Index

α chains, 292–293 Amynthas spp., 269 Acanthobdella, 23, 117, 317 Amynthas tokioensis, 273 Acanthobdella peledina, 320–321, 342 Anaitides, 219 Acanthobdellida peledina, 320 Andiorrhinus amazonicus, 232 Acanthodrilidae, 258, 272 Anisochaeta spp., 268 acetylated tubulin (ACT), 159–160 Annelida, 4. See also Aciculata, 14, 16 and //nemertean clade, 5 , 4 cellular and behavorial properties of learning in, Acrocirridae, 22 135–150 acrosome tube, 314, 318, 324 culture techniques for, 48–50 ACT. See acetylated tubulin developmental events, heterochronic shift in, 79 actin, 172 diversity in cleavage of, 78–79 actin microfi laments, 121–122 in evo-devo biology, 65–82 adenosine triphosphate (ATP), 310–311 genetic and developmental tools for, 5–6 citrate lyase, 289 genomics, 108–109 afterhyperpolarization (AHP), 143 groups within, 78 agriculture, 274 hydrothermal vent, 287–298 AHP. See afterhyperpolarization life cycle cultures of, 47–59 albumenotrophy, 333 major lineages of, 47 Alciopidae, 212 and metazoan phylogeny, 67–68 algal “gardening” behavior, 214 as model organisms, 6–9, 23–24 Alitta virens, 206 criteria for selection of, 48 Allolobophora celtica, 274 molecular analysis of, 13–26 Allolobophora chlorotica, 39–40 molecular relationships with allies, 16–22 alpha-tubulin, 94 phylogeny of, 4–5, 13–26, 24–25 Alvinella pompejana, 211, 287–290, 341 molecular, 16–22 adaptation studies of, 297–298 problems inferring, 23–24 amino acid composition of, 293 radiation, 25 cellular temperature adaptation, 293–297 segmentation, molecular mechanism of, 73–74 chemical and thermal stability of, 291–299 studies of subtaxa relationships among, 16 temperature comfort of, 290–291 subtaxa, 23 tubes, 291–292 summary of representative studies of, 15 Alvinella spp., integument types in, 291 temperature adaption in, 290–291 Ampharetidae, 211 anoxia, 207 Amphinomidae, 22, 50, 211COPYRIGHTEDanterior-posterior MATERIAL (AP) axis, 70, 72, 186 Amphitrite spp., 213 polarity, 106–108 Amynthas, 275 Aphelochaeta sp., 212 Amynthas corticis, 273 Aplysia, 139, 143 Amynthas diffringens, 273 Aplysia californica, 5 Amynthas gracilis, 273 Aporrectodea caliginosa, 39–40, 237 Amynthas hawayanus, 273 Aporrectodea rosea, 39–40 Amynthas hendersonianus, 276 Aporrectodea turgida, 273 Amynthas minimus, 269 Archiannelida, 16

Annelids in Modern Biology, Edited by Daniel H. Shain Copyright © 2009 John Wiley & Sons, Inc.

347 348 Index

Arenicola marina, 207, 213–214, 221, 293 Caenorhabditis elegans, 3, 13, 23–24, 65, 68, 76, Arenicolidae, 16 81, 99, 108, 124, 150, 169, 179 Arthopoda, 4, 68, 81, 339 calcium sensors/signaling, 173, 174 astral microtubules, 122 Californian blackworm. See Lumbriculus variegatus ATP. See adenosine triphosphate Calopteryx, 35 Aulodrilus, 322 cAMP/PKA messenger pathway, 141 Autolytinae, 49 Canalipalpata, 14, 16 Autolytus, 49 Capilloventridae, 23 Autolytus brachycephalus, 56 Capitella capitata, 7, 16, 23–24, 50, 76, 89, 208, Autolytus prolifera, 56 216, 218 axial patterning, 4, 66, 77 chromosome numbers in, 90 life history traits in, 90–91 BAFs. See bioaccumulation factors mesoderm and nervous system of, 95–97 “bandlet,” 71, 121, 124–127 Capitella sp., 207, 209, 219 bar coding, 34, 110 Capitella sp. I, 5–8, 13, 23–24, 81, 104, 220 basal taxon, 66 advantages of, for evo-devo studies, 89 Begemius, 258, 272, 275 allozyme patterns and species designations, behavioral neurobiology, 82 89–90 bending, local, 166 colonizing polluted areas, 109 benthos, states of change in, 218–219 developmental stages in, 92, 94 BI. See biotic index evolution, development and ecology of origin of, bilateral organisms, clades of, 4–5 88–111 Bimastos parvus, 273 gene expression during gut development of, 98 Bimastos spp., 268, 273–274 metabolizing polycyclic aromatic hydrocarbons, bioaccumulation, 220 109 factors, 234 metamorphosis in, 93 test morphology, 91–92 Enchytraeid, 233–235 phylogeny of, 110–111 oligochaete, 233–235 vasa expression in, 101 sediment, 235–236 Capitella sp. II, 7 biodiversity comparisons, requirements of, Capitella sp. IIIa, 90 269–271 Capitellidae, 211 biomonitoring, 217–219 carabid beetles, 234 biotic index (BI), 217 carbendazim, 232 bioturbation, 214, 216 cathepsin L (Tt-catl), 177 blast cells, 71, 72, 126–127 cell division, asymmetric, 123–124 blastomeres, 74–78, 94, 117–118, 120 centipedes, 234 Boccardia hamata, 210 central arbors, 163–164 Boccardia proboscidea, 54 central nervous system (CNS) body shortening, 169–170 development of the, 157–159 Brachiopoda, 4–5 injury, 172–173 branch length, 24 leech, 156–180 branch point refl ection, 149 neuromeres in, 156 Branchiobdella italica, 321 and regeneration, 169–171 Branchiobdella pentodonta, 334 oligochaete, 187 Branchiobdellida, 320–321 central pattern generator (CPG), 144 Branchiomaldane vincentii, 208 amphioxus, 107 Branchiomma luctuosum, 215, 219 CGCs. See clitellar gland cells Branchiomma vesiculosum, 147 chaetal structures, 5 Branchiura sowerbyi, 235 Chaetopteridae, 16, 22–23, 211 BrdU, 73, 91, 95 Chaetopterus, 24, 76, 81, 104, 107 brooding behavior, 334–335 Chaetopterus variopedatus, 8–9, 13, 24, 207 , 4 Chaetozone setosa, 216 Index 349

Chironomidae, 230 consistency index (CI), 324 Chironomus riparius, 230 Cossura sp., 207 chitin, 291 cox1, 8, 14, 22–23 Chlamydomonas nivalis, 305 cox3, 14 Chordata, 4–5, 96 C-propeptide, 294 CI. See consistency index crack propagation, 215 ciliary band markers, 94 Crassiclitellata, 318 Ciona intestinalis, 5 Crassostrea virginica, 210 circumferential indentation, 167 Ctenodrilidae, 51 Cirratulidae, 16, 50–51, 215–216 Ctenodrilus, 16 Cirratulus cirratus, 209 Ctenodrilus serratus, 51, 220 Cirriformia, 219 ctenograths, 243 Cirriformia spirobrancha, 50–51 ctenophores, 4 cleavage, 120, 123–124 culture techniques, 48–50 equal, 75–76, 94 Cupiennius salei, 105 polar lobe-dependent, 76 Cupressus sempervirens, 276 spiral, 74–77, 117–121 cystatin B (Tt-cysb), 177 unequal, 76, 91, 117 cysteine-rich intestinal protein (CRIP), 172 clitellar gland cells (CGCs), 329–332 cytaster, 122 , 5, 14, 16, 22, 25, 48, 71–73, 78, 305 cytb, 14 assessing phylogenetic relationships in, 314–325 cytokinesis, 76 cocoons, 328–342 proinfl ammatory, 178 biomaterials applications for, 342 cytoplasm, 76 cryptic speciation in, 31–42 cytotoxic lymphocytes, 177 ectoparasitic, 320 examples of model organisms of, 34–36 Danio rerio, 3 hermaphroditic reproduction of, 314, 329–330 DD-PCR. See differential display-PCR as model organisms, 34–40 dendritic arbor, 156–158 morphology, 32–34 Dendrobaena attemsi, 274 patterns of spermatological characters in, Dendrobaena octaedra, 39–40, 274 324–325 Dendrobaena spp., 269 phylogeny of, 23, 318–319 Dendrodrilus rubidus, 39–40 polyploidy of, 33–34 deoxyribonucleic acid (DNA), 34 popularity of, for research, 31–32 RAPD, 40 reproductive biology in, 330 destabilase, 177–178 spermatozoal characters of, 315 Deuterostoma, 3–4, 67–70 spermatozoon, 324 developmental system drift (DSD), 126–127 variation, sources and kinds of, 32–34 5,7-DHT, 138–139 clitellum, 314, 328–331 Diachaeta, 258 Clymenella torquata, 14, 206, 213 Dichogaster curgensis, 277 cnidarians, 4 Dichogaster spp., 258, 269, 275 CNQX-sensitive synapse, 142 differential display-PCR (DD-PCR), 128 CNS. See central nervous system differentiation, 95, 96 cocoons 5,7-dihydroxytryptamine (5,7-DHT), Clitellata, 331–336 138–139 T. tessulatum, 336–338 Dinophilidae, 16 types of, 329 Dinophilus gyrociliatus, 51–52, 220 coelom, 25, 95, 176–177 Diopatra, 206, 210 Cognettia sphagnetorum, 228 dipteran insects, 70 COI. See genes, cytochrome oxidase I DM. See proteloblast, mesodermal DM collagen, 291–295 D-NAME, 171 Collembola, 304 DNOPQ. See proteloblast, ectodermal DNOPQ conduction, 149–150 Dodecaceria sp., 210 350 Index

Doliodrilus, 322 Embryology of Clepsine, The (Whitman), 116 “dorsal ,” 97 EML. See external mucous layer dorsal-posterior (DP) nerve, 157, 165 Enchytraeid, 233 Dorvillea, 219, 243–244 enchytraeid reproduction test (ERT), 233 Dorvillea articulata, 51 Enchytraeidae, 32–33, 37–39, 56–57, 301–311, Dorvillea (Schistomeringos) longicornis, 51 317–318, 330 Dorvilleidae, 49, 51, 211, 243–253 + Crassiclitellata, 23 double-P triplets (GPP), 293 in soil ecotoxicology, 237–238 DP. See dorsal-posterior nerve soil tests with, 232–235 D-quadrant, 75–77 Enchytraeus, 37–38 draft genome sequence, 108–109 Enchytraeus albidus, 33, 56–57, 233–234 Drawida barwelli, 273 genotypes for, 38–39 Drawida japonica, 275 Enchytraeus buchholzi, 38–39 Drawida spp., 269 Enchytraeus bulbosus, 38–39 Drosophila melanogaster, 3, 13, 23–24, 65, 68, 70, Enchytraeus christenseni, 38–39 73–74, 81, 96–97, 99, 101–103, 105–106, Enchytraeus coronatus, 95 108, 125, 150, 169, 177, 179 Enchytraeus crypticus, 38, 56, 233, 237 DSCAM, 161 Enchytraeus japonensis, 186 DSD. See developmental system drift Enchytraeus luxuriosus, 56, 234 Dunaliella sp., 49 Enchytraeus variatus, 38 endoderm formation, 4 earthworm, 7, 230–232 ENSO. See El Niño-Southern Oscillation index bioaccumulation test, 232 Enteromorpha intestinalis, 214 common, see megadrile oligochaete Enteromorpha spp., 49 cosmopolitan, 257–278 environmental risk assessment (ERA), 229–230 fi eld test, 231 epsilon proteobacteria, 289 megascolecid, 319 EPSP. See excitatory postsynaptic potential in soil ecotoxicology, 237 ERA. See environmental risk assessment standardized tests for, 231 Erpobdella, 333 “true,” 267 Erpobdella lineata, 334 , 3–4, 13, 67–70, 96 Erpobdella loctoculata, 321 Echinodermata, 4, 68 Erpobdellidae, 330, 334 Echiura, 5, 14, 16, 22, 25, 74, 78 Erpodbella punctata, 334, 336–337, 340 Eclysippe vanelli, 14 Errantia. See Aciculata “ecosystem engineers,” 228 EST. See express sequence tag ectodermal ablations, 159 estuaries, 206–207 EF-1α. See elongation factor-1α Eteone, 219 eggs Eucalyptus marginata, 269 hermaphroditic, 91, 100 Euclymene sp., 207 lecithotrophic, 91 Eukerria, 258 Eirene, 49 Eukerria saltensis, 269, 276–277 Eisenia albidus, 41, 234, 237 Eumida, 219 Eisenia andrei, 39–40, 230, 232, 237 Eunicidae, 16, 211, 244–253 Eisenia fetida, 37, 39–41, 57, 71, 186–187, 228, Eurythoe complanata, 50 230, 232, 237, 273, 330, 336 eusperm, 322 Eisenia fetida Eisenia andrei, 39–40 evolutionary development (evo-devo), 66–67, 79–80 Eisenia japonica, 272, 275 Exallopus, 244, 246 Eisenia koreana, 272 excitatory motor neurons (eMNs), 166–167 Eisenoides, 273 excitatory postsynaptic potential (EPSP), 142 El Niño-Southern Oscillation (ENSO) index, 311 Exogone lourei, 208 electrophoresis, 34, 35 express sequence tag (EST), 4–5, 22, 26, 66, 298 embryogenesis, 156, 167 external mucous layer (EML), 332, 336 leech, 118–120 extracellular matrices (ECMs), 290–291 Index 351

Fabricia berkeleyi, 208 expressing, 128–130 Faivre’s nerve, 140 foxA, 99 feeding, 213, 216 “gap,” 73–74, 102–103 Ficopomatus enigmaticus, 209 GATA1/2/3, 97–99 Flabelligeridae, 16, 22 GATA4/5/6, 97–98 fl atworms, 47, 68, 74 GATAb, 98 foldases, 173 Gsx, 107 Fridericia, 39 hairy, 105–106 Hau-Pax3/7A, 125 Galathealinum brachiosum, 14 hedgehog (hh), 102–105 Galeolaria caespitosa, 48 homeobox, 107 gamasid mites, 234 “housekeeping,” 66 gamma-tubulin, 123–124 Hox, 66, 101–102, 106–107, 124–125, 128 gastrulation, 78 Hox1-Hox5, 107 genes Hro-nos, 125 18S rRNA (18S), 14, 18–20, 22–23, 25, 34, hunchback (hb), 102–103 318–319 K110, 130 + 28S rRNA + EF-1α, 21, 25 mitochondrial (mtDNA), 14–16, 31, 40 analysis of, 17–20 mitochondrial COI, 34, 37 28S rRNA (28S), 22, 25, 34 mt16S rRNA (16S), 8, 14, 22–23, 34–35, 37 brachyury, 108 nad1, nad6, 14 “candidate” approach, 73–75, 128 nanos, 125 CapI-Cdx, 100, 107 Notch, 105–106 CapI-Delta, 105 nuclear, 14, 40 CapI-en, 103 P450 (CYP), 109 CapI-foxA, 99 pair-rule, 102–103 CapI-gataA1, -gataB1, -gataB2, -gataB3, 99 ParaHox, 107–108 CapI-Gsx, 107 pax3/7, 103 CapI-hes1, -hes2, -hes3, -hesr1, -hesr2, 105 Pdu-en, 104 CapI-hh, 100, 103 Pdu-vasa, 100 CapI-nanos, 100 Pdu-wnt1, 104 CapI-Notch, 105 ProtoHox, 107 CapI-sna1, 100 runt, 103 CapI-twt1 and twt2, 100 segment polarity, 102–104 CapI-vasa, 100 snail, 96 CapI–wnt1, 99–100, 103 translocations of, 15–16 CapI-Xlox, 100, 107 wingless (wg), 102–105 Cdx, 107 wnt, 125 CH-Hox2, 107 Xlox, 107 claudal, 108 genetics, 5–6 CYP4AT, 109 germinal band, 71 CYP331, 109 germinal plate, 71 cytochrome b (cytb), 14 GFs. See giant fi bers cytochrome c oxidase subunit I (cox1), 8, 14, giant fi bers (GFs), 187 22–23 glacier ice worms, 301–311. See also cytochrome c oxidase subunit III (cox3), 14 Mesenchytraeus solifugus; Mesenchytraeus cytochrome oxidase I (COI), 31, 110, 127–128, solifugus rainierensis; Sinenchytraeus 244 glacialis Delta, 105–106 adenylate profi les of, 310 developmental regulatory, 66, 75 clades, 307–309 elongation factor-1α (EF-1α), 14, 22, 25 classifi cation and phylogenetic relationships of, engrailed (en), 73–74, 102–105, 125, 128 305–306 even-skipped, 103, 108, 125 conservation status of, 311 352 Index glacier ice worms (cont’d) Hesiolyra bergi, 297 density of, 302–305 Hesionidae, 211 natural history of, 303–305 Heteromastus, 219 origins of, 306–307 Heteromastus fi liformis, 213 phylogeny of, 307 Heteroporodrilus, 258 physiology of, 309–311 hexachlorobenzene (HCB), 234–235 Glossiphonia, 335 Higher-Tier Aquatic Risk Assessment for Pesticides Glossiphonia complanata, 118 (HARAP), 238 Glossiphoniidae, 6, 58–59, 71, 117, 330, 334 Hirudinidae, 6–7, 59, 117, 317, 320 brooding behavior in, 334–335 leeches, 156 Glossoscolecidae, 232 life cycle cultures of, 58–59 glutamine (trnQ), 14 Hirudo medicinalis, 6–7, 13, 23, 40, 50, 59, 82, Glycera dibranchiata, 48, 209–210, 221 107, 117, 135–137, 139–140, 160, 163, glycosylation, 243, 341 165–166, 169, 186, 330, 333 mannosidic, 163–164 segmental nerves in, 158–159 Glyphidrilus, 275 Hirudo spp., 23–24, 333 Glyphidrilus kuekenthali, 275 Hirudo verbana, 7, 40 Glyphidrilus stuhlmanni, 275 histone H3, 14, 22 Gly-X-Y amino acid triplets, 292–294 Hm-inx1, -inx2, -inx5, -inx6, -inx8, 169 Goniada, 219 HmLAR1, -2, 161–163 Gordiodrilus elegans, 269 HNK-1, 94 Grania, 32 Hodgkin, Alan, 117 growth zones, 95 Hrabeiella periglandulata, 206 guanylate cyclase inhibitor, 171 HSPs. See heat shock proteins 5-HT. See serotonin

Haementeria, 334 5-HT7, 141 Haementeria ghilianii, 6, 139, 333 Hudineans, sperm ultrastructure in, 320–321 Haemopis spp., 117 human migration routes, 275 Haliotis rufescens, 210 Huxley, Andrew, 117 Halosydna, 208 HXXE divalent ion, 177 Halosydna brevisetosa, 53 Hydra, 185 Halosydna johnsoni, 53, 208 Hydra magnipapillata, 97 Haplotaxidae, 318–319 Hydra vulgaris, 97 HARAP. See Higher-Tier Aquatic Risk Assessment Hydroides dianthus, 209 for Pesticides Hydroides elegans, 8–9, 53, 81, 94, Harmothoë imbricata, 48, 206 103–104 HCB. See hexachlorobenzene Hydroides norvegica, 208 heat shock proteins (HSPs), 297–298 hydrothermal vent, 211 Helobdella, 122, 335 communities, 287–298 Helobdella europaea, 128 hymenopteran insects, 70 Helobdella papillornata, 329 Helobdella robusta, 5–7, 13–14, 40, 72, 77, 82, Ilyanassa, 6 101, 104, 109–110, 122–123, 126–128, 169, Ilyanassa obsoleta, 75–76, 125 179 Ilyodrilus, 33 Helobdella spp., 7, 23–24, 40–41, 50, 58, 65, 71, immune responses, in leech nervous system, 76, 81–82, 117–118, 125, 334 156–180 Helobdella stagnalis, 127, 330, 335 immunity, 176–178 Helobdella triserialis, 6–7, 58, 107, 126–128 immunoreactivity, cGMP, 171 Helodrilus oculatus, 269, 274 iMN. See inhibitory motor neuron hemerythrin message, 175 infaunal macrobenthos, 216 Hemichordata, 4–5 inhibitory motor neuron (iMN), 165–167 hermaphrodites, 90–91, 118 Insulodrilus bifi dus, 314 Hesiocaeca methanicola, 212 spermatozoa, 316 Index 353 intercellular communication, 169 local bend interneurons (LBIs), 166–168 interleukin-1β, 178 Loligo pealei, 6 interleukin-6, 178 long-branch attraction (LBA), 15, 22, 24 internal transcriber spacers (ITS1–2), 34 long-term depression (LTD), 148–149 International Code of Zoological Nomenclature long-term potentiation (LTP), 148–149 (ICZN 1999), 278 Lopadorhynchidae, 212 interneuron S, 170 Lophotrochozoa, 3–9, 67–70, 74 intertides, 207 considerations for a model system for, 4–5 introns, 26 early diversifi cation of, 4 introsperm, 314 genetic and developmental tools for, 5–6 Iospilidae, 212 model taxa, 13 Iphitime, 244 phylogenetic consderations for, 4–5 Iphitime paguri, 244–246 Lottia gigantea, 5 Iphitimidae, 244 LSU. See large ribosomal subunit Isochrysis sp., 49 LTD. See long-term depression isolates, tools to compare, 42 LTP. See long-term potentiation “ITS region,” 34 Lumbricidae, 39–40, 57, 186, 230, 267, 271–274 ITS1–2. See internal transcriber spacers soil tests with, 230–232 Lumbriculidae, 23, 37, 186, 317, 320 Janua (Dexiospira) brasiliensis, 54 bioaccumulation test, 235–236 Lumbriculus spp., 37 karyotypes, 90 Lumbriculus variegatus, 33, 37, 41, 58, 140, 186, kinases, 173 228, 236, 310 , 4 asexual reproduction of, 191–192 life cycle cultures of, 192 lambda-cyhalothrin, 232 as a model, 185–199 Lamellibranchia cf. luymesi, 211 and PCP toxicity, 235–236 large fat cells (LFCs), 177 respiratory apparatus of, 190 large ribosomal subunit (LSU), 4 Lumbricus rubellus, 7, 232 learning, nonassociative, 146–147 Lumbricus spp., 24 leech, 6–7, 40, 48, 78 Lumbricus terrestris, 7, 13–14, 228–229, 237, cellular and behavorial properties of learning in, 272 135–150 Lumbrineridae, 16 differentiation in, 116–130 Lumbrineris, 208, 219 immune responses of nervous system of, 156–180 ”lumping,” 32 innexins, 169 Lymnaea, 140 sanguivorous, 332 Lymnaea stagnalis, 75–76, 174 swim circuit, 144–146 Lysidice collaris, 208 touch-based swim response, 145 Lysidice ninetta, 208 whole-body shortening and the s interneuron, 137–139 mAbs. See monoclonal antibodies leucopoiesis, 177 macromeres, 77–78, 121, 122 Leucosticte tephrocitis, 306–307 magelonids, 215 leukemia, 6 malate dehydrogenase-1, 90 Limnodriloides monothecus, 322–323 Maldanidae, 211, 216 Limnodriloidinae, 317, 321–323, 322 Manayunkia speciosa, 206 Limnodrilus hoffmeisteri, 35–36, 41 Marphysa leidi, 221 Limnodrilus sp., 33, 95 Marphysa sanguinea, 221 lindane, 234 Marsupiobdella africana, 335 lineage tracer, intracellular, 73 matrix-assisted laser desorption / ionization linkage, M, N, O, P, Q, 125–128 (MALDI), 179–180 litterbag test, 231 mechanoreceptor, T, 146 L-NAME, 171 Mediomastus sp., 207–208, 212, 219 354 Index megadrile oligochaete, 330 monoclonal antibodies (mAbs), 157, 160 in archeology and human history, 273–277 Monticellina spp., 206, 217 biodiversity of, 268–273 Mooreonuphis stigmatis, 208 characteristics of, 268 morphallaxis, 186–187, 192–198 cosmopolitan, 258–278 morphology exotic, 268–273 and clades, 13–14 global and historical perspective on, 257–278 plasticity of, 25, 35 introduced exotic, 258 MPGZ. See mesodermal presumptive growth zone nonendemic species of, 258–266 mRNA, 123 number of species of, 267–273 Murchieona minuscula, 269 population, components of, 258 Mus musculus, 3 resident native, 258 mussel beds, 207–208 translocated native, 258 Mya arenaria, 6 transportations, 273–279 myohemerythrin, 172 Megascolecidae, 258, 267, 271–272 myomodulin, 141, 143 Megascolides australis, 331 Mytilus edulis, 208 Melanenchytraeus, 305 Myxobolus cerebralis, 35 Mesenchytraeus altus, 301 Myzostoma seymourcollegiorum, 14 Mesenchytraeus armatus, 306 myzostomids, 5, 14 Mesenchytraeus fl avus, 306 Mesenchytraeus gelidus, 301, 305–307 nad1, nad6, 14 Mesenchytraeus hydrius, 301 Naididae, 23, 57, 186, 330 Mesenchytraeus niveus, 305 Naidinae, 33, 41, 317 Mesenchytraeus pedatus, 306–307 Neanthes acuminata, 52 Mesenchytraeus solifugus, 39, 55, 305, 307–310 Neanthes arenaceodentata, 52, 220–221 Mesenchytraeus solifugus rainierensis, 305, Neanthes caudata, 52 309 Neanthes lighti, 207 mesodermal presumptive growth zone (MPGZ), Neanthes limnicola, 52 100–101 Neanthes spp., 219 metamerism, 70 Neanthes succinea, 52 metamorphosis, 94 Nematoda, 4, 68, 81, 339 Metaphire, 275 Nematonereis unicornis, 208 Metaphire hilgendorfi , 273 Nematostella vectensis, 97, 99 metatrochophore, 93 CapI-Post1, 106 gut formation in, 97–100 , 4–5, 74 methylene blue, 171 Nephasoma minuta, 108 methysergide, 146 Nepthys spp., 14, 219 Michaelsen, W., 267 Nereididae, 8, 23, 52–53, 206, 211–212, 214 Microdorvillea, 244 nereids, 208 microdriles, 330 Nereis crigognatha, 52 microglial cells, 171–172 Nereis diversicolor, 214, 221 micromeres, 75, 95 Nereis grubei, 52 Microscolex dubius, 258 Nereis spp., 8, 76, 213, 219 Microscolex kerguelarum, 258 Nereis succinea, 206 Microscolex macquariensis, 258 Nereis vexillosa, 214 Microscolex spp., 258 Nereis virens, 107, 210, 214 microtubules, 122 Nerillidae spp., 16, 211, 212 mitochondria, 314–317, 324 nerve injury, response to, 171–172 model organisms, 24, 31 netrin, 162 model system, choosing, 3–9 receptor unc-5, 163 model waxing, for study, siRNA, 163 88–111 neurohemerythrin, 173–174 , 4–5, 47, 68, 74 neuroimmunity, 173–176 Moniligastridae, 272 neuronal circuits, 165–169 Index 355 neuronal formation, 4 Ophryotrocha spp., 8, 13, 23, 49, 211–212, 220 neurons characteristics of, 243–244 AE and AP, 165 described, 245 CD8+ cells, 176 ecology of, 250–252 L, 137, 143 evolution and ecology of, 243–253 mechanosensory, 137 phylogeny of, 247 NK-like, 176 reproductive biology of, 247–250 nociceptive (N), 137, 144 of, 243–247

PD, 160–161 undescribed, 246 pressure (P) sensory, 137, 144, 147, 163, Orbinia latreillii, 14 166–168 Orbiniidae, 14, 16, 22

PV, 161–163 Osedax spp., 210–211 S, 147 OTU. See operational taxonomic unit touch (T) sensory, 137, 144, 163 Ougia spp., 244 neuropil, 163–164 Oweniidae, 22 nitric oxide (NO), 171–172, 175 2-oxoglutarate:acceptor oxidoreductase, 289 NMDA-R, 148–149 oxygen minimum zones (OMZs), 206 notch signaling pathway, 105–106 nuclear factor κB, 178 PAHs. See polycyclic aromatic hydrocarbons Palola viridis, 221 OBT. See oligochaete bioaccumulation test Palpata, 14 Ocnerodrilus occidentalis, 269 Paralvinella sulfi ncola, 287, 297 Octochaetidae, 258, 272 Paranais litoralis, 9, 57 Octochaetus ambrosensis, 258 Paraonidae, 211 Octodrilus spp., 276 Paraonidae sp., 212 octopamine, 149 Parapionosyllis sp., 208 Oligochaeta, 48, 71–72, 78, 97, 330 Parapodrilus, 244 central nervous system (CNS) of, 187–189 parasperm, 322 for ecotoxicological assessment of soil and Parergodrillidae, 22 sediment, 228–238 Parergodrilus heideri, 206 escape refl ex anatomy, 188–189 Parophryotrocha, 244 life cycle cultures of, 56–58 Parougia, 244 neural regeneration in, 187 parthenogenesis, 268 in sediment ecotoxicology, 238 Patella vulgate, 75–76 Onuphidae, 214 Pax-6, 66 Onuphis teres, 221 PCP. See pentachlorophenol operational taxonomic unit (OTU), 24 PCs. See precursor cells opercula, 329, 334, 338–339, 342–343 PEA. See proenkephalin A Ophryotrocha, 243–253 PEC. See predicted environmental concentration Ophryotrocha adherens, 251–252 Pectinaria koreni, 214 Ophryotrocha akessoni, 252 Penaeus kerathurus, 221 Ophryotrocha alborana, 246 Penaeus vannani, 221 Ophryotrocha cf. vivipara, 251 pentachlorophenol, 232 Ophryotrocha claparedii, 251 peptide B, 177 Ophryotrocha diadema, 8, 51, 95, 246, 250 perforatorium, 314 Ophryotrocha gracilis, 246–248, 250 Perinereis nuntia, 221 Ophryotrocha hartmanni, 244–248, 251 Perionyx excavatus, 14 Ophryotrocha labidion, 243 peripheral nervous system (PNS), 156 Ophryotrocha labronica, 8, 24, 246, 248, 252 and DP nerve formation, 160 Ophryotrocha mandibulata, 243 and ectopic ganglia, 160–161 Ophryotrocha platykephale, 252 genesis of, 159–160 Ophryotrocha puerilis, 8, 250, 252 P-neuron arbor of, 161–163 Ophryotrocha sp. japonica, 252 target, disruption of, and abnormal central Ophryotrocha spatula, 252 projections, 165 356 Index

PGCc. See primordial germ cells shell-boring, 210 Pharmacopia californica, 339 skeletons, 210–211 Phascolion strombus, 107 and toxicological testing, 219–221 Phascolopsis gouldii, 14 Polychaete, 291, 342 phenotype, and developmental change, 66 polycyclic aromatic hydrocarbons (PAHs), 216 Pheretima, 275, 276 Polydora, 219 , 5 Polydora ciliata, 55, 209 phosphoglucomutase, 89 Polydora commensalis, 210 phosphohexose isomerase, 90 Polydora concharum, 210 Phreodrilidae, 314, 318–319 Polydora cornuta, 55 Phyllodocidae, 211 Polydora ligni, 55, 208, 210 phylogenetic analysis, characters for, 15 Polydora socialis, 210 phylogeny Polydora spp., 210 metazoan, typical, 68–69 Polydora websleri, 210 molecular, 67 Polygordiidae, 16 Annelida, 16–22 Polynoidae, 53, 206, 208, 211 Piscicola geometra, CGC types in, 331 Polypheretima, 275 Piscicolidae, 330 Polypheretima brevis, 275 Pisione, 16 Polypheretima elongata, 276, 277 Pista cristata, 14 Polypheretima pentacystis, 275 Placobdella papillifera, 331–332 Polypheretima voeltzkowi, 276 Placopeclen magellanicits, 210 polyploidy, 33–34, 39–41, 268 plankton, 212 Pomatoceros triqueter, 54 platyhelminthes, 5 Pompeii worm. See Annelida pompejana Platynereis, 8, 24, 219 Pontodoridae, 212 Platynereis bicanaliculata, 208, 214 Pontodrilus, 258 Platynereis dumerilii, 5, 8, 13–14, 24, 53, 75–76, Pontodrilus litoralis, 258 81, 97, 99–100, 102, 104–105, 107–109 Pontoscolex, 258 germ cell specifi cation and differentiation in, Pontoscolex corethrurus, 258, 272, 277 100–101 Potamothrix, 33 vasa expression in, 101 PPCs. See pro-precursor cells Platynereis massiliensis, 53 precursor cells (PCs), 331 , 4 predicted environmental concentration (PEC), Playthelminthes, 4 229–230 Poebius, 16 , 4 Poeobiidae, 212 primordial germ cells (PGCs), 125 Pogonophora, 5, 22, 25, 78 Prionospio, 219 pollution, 215, 218, 219, 222 Pristina leidyi, 57, 104, 186 Polychaeta, 5, 14, 22, 25, 47–48, 71–72, 78, 267 proenkephalin A (PEA), 177 and cold methane environments, 211–212 proline hydroxylation, 293 economic importance of, 221 Propappidae, 23 as environmental indicators and mediators, Propappus, 317 214–217 Propappus glandulosus, 317–319 in environmental studies, 205–222 Propappus Volki, spermatozoon of, 317–319 feeding guilds, 213 pro-precursor cells (PPCs), 331 gonochoristic, 212 proteins, 291 habitats and demographics of, 205–206 cytoskeletal and metabolic, 173 holopelagic, 212 gliarin, 173 importance of, 205–206, 222 immunity and regeneration, 176 life cycle cultures of, 50–56 modulated after bacterial challenge, 174 metals toxic to, 219–220 ReN3, 172–173 model, waxing, 88–111 proteloblast, 122–124 selected metals and, 220 Protodrilidae, 16 Index 357

Psammoryctides, 33 Sabellaria cementarium, 76 Pseudonereis, 208 Sabellidae, 16, 22, 209, 211 Pseudophryotrocha, 244 Saccocirridae, 16 Pseudopolamilla reniformis, 210 Salmacina (or Filograna) spp., 54 Pygospio, 219 Satchellius mammalis, 274 Pygospio elegans, 55, 209, 214 scanning electron microscopy (SEM), 340 Schistomeringos rudolphii, 252 Questa, 16 Schistosoma mansoni, 6 Schmidtea mediterranea, 5 receptor protein tyrosine phosphatase (RPTP), Scolecida, 14, 16 161 Scolelepis, 219 redox potential discontinuity (RPD), 219 Scoloplos, 208 regeneration, 71–72, 140, 169–170 Scoloplos armiger, 14 and leech, 185–186 Scoloplos robustus, 210 in Capitella spp., 92–93 sea grasses, 208 epimorphic, 186 Seepiophyla jonesi, 211 in Lumbriculus veriegatus, 185–199 segmental circuitry, and local bending, 166 morphallaxis, 186–187 segmental nerves, 159–160 neuronal, 169–170, 172 segmentation, 24–25, 68–74, 88, 97 segmental, 186 Annelida, 101–102 reproduction, 41, 48 cellular mechanism of, 71–73 in Clitellata, 330 molecular mechanism of, 73–74 fi ssion, 330 CapI-hbnl, 97 fragmentation, 330 CapI-hes2, 97 gonochoristic, 248, 250 Capl-hes3, 97 hermaphrodic, 330 de novo, 72 sequential, 248, 250 engrailed, 97 simultaneous, 248–250 evolution origin of, 70–71 in Lumbriculus veriegatus, 185–199 homologous, 70 parthenogenesis, 330 hunchback , 97 self-fertilizating, 118, 330 HyAlx, 97 retention index (RI), 324 interpreting confl icting comparative data about, Retzius cells, 141–143, 146–147 104–106 reverse tricarboxylic acid (rTCA) cycle, 289 leech, ganglion, 156 Rhododrilus, 258, 272 muscle, 95 Rhododrilus edulis, 276 periodicity in, 71 Rhododrilus kermadecensis, 276 , 70 Rhododrilus queenslandicus, 276 sensitization, 141, 146–147, 170 Rhodomonas sp., 49 “sensory tile,” 161–163 Rhyacodrilus, 33 serotonin (5-HT), 138, 141–144, 146–147, 149 RI. See retention index (RI) Serpula narconensis, 209 , 14, 287, 289, 293–295, 296 Serpulidae, 16, 53–54, 209, 211 rough endoplasmic reticulum protein 1 (RER-1), shortening, whole-body, 138 172 Siboglinidae, 5, 14, 16, 22–23, 25, 211 RPD. See redox potential discontinuity , 5, 14, 16, 22, 25, 74 siRNA, 173 16S, 8, 14, 22–23, 34–35, 37 small subunit (SSU), 4 18S rRNA (18S), 14, 18–20, 22–23, 25, 34, Smithsonidrilus spp., 322 318–319 snowfl eas, 304 S cell. See S interneuron Solea senegalensis, 221 S interneuron, 137–144 Solea vulgaris, 221 Sabella pavonina, 186 Solenopsis richteri, 207 Sabella spallanzanii, 215, 219–220 solifugus, 304 358 Index speciation, in Clitellate model organisms, Terebellides stroemi, 14 31–42 Terebrasabella heterouncinata, 210 species terrestrial model ecosystems (TMEs), 237 novel, using as models, 47 Thalassia testudinum, 208 number of, in , 47 Thalassodrilides, 322 opportunistic, characteristics of, 218 theromacin, 177 specifi cation Theromyzon rude, 127 axis, 74–77 Theromyzon spp., 58–59, 117–118, 122 gene expression during, 95–100 Theromyzon tessulatum, 6, 58, 81, 104, 126, spermathecae, 330 128–130, 177, 334–340 spermine NONOate (SPNO), 171 cocoon, 338–341 Spio spp., 209 CGC types in, 330–333 Spionidae, 49, 54–56, 210–211, 219 segmental nerves in, 159 , 68, 73–77 Theromyzon trizonare, 329 Spirobranchus corniculatus, 48 Tierreich, Das: Vermes (Michaelsen), 267 Spirorbis spirorbis, 206 thioredoxin (TRX), 172 “splitting,” 32 TMEs. See terrestrial model ecosystems springtails, 304 Tomopteridae, 212 SSU. See small subunit toxic concentration (TC), 229 Stauronereis ruldolphi, 51 toxic exposure ratio (TER), 229 stem cell transposable elements, 26 cytoplasmic rearrangement of, 121 trichobranchids, 215–216 development, 117–121 Trilobodrilus axi, 206 differentiation, 124–125 triploblasty, 95 genesis, 116–130 Tritonia diomedea, 6 and development, 117–120 trochophore larva, 88, 93 factors affecting, 121–124 trotifers, 47 research, tenets of, 116 Tt-catl. See cathepsin L Sternaspidae, 215 Tt-cysb. See cystatin B Sternaspis, 216 Tubifex hattai, 57–58, 126, 337 Sternaspis scutata, 207 Tubifex spp., 117 strand asymmetry, 15 Tubifex tubifex, 8, 9, 32–33, 35, 41, 57, 65, 71, Streblospio, 219 76–77, 81, 107, 124, 218, 235, 330, Streblospio benedicti, 55–56, 206, 210 336 Stygocapitella subterranean, 206 neighbor-joining tree for, 36 subtidal benthic communities, 209–212 Tubifi cidae, 23, 34–35, 41, 57–58, 318–319, 330 “subzero effect,” 144 bioaccumulation test, 235–236 sulfated glycosaminoglycans, 291 physiology of, 320–323 swim circuit, 168–169 sperm ultrastructure in, 320–322 Syllidae, 16, 56, 212 Tubifi cinae, 317, 322 synapsin, 172 tubulin, 172 synaptic plasticity, in leech CNS, 147–150 tumor necrosis factor-α, 178 28S, 22, 25, 34 Taenia solium, 5 Typhloscolecidae, 212 tandem cystatin B (Tt-cysb)–cathepsin L (Tt-catl), Typosyllis pulchra, 56 177 Typosyllis spp., 208 TC. See toxic concentration Tcp. See Theromyzon tessulatum, cocoon protein U2 snRNA, 22 (Tcp) Ulva sp., 49 teloblasts, 71–73, 95, 120–121, 126, 156 Urechis caupo, 14 teloplasm, 121–123 TER. See toxic exposure ratio vent shrimps, 298 Terebellida, 14, 215–217 ventral nerve cord (VNC), 187 Index 359 vermes, sex among, 100–101 Xenopus laevis, 96 Vibrio alginolyticus, 215 , 4 VNC. See ventral nerve cord zebra fi sh, 179 Whitman, Charles O., 116, 121, 124 Zostera noltii, 214