 The ceratostyles lac
Page 321 and 322:
Page 323 and 324:
NUMBER 523 311 184. Eunice stigmatu
Page 325 and 326:
NUMBER 523 313 Eunice stigmalura is
Page 327 and 328:
NUMBER 523 31! .1 .5 a, b, c t n h
Page 329 and 330:
NUMBER 523 317 Hartman (1956, as E.
Page 331 and 332:
NUMBER 523 319 REMARKS.—Eunice th
Page 333 and 334:
NUMBER 523 321 without articulation
Page 335 and 336:
Page 337 and 338:
NUMBER 523 325 from setiger 5 throu
Page 339 and 340:
NUMBER 523 327 appendages long, kni
Page 341 and 342:
NUMBER 523 329 superior aciculae li
Page 343 and 344:
NUMBER 523 331 b. c, e, f, g, h. i,
Page 345 and 346:
NUMBER 523 333 paratype consisted o
Page 347 and 348:
Page 349 and 350:
NUMBER 523 337 always single (excep
Page 351 and 352:
Page 353 and 354:
NUMBER 523 341 1 .05 I 1 h a FIGURE
Page 355 and 356:
NUMBER 523 343 3 A of total peristo
Page 357 and 358:
NUMBER 523 345 Characters: 1, 2, 13
Page 359 and 360:
NUMBER 523 347 TABLE 21.—Comparis
Page 361 and 362:
Page 363 and 364:
Page 365 and 366:
Page 367 and 368:
Page 369 and 370:
Page 371 and 372:
Page 373 and 374:
Page 375 and 376:
Page 377 and 378:
Page 379 and 380:
Page 381 and 382:
Page 383 and 384:
Page 385 and 386:
Page 387 and 388:
Appendix A Character List for Eunic
Page 389 and 390:
NUMBER 523 377 4. digitiform 5. cla
Page 391 and 392:
NUMBER 523 379 5.03 13.U 14,U 15,3
Page 393 and 394:
NUMBER 523 381 16.07 16.08 16.09 17
Page 395 and 396:
NUMBER 523 383 28.01 biformicirrata
Page 397 and 398:
NUMBER 523 385 39.03 39.04 39.05 39
Page 399 and 400:
NUMBER 523 387 50.07 50.08 50.09 51
Page 401 and 402:
NUMBER 523 389 62.01 62.02 62.03 62
Page 403 and 404:
NUMBER 523 391 73.04 73.05 73.06 73
Page 405 and 406:
NUMBER 523 393 84.07 84.08 84.09 85
Page 407 and 408:
NUMBER 523 395 96.01 kerguelensis 9
Page 409 and 410:
NUMBER 523 397 107.04 25,3 26,3 27,
Page 411 and 412:
NUMBER 523 399 118.07 118.08 118.09
Page 413 and 414:
NUMBER 523 401 130.01 mutilata 130.
Page 415 and 416:
NUMBER 523 403 141.03 15.U 16,U 17.
Page 417 and 418:
NUMBER 523 405 152.05 35,5 36,28 37
Page 419 and 420:
NUMBER 523 407 164.07 164.08 164.09
Page 421 and 422:
NUMBER 523 409 176.01 savignyi 176.
Page 423 and 424:
NUMBER 523 411 187.04 24,U 25,4 26.
Page 425 and 426:
NUMBER 523 413 198.07 55,2 56- 58-
Page 427 and 428:
Appendix C Important publications o
Page 429 and 430:
NUMBER 523 417 1919c. Pacific Coast
Page 431 and 432:
NUMBER 523 419 Videnskabers Selskab
Page 433 and 434:
NUMBER 523 421 38:1-74,37 figures.
Page 435:
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