À.Â. ÀÄÐÈÀÍÎÂ, Â.Â. ÌÀËÀÕÎÂ

ÃÎËÎÂÎÕÎÁÎÒÍÛÅ ×ÅÐÂÈ (CEPHALORHYNCHA) ÌÈÐÎÂÎÃÎ ÎÊÅÀÍÀ (Îïðåäåëèòåëü ìîðñêîé ôàóíû)

A.V. ADRIANOV & V.V. MALAKHOV CEPHALORHYNCHA OF THE WORLD OCEAN

KMK Scientific Press Ltd. Ìîñêâà  1999 ÓÄÊ 595.13:591.4:575.86

À.Â. Àäðèàíîâ, Â.Â. Ìàëàõîâ. Ãîëîâîõîáîòíûå ÷åðâè (Cephalorhyncha) Ìèðîâîãî Îêåàíà (Îïðåäå- ëèòåëü ìîðñêîé ôàóíû). Ìîñêâà: Òîâàðèùåñòâî íàó÷íûõ èçäàíèé ÊÌÊ. 1999. 328 ñòð. A.V. Adrianov & V.V. Malakhov. Cephalorhyncha of the World Ocean. Moscow: KMK Scientific Press Ltd. 1999. 328 pp.

Ìîíîãðàôèÿ ïðåäñòàâëÿåò ñîáîé ïåðâóþ â ìèðîâîé ëèòåðàòóðå ïîëíóþ ñâîäêó ïî ìîðôîëîãèè è ñèñòåìàòèêå ìîðñêèõ ãîëîâîõîáîòíûõ ÷åðâåé — ïðåäñòàâèòåëåé òèïà Cephalorhyncha: ïðèàïó- ëèä, ëîðèöèôåð, êèíîðèíõ, ìîðñêèõ íåìàòîìîðô (âîëîñàòèêîâ). Ïðèâåäåíû ïîäðîáíûå îïðåäåëè- òåëüíûå òàáëèöû êàê íàäâèäîâûõ òàêñîíîâ, òàê è âèäîâ; äëÿ êàæäîãî òàêñîíà äàíû ïîäðîáíûå äèàãíîçû; äëÿ êàæäîãî âèäà ïðèâåäåíû ïîäðîáíûå èëëþñòðàöèè îïðåäåëèòåëüíûõ ïðèçíàêîâ, ýêî- ëîãè÷åñêèå è áèîãåîãðàôè÷åñêèå õàðàêòåðèñòèêè. 1 ñåìåéñòâî, 1 ðîä è 14 âèäîâ îïèñàíû êàê íîâûå äëÿ íàóêè. Ïîäðîáíî îïèñàíû íàõîäêè ðÿäà ðàíåå èçâåñòíûõ âèäîâ. Ìîíîãðàôèÿ áîãàòî èëëþñòðèðîâàíà ðèñóíêàìè è ìèêðîôîòîãðàôèÿìè. Äëÿ çîîëîãîâ, ãèäðîáèîëîãîâ è ýêîëîãîâ, àñïèðàíòîâ è ñòóäåíòîâ. Òàá. 8, Èë. 193, Áèáë. 431.

The monograph represents the first treatise on both morphology and taxonomy of marine cephalorhynch worms (phylum Cephalorhyncha): Priapulida, Loricifera, Kinorhyncha, and marine Nematomorpha. Identification keys and diagnoses to all taxa are provided. Detailed illustartions of distinguishing characters, as well as data on ecology and distribution are given for all species. One family, one genus, and 14 species are described as new for science. New records of previously known species are described in details. The monograph is abundantly illustrated by drawings, SEM- & TEM photos. Destined to zoologists, hydrobiologists and ecologists, postgraduate and graduate students. 8 tabs., 193 figs., 431 references.

ISBN 5-87317-067-3  À.Â. Àäðèàíîâ, Â.Â. Ìàëàõîâ, òåêñò, èëëþñòðàöèè  Ê.Ã. Ìèõàéëîâ, îðèãèíàë-ìàêåò  KMK Scientific Press Ltd., èçäàíèå Ëîäåéíèêîâ ñêëîíèëñÿ íàä ëèñòàìè, È â ýòîò ìèã ïðèâèäåëñÿ åìó Îãðîìíûé ÷åðâü, æåëåçíûìè çóáàìè Ñõâàòèâøèé ëèñò è ïðÿíóâøèé âî òüìó. Òàê âîò îíà, ãàðìîíèÿ ïðèðîäû, Òàê âîò îíè, íî÷íûå ãîëîñà! Òàê âîò î ÷åì øóìÿò âî ìðàêå âîäû...!

Í. Çàáîëîöêèé, “Ëîäåéíèêîâ” PHYLOGENY, AND CLASSIFICATION PRIAPULIDA: STRUCTURE, DEVELOPMENT,

KMK LTD. ÌÎÑÊÂÀ 1996 3 ÑÎÄÅÐÆÀÍÈÅ

1. ÏÐÅÄÈÑËÎÂÈÅ ...... 12 2. ÂÂÅÄÅÍÈÅ. Ïëàí ñòðîåíèÿ ãîëîâîõîáîòíûõ ÷åðâåé ...... 13 3. ÏÐÈÀÏÓËÈÄÛ ...... 14 3.1. Èñòîðè÷åñêèé îáçîð ñèñòåìû ïðèàïóëèä ...... 14 3.2. Ïëàí ñòðîåíèÿ è îñîáåííîñòè ìîðôîëîãèè, èñïîëüçóåìûå â ñèñòåìàòèêå ...... 17 3.2.1. Ïëàí ñòðîåíèÿ, âíåøíÿÿ ìîðôîëîãèÿ è ñèììåòðèÿ ...... 17 3.2.2. Ìèêðîñêîïè÷åñêàÿ àíàòîìèÿ ...... 32 3.2.3. Ëè÷èíêè ïðèàïóëèä ...... 32 3.3. Ôèëîãåíèÿ ïðèàïóëèä ...... 40 3.4. Ñèñòåìà, îïðåäåëèòåëüíûå òàáëèöû è äèàãíîçû òàêñîíîâ ...... 40 Êëàññ Priapulida ...... 40 Êëþ÷ äëÿ îïðåäåëåíèÿ íàäâèäîâûõ òàêñîíîâ ïî âçðîñëûì ôîðìàì ...... 41 Êëþ÷ äëÿ îïðåäåëåíèÿ ëè÷èíîê...... 41 Îòðÿä Priapulomorpha ...... 42 Ñåìåéñòâî Priapulidae ...... 42 Ðîä Priapulus ...... 42 Ðîä Acanthopriapulus ...... 47 Ðîä Priapulopsis ...... 48 Ñåìåéñòâî Tubiluchidae ...... 49 Ðîä Tubiluchus ...... 49 Îòðÿä Meiopriapulomorpha ...... 52 Ñåìåéñòâî Meiopriapulidae ...... 52 Ðîä Meiopriapulus...... 52 Îòðÿä Halicryptomorpha ...... 52 Ñåìåéñòâî Halicryptidae ...... 52 Ðîä Halicryptus ...... 52 Îòðÿä Seticoronaria ...... 53 Ñåìåéñòâî Chaetostephanidae ...... 53 Ðîä Maccabeus ...... 53 3.5. Çîîãåîãðàôèÿ ...... 54 3.5.1. Ìàêðîáåíòîñíûå ïðèàïóëèäû ...... 54 3.5.2. Ìåéîáåíòîñíûå ïðèàïóëèäû ...... 57 3.5.3. Ãèïîòåçà ãëÿöèàëüíûõ ðåëèêòîâ ...... 58 3.6. Summary of chapter 3 ...... 59 3.6.1. Classification of the class Priapulida ...... 59 Class Priapulida ...... 59 Key to superspecies taxa (adults) ...... 59 Key to superspecies taxa (larvae) ...... 59 Order Priapulomorpha ...... 60 Family Priapulidae ...... 60 Genus Priapulus ...... 60 Genus Acanthopriapulus ...... 61 Genus Priapulopsis ...... 62 Family Tubiluchidae ...... 62 Genus Tubiluchus ...... 62 Order Meiopriapulomorpha ...... 64 Family Meiopriapulidae ...... 64 Genus Meiopriapulus...... 64 Order Halicryptomorpha ...... 64 Family Halicryptidae ...... 64

4 Genus Halicryptus ...... 64 Order Seticoronaria ...... 65 Family Chaetostephanidae ...... 65 Genus Maccabeus ...... 65 3.6.2. Zoogeography ...... 66 4. ËÎÐÈÖÈÔÅÐÛ ...... 67 4.1. Èñòîðèÿ ñèñòåìàòèêè è ñîâðåìåííîå ñîñòîÿíèå ñèñòåìû ...... 67 4.2. Ïëàí ñòðîåíèÿ è îñîáåííîñòè ìîðôîëîãèè, èñïîëüçóåìûå â ñèñòåìàòèêå ...... 67 4.2.1. Âíåøíÿÿ ìîðôîëîãèÿ ...... 67 4.2.2. Ìèêðîñêîïè÷åñêàÿ àíàòîìèÿ ...... 75 4.2.3. Ðàçâèòèå ëîðèöèôåð ...... 75 4.3. Ñèñòåìà, îïðåäåëèòåëüíûå òàáëèöû è äèàãíîçû òàêñîíîâ ...... 76 Êëàññ Loricifera ...... 76 Îòðÿä Nanaloricida ...... 76 Êëþ÷ äëÿ îïðåäåëåíèÿ òàêñîíîâ ïî âçðîñëûì ôîðìàì ...... 76 Êëþ÷ äëÿ îïðåäåëåíèÿ òàêñîíîâ ïî ëè÷èíêàì ...... 77 Ñåìåéñòâî Nanaloricidae ...... 77 Ðîä Nanaloricus ...... 77 Ñåìåéñòâî Pliciloricidae...... 78 Ðîä Pliciloricus ...... 78 Ðîä Rugiloricus ...... 80 4.4. Çîîãåîãðàôèÿ ...... 81 4.5. Summary of chapter 4 ...... 81 5. ÊÈÍÎÐÈÍÕÈ ...... 82 5.1. Èñòîðèÿ ñèñòåìàòèêè è ñîâðåìåííîå ñîñòîÿíèå ñèñòåìû ...... 82 5.2. Ïëàí ñòðîåíèÿ è îñîáåííîñòè ìîðôîëîãèè, èñïîëüçóåìûå â ñèñòåìàòèêå ...... 85 5.2.1. Âíåøíÿÿ ìîðôîëîãèÿ ...... 85 5.2.2. Ìèêðîñêîïè÷åñêàÿ àíàòîìèÿ ...... 87 5.2.2.1. Ïîêðîâû òåëà ...... 87 5.2.2.2. Ìóñêóëàòóðà ...... 88 5.2.2.3. Íåðâíàÿ ñèñòåìà ...... 90 5.2.2.4. Îðãàíû ÷óâñòâ ...... 91 5.2.2.5. Ïèùåâàðèòåëüíàÿ ñèñòåìà ...... 92 5.2.2.6. Ïîëîñòü òåëà...... 93 5.2.2.7. Âûäåëèòåëüíàÿ ñèñòåìà ...... 93 5.2.2.8. Ïîëîâàÿ ñèñòåìà ...... 93 5.2.3. Ðàçâèòèå è ñòàäèè æèçíåííîãî öèêëà ...... 94 5.3. Ôèëîãåíèÿ êèíîðèíõ ...... 96 5.4. Ñèñòåìà, îïðåäåëèòåëüíûå òàáëèöû è äèàãíîçû òàêñîíîâ ...... 104 Êëàññ Kinorhyncha ...... 104 Êëþ÷ äëÿ îïðåäåëåíèÿ íàäâèäîâûõ òàêñîíîâ ...... 104 Îòðÿä Cyclorhagida ...... 105 Ïîäîòðÿä Protorhagae ...... 105 Ñåìåéñòâî Zelinkaderidae ...... 106 Ðîä Zelinkaderes ...... 106 Ñåìåéñòâî Antigomonidae ...... 107 Ðîä Antigomonas ...... 108 Ïîäîòðÿä Cryptorhagae ...... 108 Ñåìåéñòâî Cateriidae ...... 108 Ðîä Cateria ...... 108 Ïîäîòðÿä Conchorhagae ...... 109 Ñåìåéñòâî Semnoderidae ...... 109

5 Ðîä Semnoderes ...... 109 Ðîä Sphenoderes ...... 111 Ïîäîòðÿä Cyclorhagae ...... 112 Ñåìåéñòâî Centroderidae ...... 112 Ðîä Centroderes ...... 112 Ðîä Condyloderes ...... 114 Ðîä Campyloderes ...... 115 Ñåìåéñòâî Echinoderidae ...... 116 Ðîä Echinoderes ...... 116 Ñåìåéñòâî Dracoderidae ...... 157 Ðîä Dracoderes ...... 158 Ñåìåéñòâî Cephalorhynchidae ...... 161 Ðîä Cephalorhyncha ...... 161 Îòðÿä Homalorhagida ...... 164 Ïîäîòðÿä Homalorhagae ...... 164 Ñåìåéñòâî Pycnophyidae ...... 164 Ðîä Pycnophyes...... 164 Ðîä Kinorhynchus ...... 211 Ñåìåéñòâî Neocentrophyidae ...... 228 Ðîä Neocentrophyes ...... 228 Ðîä Paracentrophyes ...... 229 5.5. Çîîãåîãðàôèÿ ...... 230 5.6. Summary of chapter 5 ...... 232 5.6.1. History of classification of the class Kinorhyncha ...... 232 5.6.2. Body plan and morphological characters used in taxonomy ...... 234 5.6.2.1. External morphology ...... 234 5.6.2.2. Microscopical anatomy ...... 235 5.6.2.2.1. Integument ...... 235 5.6.2.2.2. Musculature ...... 235 5.6.2.2.3. Nervous system, sensory organs and glands ...... 236 5.6.2.2.4. Digestive system...... 237 5.6.2.2.5. Body cavity and excretory system ...... 237 5.6.2.2.6. Reproductive system ...... 238 5.6.2.3. Development ...... 238 5.6.3. Phylogenetic relationships within Kinorhyncha ...... 239 5.6.4. Classification, keys for adults and diagnoses of taxa ...... 248 Classification of the class Kinorhyncha...... 248 Class Kinorhyncha ...... 248 Key to supraspecies taxa...... 248 Order Cyclorhagida...... 249 Suborder Protorhagae ...... 249 Family Zelinkaderidae ...... 249 Genus Zelinkaderes ...... 249 Family Antigomonidae ...... 250 Genus Antigomonas ...... 250 Suborder Cryptorhagae ...... 250 Family Cateriidae ...... 251 Genus Cateria ...... 251 Suborder Conchorhagae ...... 251 Genus Semnoderes ...... 251 Genus Sphenoderes ...... 252 Suborder Cyclorhagae ...... 252

6 Family Centroderidae ...... 253 Genus Centroderes ...... 253 Genus Condyloderes ...... 253 Genus Campyloderes ...... 254 Family Echinoderidae ...... 255 Genus Echinoderes ...... 255 Family Dracoderidae ...... 274 Genus Dracoderes ...... 274 Family Cephalorhynchidae ...... 275 Genus Cephalorhyncha ...... 276 Order Homalorhagida ...... 277 Family Pycnophyidae ...... 277 Genus Pycnophyes ...... 277 Genus Kinorhynchus ...... 296 Family Neocentrophyidae ...... 304 Genus Neocentrophyes ...... 304 Genus Paracentrophyes ...... 305 5.6.5. Zoogeography ...... 305 6. ÌÎÐÑÊÈÅ ÍÅÌÀÒÎÌÎÐÔÛ ...... 306 6.1. Èñòîðèÿ ñèñòåìàòèêè è ñîâðåìåííîå ñîñòîÿíèå ñèñòåìû ...... 306 6.2. Ïëàí ñòðîåíèÿ è îñîáåííîñòè ìîðôîëîãèè, èñïîëüçóåìûå â ñèñòåìàòèêå ...... 307 6.3. Ôèëîãåíèÿ âîëîñàòèêîâ ...... 310 6.4. Ñèñòåìà, îïðåäåëèòåëüíûå òàáëèöû è äèàãíîçû òàêñîíîâ ...... 311 Êëàññ Nematomorpha ...... 311 Ïîäêëàññ Nectonematoidea ...... 311 Îòðÿä Nectonematida ...... 311 Ñåìåéñòâî Nectonematidae ...... 311 Ðîä Nectonema ...... 311 6.5. Çîîãåîãðàôèÿ ...... 313 6.6. Summary of chapter 6 ...... 314 Class Nematomorpha ...... 314 Subclass Nectonematoidea ...... 314 Order Nectonematida ...... 314 Family Nectonematidae ...... 314 Genus Nectonema ...... 314 7. ËÈÒÅÐÀÒÓÐÀ ...... 316

7 CONTENTS

1. PREFACE ...... 12 2. INTRODUCTION. Body plan of cephalorynch worms ...... 13 3. PRIAPULIDA ...... 14 3.1. Historical backgroung of classification ...... 14 3.2. Body plan and morphological features used in taxonomy ...... 17 3.2.1. Body plan, external morphology and symmetry ...... 17 3.2.2. Microscopical anatomy ...... 32 3.2.3. Larvae of priapulids ...... 32 3.3. Phylogeny of priapulids ...... 40 3.4. Classification, keys and diagnoses of taxa ...... 40 Class Priapulida ...... 40 Key to supraspecies taxa (adults) ...... 41 Key to supraspecies taxa (larvae) ...... 41 Order Priapulomorpha ...... 42 Family Priapulidae ...... 42 Genus Priapulus ...... 42 Genus Acanthopriapulus ...... 47 Genus Priapulopsis ...... 48 Family Tubiluchidae ...... 49 Genus Tubiluchus ...... 49 Order Meiopriapulomorpha ...... 52 Family Meiopriapulidae ...... 52 Genus Meiopriapulus...... 52 Order Halicryptomorpha ...... 52 Family Halicryptidae ...... 52 Genus Halicryptus ...... 52 Order Seticoronaria ...... 53 Family Chaetostephanidae ...... 53 Genus Maccabeus ...... 53 3.5. Zoogeography ...... 54 3.5.1. Macrobenthic species ...... 54 3.5.2. Meiobenthic species ...... 57 3.5.3. Hypothesis of glacial relicts ...... 58 3.6. English summary of chapter 3 ...... 59 Classification of the class Priapulida ...... 59 Class Priapulida ...... 59 Key to supraspecies taxa (adults) ...... 59 Key to supraspecies taxa (larvae) ...... 59 Order Priapulomorpha ...... 60 Family Priapulidae ...... 60 Genus Priapulus ...... 60 Genus Acanthopriapulus ...... 61 Genus Priapulopsis ...... 62 Family Tubiluchidae ...... 62 Genus Tubiluchus ...... 62 Order Meiopriapulomorpha ...... 64 Family Meiopriapulidae ...... 64 Genus Meiopriapulus...... 64 Order Halicryptomorpha ...... 64 Family Halicryptidae ...... 64

8 Genus Halicryptus ...... 64 Order Seticoronaria ...... 65 Family Chaetostephanidae ...... 65 Genus Maccabeus ...... 65 3.6.2. Zoogeography ...... 66 4. LORICIFERA ...... 67 4.1. Historical background and modern classification ...... 67 4.2. Body plan and morphological features used in taxonomy ...... 67 4.2.1. External morphology ...... 67 4.2.2. Microscopical anatomy ...... 75 4.2.3. Development ...... 75 4.3. Classification, keys and diagnoses of taxa...... 76 Classification of the class Loricifera ...... 76 Class Loricifera...... 76 Order Nanaloricida ...... 76 Key to supraspecies taxa (adults) ...... 76 Key to supraspecies taxa (larvae) ...... 77 Family Nanaloricidae ...... 77 Genus Nanaloricus ...... 77 Family Pliciloricidae ...... 78 Genus Pliciloricus ...... 78 Genus Rugiloricus ...... 80 4.4. Zoogeography ...... 81 4.5. English summary of chapter 4 ...... 81 5. KINORHYNCHA ...... 82 5.1. Historical background and modern classification ...... 82 5.2. Body plan and morphological features used in taxonomy ...... 85 5.2.1. External morphology ...... 85 5.2.2. Microscopical anatomy ...... 87 5.2.2.1. Integument ...... 87 5.2.2.2. Musculature ...... 88 5.2.2.3. Nervous system ...... 90 5.2.2.4. Sensory organs ...... 91 5.2.2.5. Digestive system ...... 92 5.2.2.6. Body cavity ...... 93 5.2.2.7. Excretory system...... 93 5.2.2.8. Reproductive system ...... 93 5.2.3. Development and stages of life history...... 94 5.3. Phylogeny ...... 96 5.4. Classification, keys and diagnoses of taxa...... 104 Class Kinorhyncha ...... 104 Key to supraspecies taxa...... 104 Order Cyclorhagida...... 105 Suborder Protorhagae ...... 105 Family Zelinkaderidae ...... 106 Genus Zelinkaderes ...... 106 Family Antigomonidae ...... 107 Genus Antigomonas ...... 108 Suborder Cryptorhagae ...... 108 Family Cateriidae ...... 108 Genus Cateria ...... 108 Suborder Conchorhagae ...... 109

9 Family Semnoderidae ...... 109 Genus Semnoderes ...... 109 Genus Sphenoderes ...... 111 Suborder Cyclorhagae ...... 112 Family Centroderidae ...... 112 Genus Centroderes ...... 112 Genus Condyloderes ...... 114 Genus Campyloderes ...... 115 Family Echinoderidae ...... 116 Genus Echinoderes ...... 116 Family Dracoderidae ...... 157 Genus Dracoderes ...... 158 Family Cephalorhynchidae ...... 161 Genus Cephalorhyncha ...... 161 Order Homalorhagida ...... 164 Suborder Homalorhagae ...... 164 Family Pycnophyidae ...... 164 Genus Pycnophyes ...... 164 Genus Kinorhynchus ...... 211 Family Neocentrophyidae ...... 228 Genus Neocentrophyes ...... 228 Genus Paracentrophyes ...... 229 5.5. Zoogeography ...... 230 5.6. English summary of chapter 5 ...... 232 5.6.1. History of classification of the class Kinorhyncha ...... 232 5.6.2. Body plan and morphological characters used in taxonomy ...... 234 5.6.2.1. External morphology ...... 234 5.6.2.2. Microscopical anatomy ...... 235 5.6.2.2.1. Integument ...... 235 5.6.2.2.2. Musculature ...... 235 5.6.2.2.3. Nervous system, sensory organs and glands ...... 236 5.6.2.2.4. Digestive system...... 237 5.6.2.2.5. Body cavity and excretory system ...... 237 5.6.2.2.6. Reproductive system ...... 238 5.6.2.3. Development ...... 238 5.6.3. Phylogenetic relationships within Kinorhyncha ...... 239 5.6.4. Classification, keys to adults and diagnoses of taxa ...... 248 Classification of the class Kinorhyncha...... 248 Class Kinorhyncha ...... 248 Key to supraspecies taxa...... 248 Order Cyclorhagida...... 249 Suborder Protorhagae ...... 249 Family Zelinkaderidae ...... 249 Genus Zelinkaderes ...... 249 Family Antigomonidae ...... 250 Genus Antigomonas ...... 250 Suborder Cryptorhagae ...... 250 Family Cateriidae ...... 251 Genus Cateria ...... 251 Suborder Conchorhagae ...... 251 Genus Semnoderes ...... 251 Genus Sphenoderes ...... 252

10 Suborder Cyclorhagae ...... 252 Family Centroderidae ...... 253 Genus Centroderes ...... 253 Genus Condyloderes ...... 253 Genus Campyloderes ...... 254 Family Echinoderidae ...... 255 Genus Echinoderes ...... 255 Family Dracoderidae ...... 274 Genus Dracoderes ...... 274 Family Cephalorhynchidae ...... 275 Genus Cephalorhyncha ...... 276 Order Homalorhagida ...... 277 Family Pycnophyidae ...... 277 Genus Pycnophyes ...... 277 Genus Kinorhynchus ...... 296 Family Neocentrophyidae ...... 304 Genus Neocentrophyes ...... 304 Genus Paracentrophyes ...... 305 5.6.5. Zoogeography ...... 305 6. MARINE NEMATOMORPHA ...... 306 6.1. Historical background and modern classification ...... 306 6.2. Body plan and morphological features used in taxonomy ...... 307 6.3. Phylogeny ...... 310 6.4. Classification, keys and diagnoses of taxa ...... 311 Class Nematomorpha ...... 311 Subclass Nectonematoidea ...... 311 Order Nectonematida ...... 311 Family Nectonematidae ...... 311 Genus Nectonema ...... 311 6.5. Zoogeography ...... 313 6.6. English summary of chapter 6 ...... 314 Class Nematomorpha ...... 314 Subclass Nectonematoidea ...... 314 Order Nectonematida ...... 314 Family Nectonematidae ...... 314 Genus Nectonema ...... 314 7. LITERATURE ...... 316

11 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

1. Ïðåäèñëîâèå

Ýòà êíèãà ïðîäîëæàåò ñåðèþ ìîíîãðàôè÷åñêèõ èçäàíèé, ïîñâÿùåííûõ ìèêðîñêîïè÷åñêîé àíàòîìèè, óëüòðàñòðóêòóðå, ôèëîãåíèè è ñèñòåìàòèêå öåôàëîðèíõ - øèðîêî ðàñïðîñòðàíåííîé ãðóïïû ìîðñêèõ è ïðåñíîâîäíûõ ãîëîâîõîáîòíûõ ÷åðâåé, ïðåäñòàâëåííûõ êàê ñîâðåìåííûìè, òàê è èñêîïàåìûìè îðãàíèçìàìè. Òèï Cephalorhyncha âêëþ÷àåò â ñåáÿ ÷åòûðå êëàññà ãîëîâîõî- áîòíûõ ÷åðâåé: ïðèàïóëèäû (Priapulida), ëîðèöèôåðû (Loricifera), êèíîðèíõè (Kinorhyncha) è âîëîñàòèêè (Nematomorpha) (ñì. Ìàëàõîâ, 1980, 1986; Àäðèàíîâ, Ìàëàõîâ, 1994, 1995, 1996; Ìàëàõîâ, Àäðèàíîâ, 1995; Adrianov, Malakhov, 1994, 1995, 1996a–b). Ïåðâûå òðè ãðóïïû ïðåä- ñòàâëåíû èñêëþ÷èòåëüíî ìîðñêèìè ðîþùèìè ìàêðîáåíòîñíûìè èëè ìåéîáåíòîñíûìè ôîðìà- ìè, òîãäà êàê êëàññ Nematomorpha âêëþ÷àåò òîëüêî ïàðàçèòè÷åñêèõ ìîðñêèõ (ïîäêëàññ Nectonematoidea) è ïðåñíîâîäíûõ (ïîäêëàññ Gordioidea) ïðåäñòàâèòåëåé.  íàñòîÿùåé ñèñòåìà- òè÷åñêîé ñâîäêå ìû îãðàíè÷èâàåìñÿ ðàñìîòðåíèåì òîëüêî ìîðñêèõ âîëîñàòèêîâ èç ïîäêëàññà Nectonematoidea. Ïåðâûå òðè êíèãè ñåðèè â îñíîâíîì ïîñâÿùåíû âîïðîñàì òîíêîé ìîðôîëîãèè è ôèëîãåíèè ðàçëè÷íûõ ãðóïï ãîëîâîõîáîòíûõ ÷åðâåé è òèïà Cephalorhyncha âöåëîì (Àäðèàíîâ, Ìàëàõîâ, 1994, 1996; Ìàëàõîâ, Àäðèàíîâ, 1995).  ýòîì èçäàíèè ìû ïðåäëàãàåì ÷èòàòåëþ ïîëíûé îïðåäå- ëèòåëü ìîðñêîé ôàóíû öåôàëîðèíõ, âïåðâûå ïîäãîòîâëåííûé äëÿ ýòîé ãðóïïû ìîðñêèõ ÷åðâåé. Âñå ðàçäåëû êíèãè ïðåäñòàâëÿþò ñîáîé ïëîä ñîâìåñòíîé ðàáîòû àâòîðîâ çà èñêëþ÷åíèåì ãëàâû 5 "Êèíîðèíõè", àâòîðñòâî êîòîðîé ïðèíàäëåæèò òîëüêî À.Â. Àäðèàíîâó. Ìû áëàãîäàðíû èçäàòåëüñòâó “Smithsonian Institution Press”, à òàêæå ðåäàêöèÿì íàó÷íûõ æóð- íàëîâ “Biological Society of Washington” è “American Microscopical Society”, ïðåäîñòàâèâøèì íàì ïèñüìåííûå ðàçðåøåíèÿ íà èñïîëüçîâàíèå èëëþñòðàöèé èç ýòèõ èçäàíèé â íàøèõ îïðåäåëèòåëü- íûõ òàáëèöàõ. Âûïîëíåíèå èññëåäîâàíèé, ïîëîæåííûõ â îñíîâó íàñòîÿùåé ðàáîòû ïîääåðæèâàëîñü Ðîñ- ñèéñêèì ôîíäîì ôóíäàìåíòàëüíûõ èññëåäîâàíèé (ÐÔÔÈ) (ïðîåêò N 96-15-97037). ×àñòè÷íî ðàáîòà ïîääåðæèâàëàñü Ðîññèéñêèì ôîíäîì "Áèîðàçíîîáðàçèå". Èçäàíèå îñóùåñòâëåíî â ðàìêàõ èçäàòåëüñêîãî ïðîåêòà ÐÔÔÈ (ïðîåêò N 98-04-62148). Íåäîñòàþùèå ñðåäñòâà âíåñåíû èçäàòåëü- ñòâîì ÊÌÊ è àâòîðàìè.

Áëàãîäàðíîñòè

Àâòîðû ïðèíîñÿò áëàãîäàðíîñòè êîëëåãàì — ïðîô. Ð. Õèããèíñó (ÑØÀ, Âàøèíãòîí), ïðîô. Ô. Øòîðõó (ÔÐÃ, Ãåéäåëüáåðã), ä-ðó Ñ.Ý. Ñïèðèäîíîâó (Ðîññèÿ, Ìîñêâà), Ñ.À. Ìàñëàêîâîé (Ðîñ- ñèÿ, Ìîñêâà), îêàçàâøèì áîëüøóþ ïîìîùü íà ðàçëè÷íûõ ýòàïàõ ðàáîòû íàä êíèãîé. Àâòîðû áëàãîäàðíû êîëëåêòèâó Ìåæêàôåäðàëüíîé ëàáîðàòîðèè ýëåêòðîííîé ìèêðîñêîïèè Áèîëîãè÷åñêîãî ôàêóëüòåòà ÌÃÓ (Ìîñêâà) âî ãëàâå ñ Ã.Í. Äàâèäîâè÷åì è À.Ã. Áîãäàíîâûì çà ïîìîùü â ïðîâåäåíèè óëüòðàñòðóêòóðíûõ èññëåäîâàíèé. Àâòîðû áëàãîäàðÿò Ðîññèéñêèé ôîíä ôóíäàìåíòàëüíûõ èññëåäîâàíèé çà ôèíàíñîâóþ ïîä- äåðæêó íàøåé ðàáîòû. PHYLOGENY, AND CLASSIFICATION PRIAPULIDA: STRUCTURE, DEVELOPMENT,

KMK LTD. ÌÎÑÊÂÀ 12 1996 2. Ââåäåíèå

Òèï Cephalorhyncha áûë ïðåäëîæåí â 1980 ã. îäíèì èç àâòîðîâ (Ìàëàõîâ, 1980) äëÿ îáúåäèíå- íèÿ òðåõ êëàññîâ ïåðâè÷íîïîëîñòíûõ ÷åðâåé — ïðèàïóëèä (Priapulida), êèíîðèíõ (Kinorhyncha) è âîëîñàòèêîâ (Nematomorpha). Ïîçäíåå, ïîñëå îòêðûòèÿ ëîðèöèôåð (Loricifera), îíè òàêæå áûëè âêëþ÷åíû â ñîñòàâ ãîëîâîõîáîòíûõ ÷åðâåé íà ïðàâàõ êëàññà (Ìàëàõîâ, 1986; Àäðèàíîâ è äð., 1989). Èñòîðèÿ ïðîáëåìû îáúåäèíåíèÿ ýòèõ ãðóïï è êîíöåïöèÿ ñðàâíèòåëüíî-ìîðôîëîãè÷åñêîãî åäèíñòâà âñåõ ãîëîâîõîáîòíûõ ÷åðâåé ïîäðîáíî ðàññìîòðåíû íàìè â ïðåäøåñòâóþùèõ èçäàíèÿõ (ñì. Ìàëàõîâ, 1980, 1986; Àäðèàíîâ è äð., 1989; Àäðèàíîâ, Ìàëàõîâ, 1994; 1995à–å; Ìàëàõîâ, Àäðèàíîâ, 1995; Adrianov, Malakhov, 1994, 1995, 1996a–b). Íàçâàíèå Cephalorhyncha ïðîèñõîäèò îò ãðå÷åñêèõ ñëîâ “kephalon” — ãîëîâà è “rhynchos” — ðûëî, õîáîò. Âñåì öåôàëîðèíõàì, èëè ãîëîâîõîáîòíûì ÷åðâÿì, ñâîéñòâåíåí åäèíûé ïëàí ñòðîåíèÿ, îò- ëè÷àþùèé èõ îò äðóãèõ ãðóïï ïåðâè÷íîïîëîñòíûõ ÷åðâåé. Òåëî âñåõ ãîëîâîõîáîòíûõ ÷åðâåé ïîä- ðàçäåëÿåòñÿ íà äâà îòäåëà: õîáîòíûé (èíòðîâåðò) è òóëîâèùíûé. Ó âñåõ ñîâðåìåííûõ ãîëîâîõî- áîòíûõ ÷åðâåé èíòðîâåðò (õîòÿ áû íà ëè÷èíî÷íîé ñòàäèè) ñïîñîáåí ââîðà÷èâàòüñÿ âíóòðü òóëî- âèùà. Õîáîòíûé îòäåë, â ñâîþ î÷åðåäü, ïîäðàçäåëÿåòñÿ íà ïîäîòäåëû: ðîòîâîé êîíóñ, íåñóùèé íàïðàâëåííûå âïåðåä ïðèäàòêè (ñòèëåòû èëè çóáû), ñðåäíþþ ÷àñòü, âîîðóæåííóþ íàïðàâëåí- íûìè íàçàä ñåíñîðíî-ëîêîìîòîðíûìè ïðèäàòêàìè — ñêàëèäàìè, è øåéíóþ îáëàñòü (íàçûâàå- ìóþ èíîãäà òîðàêñîì), îáû÷íî ëèøåííóþ ïðèäàòêîâ. Ðàñïîëîæåíèå ïðèäàòêîâ ðîòîâîãî êîíóñà è ñðåäíåé ÷àñòè õîáîòà ïîä÷èíÿåòñÿ ðàäèàëüíîé ñèììåòðèè. Íà âåðøèíå ðîòîâîãî êîíóñà îòêðû- âàåòñÿ òåðìèíàëüíîå ðîòîâîå îòâåðñòèå, âåäóùåå â ñêâîçíîé êèøå÷íèê, çàêàí÷èâàþùèéñÿ òåð- ìèíàëüíûì èëè ñóáòåðìèíàëüíûì (âåíòðàëüíûì) àíóñîì. Åäèíñòâî ãîëîâîõîáîòíûõ ÷åðâåé ïîä÷åðêèâàåòñÿ îáùèì ïëàíîì ñòðîåíèÿ íåðâíîé ñèñòåìû, êîòîðûé ñî âðåìåí Êþâüå ñ÷èòàåòñÿ âàæíåéøåé õàðàêòåðèñòèêîé òèïîâîãî ðàíãà. Íåðâíàÿ ñèñ- òåìà âñåõ ãîëîâîõîáîòíûõ ÷åðâåé ïðåäñòàâëåíà îêîëîãëîòî÷íûì íåðâíûì êîëüöîì, çàëåãàþùèì îáÿçàòåëüíî â õîáîòíîì îòäåëå â îñíîâàíèè ðîòîâîãî êîíóñà, è âåíòðàëüíûì íåðâíûì ñòâîëîì, èäóùèì â ýêòîäåðìå áðþøíîé ñòîðîíû. Õîáîòíûé îòäåë (èíòðîâåðò) Ñephalorhyncha — ñïåöèôè÷åñêîå îáðàçîâàíèå, ñâîéñòâåííîå òîëüêî ïðåäñòàâèòåëÿì ýòîãî òèïà. Îí ðåçêî îòëè÷àåòñÿ îò âûâîðà÷èâàþùèõñÿ è âûäâèãàþùèõñÿ îáðàçîâàíèé â äðóãèõ ãðóïïàõ æèâîòíîãî öàðñòâà (õîáîòêîâ òóðáåëëÿðèé-êàëèïòîðèíõèé, ëîâ÷å- ãî õîáîòà íåìåðòèí è ïðèêðåïèòåëüíîãî õîáîòà ñêðåáíåé, âûâîðà÷èâàþùèõñÿ ãëîòîê ïîëèõåò) òåì, ÷òî íåñåò â ñåáå öåíòðàëüíûé îðãàí íåðâíîé ñèñòåìû — êîëüöåâîé ìîçã è êîìïëåêñ îñíîâ- íûõ îðãàíîâ ÷óâñòâ, à òàêæå ðîòîâîå îòâåðñòèå è ïåðåäíèå îòäåëû ïèùåâàðèòåëüíîãî òðàêòà. Õîáîòíûé îòäåë Cephalorhyncha — ýòî âûâîðà÷èâàþùàÿñÿ ãîëîâà. Ïðèìåíèòåëüíî ê Cephalorhyncha ìû èñïîëüçóåì ñëîâîñî÷åòàíèÿ “õîáîò”,”õîáîòíûé îòäåë” êàê ñèíîíèì òåðìèíà èíòðîâåðò. Ãîëîâîõîáîòíûå ÷åðâè — ýòî îáøèðíàÿ è ðàçíîîáðàçíàÿ ãðóïïà æèâîòíûõ, íî âñå ïðåäñòà- âèòåëè ýòîãî òèïà ñîîòâåòñòâóþò ñôîðìóëèðîâàííûì âûøå ïîëîæåíèÿì åäèíîãî ïëàíà ñòðîåíèÿ. PHYLOGENY, AND CLASSIFICATION PRIAPULIDA: STRUCTURE, DEVELOPMENT,

KMK LTD. ÌÎÑÊÂÀ 1996 13 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

3. Ïðèàïóëèäû

Ïðèàïóëèäû (Priapulida) — ìîðñêèå áåíòîñíûå ÷åðâè, â íàñòîÿùåå âðåìÿ ðàññìàòðèâàåìûå â ðàíãå êëàññà â ïðåäåëàõ òèïà ãîëîâîõîáîòíûõ ÷åðâåé — Cephalorhyncha (Ìàëàõîâ, 1980; Àäðèà- íîâ, Ìàëàõîâ, 1995à–å, 1996; Adrianov, Malakhov, 1995; Ìàëàõîâ, Àäðèàíîâ, 1995). Ãðóïïà ïðåä- ñòàâëåíà êàê âûìåðøèìè êåìáðèéñêèìè è êàðáîíîâûìè ôîðìàìè, òàê è ñîâðåìåííûìè ìàêðî- áåíòîñíûìè è ìåéîáåíòîñíûìè ïðèàïóëèäàìè, øèðîêî ðàñïðîñòðàíåííûìè â Ìèðîâîì îêåàíå (Conway Morris, 1977; Land, 1970). Ïåðâûå ïðèàïóëèäû ïîÿâèëèñü óæå â êåìáðèè, ãäå áûëè îñíîâíûìè ðîþùèìè äåòðèòîôàãàìè è õèùíèêàìè, ïîëíîñòüþ äîìèíèðóþùèìè â øåëüôîâûõ ãðóíòàõ (Conway Morris, 1977).  êëàññå Priapulida ìû ðàññìàòðèâàåì ëèøü íàñòîÿùèõ êåìáðèéñêèõ ïðèàïóëèä — ïðåäñòàâèòåëåé Selkirkia è Ottoia, òîãäà êàê äðóãèå ïðåäñòàâèòåëè ñáîðíîãî èñêîïàåìîãî òàêñîíà Archaepriapulida (ñì. Conway Morris, 1977) — Fieldia, Ancalagon è Louisella, íåäàâíî áûëè âûäåëåíû â ñàìîñòîÿòåëüíûå òàêñîíû (êëàññû) â ïðåäåëàõ òèïà ãîëîâîõîáîòíûõ (Àäðèàíîâ, Ìàëàõîâ, 1995à–å; Adrianov, Malakhov, 1995). Ãîñïîäñòâî â áåíòîñå ïðèàïóëèäû ïîñòåïåííî óñòóïèëè àííåëèäàì, ïðåäïîëîæèòåëüíî, â Îðäî- âèêå, êîãäà ó ïîëèõåò âïåðâûå ïîÿâèëèñü ÷åëþñòè (Conway Morris, 1979; Gould, 1989). Îáùàÿ ìîðôîëîãèÿ ïðèàïóëèä ñ êåìáðèÿ èçìåíèëàñü î÷åíü ìàëî è îíè ïî ïðàâó ðàññìàòðè- âàþòñÿ êàê íàãëÿäíûé ïðèìåð òàê íàçûâàåìîé “ìåäëåííîé ýâîëþöèè” è “æèâûõ èñêîïàåìûõ”. Óæå â êàðáîíå îáíàðóæåíû õâîñòàòûå ôîðìû — Priapulites, óäèâèòåëüíî ñõîäíûå ñ ñîâðåìåííû- ìè Priapulus (Schram, 1973) è êîòîðûõ ìîæíî âïîëíå ñîïîñòàâèòü ñ òàêèìè “òèïè÷íûìè æèâû- ìè èñêîïàåìûìè”, êàê áðàõèîïîäà Lingula, õåëèöåðîâîå Limulus èëè ðàêîîáðàçíîå Triops, èìåþ- ùèìè èñêîïàåìûõ äâîéíèêîâ â ïàëåîçîéñêîé ôàóíå. Ñîâðåìåííûå ïðèàïóëèäû èìåþò ìåéîáåíòîñíûõ ëè÷èíîê. Ëè÷èíêè ïðèàïóëèä âïîëíå ñîîò- âåòñòâóþò ïëàíó ñòðîåíèÿ âçðîñëûõ ÷åðâåé, îäíàêî èõ òóëîâèùå çàêëþ÷åíî â æåñòêèé êóòèêó- ëÿðíûé ïàíöèðü — ëîðèêó, ïðåäîõðàíÿþùèé ýòè ìèêðîñêîïè÷åñêèå ôîðìû îò àáðàçèè ïðè èíòåðñòèöèàëüíîì îáðàçå æèçíè. Åäèíñòâåííàÿ ñîâðåìåííàÿ ïðèàïóëèäà ñ ïðÿìûì ðàçâèòèåì — Meiopriapulus. Âèäîâîå ðàçíîîáðàçèå ïðèàïóëèä íåâåëèêî. Îïèñàíû îíè áûëè âïåðâûå â ñåðåäèíå XVIII âåêà (Odhelius, 1754; Linnaeus, 1759), íî ñ òîé ïîðû íàóêå èçâåñòíî ÷óòü áîëåå 20 âèäîâ ýòèõ æèâîòíûõ (Land, 1970; Adrianov, Malakhov, 1996a–b). Èíòåðåñíî, ÷òî áîëåå ïîëîâèíû âèäîâ ïðèàïóëèä îòêðûòû ëèøü âî âòîðîé ïîëîâèíå íàøåãî âåêà, êîãäà îáîçíà÷èëñÿ èíòåðåñ ê ýòîìó òàêñîíó â ñâÿçè ñ íàõîäêàìè êàðëèêîâûõ ïðèàïóëèä Tubiluchus (Land, 1968), ñåäåíòàðíûõ Maccabeus (Por, 1973) è ìåéîáåíòîñíûõ Meiopriapulus (Morse, 1981): ê íà÷àëó âåêà áûëî èçâåñòíî 5 âèäîâ ñîâðåìåííûõ ïðèàïóëèä, ê 1968 ãîäó èõ áûëî óæå 8, à ê 1991 — 16 âàëèäíûõ âèäîâ. Ê íàñòîÿùåìó âðåìåíè îïèñàíû óæå 17 âèäîâ ñîâðåìåííûõ íàñòîÿùèõ ïðèàïóëèä. Íåñìîòðÿ íà íåáîëüøîå ÷èñëî âèäîâ, ïðèàïóëèäû è èõ ëè÷èíêè äåìîíñòðèðóþò ðàçíîîáðàç- íûå ìîðôîëîãè÷åñêèå îñîáåííîñòè, î÷åíü âàæíûå äëÿ ïîíèìàíèÿ ìîðôîëîãè÷åñêîãî åäèíñòâà è ôèëîãåíèè âñåõ ãîëîâîõîáîòíûõ ÷åðâåé. Ýâðèáèîíòíîñòü ïðèàïóëèä è îñîáåííîñòè èõ áèîëîãèè è ðàñïðîñòðàíåíèÿ â Ìèðîâîì îêåàíå äåëàþò ýòó ðåëèêòîâóþ ãðóïïó èíòåðåñíîé è ñ òî÷êè çðåíèÿ ýêîëîãèè è çîîãåîãðàôèè ìîðñêèõ áåíòîñíûõ îðãàíèçìîâ (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996). PHYLOGENY, AND CLASSIFICATION 3.1. Èñòîðè÷åñêèé îáçîð ñèñòåìû ïðèàïóëèä

Ïåðâîå íàó÷íîå îïèñàíèå ïðèàïóëèä ïîä èìåíåì Priapus humanus ìû íàõîäèì ó Îäõåëèóñà

(Odhelius, 1754) â åãî äèññåðòàöèè “Chinensis Lagerstromiana”,PRIAPULIDA: STRUCTURE, DEVELOPMENT, ïîñâÿùåííîé ôàóíå Êèòàÿ. Îäíà- êî, ïî ñïðàâåäëèâîìó çàìå÷àíèþ Ëàíäà (Land, 1970), ïðîèñõîæäåíèå ýòîãî ìàòåðèàëà íàõîäèòñÿ ïîä âîïðîñîì: íåò óêàçàíèé â ñîïðîâîäèòåëüíûõ çàïèñÿõ Îäõåëèóñà, ãîëîòèï óòåðÿí, äðóãèå íàõîäêè ïðèàïóëèä ó áåðåãîâ Êèòàÿ äî ñèõ ïîð íå èçâåñòíû. ×óòü ïîçæå ýòîò âèä áûë ïåðåîïèñàí Ê. Ëèííååì (Linnaeus, 1759) è ïîä òåì æå íàçâàíèåì çàíÿë ìåñòî â çíàìåíèòîé “Systema naturae”.  ðîä Priapus âìåñòå ñ P. humanis Ëèííåé âêëþ÷èë ïðåäñòàâèòåëÿ ìîðñêèõ àíåìîíîâ Priapus equinus. KMK LTD. ÌÎÑÊÂÀ 14 1996 Ãëàâà 3. Ïðèàïóëèäû

Íàçâàíèå ðîäà ïðîèñõîäèò îò ãðå÷åñêîãî ñëîâà Priapos.  àíòè÷íîé ìèôîëîãèè Ïðèàï — èòèôàë- ëè÷åñêîå áîæåñòâî ïðîèçâîäèòåëüíûõ ñèë ïðèðîäû, ïðè÷åì èçíà÷àëüíî ñîáñòâåííî ôàëëîñ (ïîä- ðîáíåå ñì. Àäðèàíîâ, Ìàëàõîâ, 1996). Âûáîð Ïðèàïà êàê ïðîòîòèïà äëÿ íàçâàíèÿ îïèñàííûõ Îäõåëè- óñîì è Ëèííååì æèâîòíûõ áûë îáóñëîâëåí íåêîòîðûì ñõîäñòâîì ýòèõ îðãàíèçìîâ ñ ìóæñêèì ïîëî- âûì ÷ëåíîì. Ïîçæå Ëèííåé ïåðåìåñòèë ðîä Priapus èç ìîðñêèõ àíåìîíîâ ê ãîëîòóðèÿì, äàâ íîâîå íàçâàíèå — Holothuria priapus (Linnaeus, 1767). Ïðèàïóëèä îïèñûâàëè è êàê ïèÿâîê: Hirudo annulis (Olafsen, 1772). Ñîâðåìåííîå íàçâàíèå — Priapulus caudatus — áûëî äàíî Ëàìàðêîì (Lamarck, 1816). Êàê è áîëüøèíñòâî äðóãèõ çîîëîãîâ, Ëàìàðê ïðîäîëæàë ñ÷èòàòü ïðèàïóëèä èãëîêîæèìè. Íàçâàíèå Priapus (P. equinus Linnaeus, 1758) âïîñëåäñòâèè ñòàëî òèïîâûì â ïðåäåëàõ êèøå÷íîïîëîñòíûõ (Poche, 1908).  ñèëó ýòîãî, à òàêæå èç-çà òîãî, ÷òî óìåíüøèòåëüíûé ñóôôèêñ ëó÷øå îòðàæàë íåáîëüøèå ðàçìåðû æèâîòíîãî, íàçâàíèå Priapulus áûëî ïðèíÿòî áîëüøèíñòâîì çîîëîãîâ.  òî æå âðåìÿ, îïðåäåëåííàÿ ïóòàíèöà â íîìåíêëàòóðå Priapulus caudatus ñîõðàíÿëàñü äîâîëü- íî äîëãî: Priapus humanus (Houttuyn, 1770), Priapulus humanus (Guerne, 1888; Thiel, 1906; Àâåðèí- öåâ, 1908), P. priapus (Äåðþãèí, 1906; Ñêîðèêîâ, 1910), P. vulgaris (Cuvier, 1836), Priapula priapus (Blainville, 1826), P. caudatus (Blainville, 1826), P. caudata (Guérin-Ménéville, 1829), Holothuria priapus (Müller, 1775; Fabricius 1780; Herbst, 1788; Cuvier, 1830; Voigt, 1840). Ýòà êàðòèíà óñëîæíÿëàñü ìíîãî÷èñëåííûìè ïîïûòêàìè îïèñàíèé íîâûõ âèäîâ Priapulus, êîòîðûå ïðè ïðîâåðêàõ îêàçûâà- ëèñü P. caudatus (P. glandifer Ehlers, 1861, P. pygmaeus Verrill, 1879, P. multidentatus Mobius, 1873, P. intermedius Lenz, 1878, P. profundus Sanders et Hessler, 1962 è äð.) (ñì. Land, 1970). Âî èçáåæàíèå ïóòàíèöû â 1932 ãîäó áûëî ñîçâàíî ñïåöèàëüíîå ñîâåùàíèå Ìåæäóíàðîäíîé Êîìèññèè ïî çîîëîãè÷åñêîé íîìåíêëàòóðå (ñì. Land, 1971), ïðèíÿâøåå ñëåäóþùèå ðåøåíèÿ: 1. Ïîñêîëüêó òèïîâîé ýêçåìïëÿð, îïèñàííûé êàê Priapus humanus, óòðà÷åí, à Priapus equinus ïðè- çíàí òèïîâûì ðîäîì àêòèíèé (ñì. Poche, 1908), ñ÷èòàòü íàçâàíèå Priapus ðîäîì êèøå÷íîïîëîñòíûõ. 2. Íàçâàíèå Holothuria ñ÷èòàòü ðîäîì èãëîêîæèõ. 3. Íàçâàíèå Priapulus è ñîïðÿæåííîå ñ íèì âèäîâîå íàçâàíèå caudatus ñ÷èòàòü âàëèäíûìè, à ïåðâîå èç íèõ — òèïîâûì äëÿ ñåìåéñòâà Priapulidae è äëÿ òèïà â öåëîì. 4. Âèäîâîå íàçâàíèå humanus âïåðâûå ñîïðÿæåíî â áèíîìåíå Priapus humanus è íå âàëèäíî äëÿ Priapulus. 5. Âèäîâîå íàçâàíèå priapus âïåðâûå ñîïðÿæåíî â áèíîìåíå Holothuria priapus è íå âàëèäíî äëÿ Priapulus. 6. Âíåñòè áèíîìåí Priapulus caudatus â îôèöèàëüíûé ñïèñîê âèäîâûõ çîîëîãè÷åñêèõ íàçâàíèé (Official List of Specific Names in Zoology). Âòîðîé âèä è ðîä ïðèàïóëèä — Halicryptus spinulosus Siebold, 1849 — áûë îïèñàí óæå â ñåðåäèíå ïðîøëîãî ñòîëåòèÿ (Siebold, 1849).  1913 ãîäó áûëè âïåðâûå îïèñàíû ëè÷èíêè ïðèàïóëèä (Hammarsten, 1913), îêàçàâøèåñÿ ïîñòýìáðèîíàëüíûìè ñòàäèÿìè H. spinulosus. Íåäàâíî áûë îïèñàí âòîðîé âèä ðîäà Halicryptus — ãèãàíòñêèé H. higginsi, íàéäåííûé íà Àëÿñêå (Storch, 1996; Shirley, Storch, 1998). Ïîìèìî îïèñàííûõ ïåðâûìè ðîäîâ Priapulus è Halicryptus â ñåìåéñòâå Priapulidae íà ñåãîä- íÿøíèé äåíü îïèñàíû åùå Priapulopsis Koren et Danielsen 1875 è Acanthopriapulus Land, 1970. Êðî- ìå íèõ â íàñòîÿùåå âðåìÿ èçâåñòíû ìåéîáåíòîñíûå ïðèàïóëèäû ðîäîâ Tubiluchus Land, 1968 è Meiopriapulus Morse, 1981. Èçâåñòåí è îäèí ðîä ñåäåíòàðíûõ ïðèàïóëèä — Maccabeus Por, 1973. Âçãëÿäû ðàçíûõ àâòîðîâ íà ñèñòåìàòè÷åñêîå ïîëîæåíèå ïðèàïóëèä ñóùåñòâåííî ðàçëè÷àëèñü. Áëýíâèëü (Blainville, 1826) ðàññìàòðèâàë ïðèàïóëèä â ñîñòàâå “îòäåëüíîé ãðóïïû æèâîòíûõ” — Subactinozoaries, áëèçêîé ê àêòèíèÿì. Îí âûäåëèë ïðèàïóëèä â îòäåëüíûé îòðÿä ñ íàçâàíèåì Proboscephale, îòðàçèâ ýòèì òåðìèíîì îñíîâíóþ ìîðôîëîãè÷åñêóþ îñîáåííîñòü, ïîëîæåííóþ â õàðàêòåðèñòèêó ñîâðåìåííîãî òèïà ãîëîâîõîáîòíûõ ÷åðâåé (Blainville, 1826). Äîëãîå âðåìÿ ïðè- àïóëèä ðàññìàòðèâàëè âìåñòå ñ ñèïóíêóëèäàìè è ýõèóðèäàìè êàê ïåðåõîäíóþ ìåæäó ÷åðâÿìè è èãëîêîæèìè ãðóïïó — Gephyrea (ãðå÷. gephyrea — ìîñò) (Quatrefages, 1847). À íåêîòîðûå àâòîðû äàæå âêëþ÷àëè èõ â ñîñòàâ ñèïóíêóëèä (Diesing, 1859; Haeckel, 1896).  1908 ãîäó Âîñìàåð (Vosmaer, 1908) âïåðâûå ïðåäëîæèë ðàññìàòðèâàòü ïðèàïóëèä â êà÷åñòâå îòäåëüíîãî òèïà.  íûíåøíåì ñòîëåòèè âî âçãëÿäàõ íà ñèñòåìàòè÷åñêîå ïîëîæåíèå ïðèàïóëèä îïðåäåëèëèñü äâå îñíîâíûå òåíäåíöèè: îòíîñèòü èõ ê öåëîìè÷åñêèì ÷åðâÿì èëè ðàññìàòðèâàòü â ïðåäåëàõ ïåðâè÷íîïîëîñòíûõ.

15 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

ÒÀÁËÈÖÀ 3.1. ÑÈÑÒÅÌÀ ÏÐÈÀÏÓËÈÄ ÏÎ ËÀÍÄÓ (LAND, 1970). ÒÀÁËÈÖÀ 3.2. ÑÈÑÒÅÌÀ ÀÐÕÅÏÐÈÀÏÓËÈÄ ÏÎ ÊÎÍÂÅÉ TABLE 3.1. CLASSIFICATION OF PRIAPULIDA AFTER LAND (1970). ÌÎÐÐÈÑÓ (CONWAY MORRIS, 1977). TABLE 3.2. CLASSIFICATION OF ARCHAEOPRIAPULIDA AFTER Семейство Род Вид CONWAY MORRIS, 1977. P.caudatus Priapulus Семейство Род Вид P.tuberculatospinosus Ottoiidae Ottoia O.prolifica P.bicaudatus Priapulidae Priapulopsis Selkirkiidae Selkirkia S.columbia P.australis Miskoiidae Louisella L.pedunculata Acanthopriapulus A.horridus Ancalagonidae Ancalagon A.minor Halicryptus H.spinulosus Fieldiidae Fieldia F.lanceolata Tubiluchidae Tubiluchus T.corallicola Priapulidae Priapulites P.konecniorum

Íåêîòîðûå àâòîðû, ïîëàãàÿ íàëè÷èå ó ïðè- Scolecofurca S.rara àïóëèä öåëîìà, ïðåäëàãàëè ðàññìàòðèâàòü èõ êàê Lecythioscopa L.simplex

äîáàâëåíèå ê òèïó êîëü÷àòûõ ÷åðâåé — àííåëèä ÒÀÁËÈÖÀ 3.3. ÑÎÂÐÅÌÅÍÍÀß ÑÈÑÒÅÌÀ ÏÐÈÀÏÓËÈÄ. (Çåíêåâè÷, 1940; Boettger, 1952). Ïðîìåæóòî÷íóþ TABLE 3.3. CLASSIFICATION OF PRIAPULIDA. ïîçèöèþ çàíèìàë Áàëüöåð (Baltzer, 1931), ðàñ- ñìàòðèâàÿ ïðèàïóëèä ñâîåîáðàçíûì ïåðåõîäíûì Отряд Семейство Род Вид çâåíîì ìåæäó íèçøèìè è âûñøèìè ÷åðâÿìè. S.columbia Selkirkio- Íàèáîëåå ìíîãî÷èñëåííû òî÷êè çðåíèÿ, Selkirkiidae Selkirkia S.spencei morpha ñáëèæàþùèå ïðèàïóëèä ñ íèçøèìè ïåðâè÷íî- S.willoughbyi ïîëîñòíûìè ÷åðâÿìè. Ïðèàïóëèä ñðàâíèâàëè ñ Ottoiomorpha Ottoiidae Ottoia O.prolifica êèíîðèíõàìè (Hammarsten, 1915; Remane, 1936; Priapulites P.konecniorum Merriman, 1985), ãàñòðîòðèõàìè (Hammarsten, 1915), êîëîâðàòêàìè (Lang, 1939; Por, Bromley, P.caudatus P.tuberculato- 1974), ñêðåáíÿìè (Schepotieff, 1908; Meyer, Priapulus 1938; Haffner, 1950; Golvan, 1958; Conway spinosus Morris, 1977). Ãàììàðñòåí âûäåëèë òèï Trochel- Priapulidae P.abyssorum minthes, âêëþ÷èâ â íåãî Rotifera, Gastrotricha, Acantho- A.horridus Kinorhyncha è Priapulida, ñ÷èòàÿ, ÷òî âñå ýòè priapulus êëàññû ïðîèçîøëè îò îäíîãî ïðåäêà, ïðè÷åì Priapulo- P.bicaudatus morpha Priapulopsis ïîñëåäíèå äâà íàèáîëåå áëèçêè äðóã ê äðóãó P.australis (Hammarsten, 1913).  1908 ãîäó Ãðîááåí (Grob- T.corallicola ben, 1908) îáúåäèíèë ïåðâè÷íîïîëîñòíûõ ÷åð- âåé â ðàíãàõ îòðÿäîâ â ñáîðíûé êëàññ Aschel- T.remanei minthes â ïðåäåëàõ ñêîëåöèä. T.philippinensis Tubiluchidae Tubiluchus  1944 ãîäó Â.Í. Áåêëåìèøåâ (1944) âêëþ÷èë T.australiensis ïðèàïóëèä â êà÷åñòâå êëàññà â ïîäòèï Nemathel- T.vanuatensis minthes òèïà Scolecida, ñáëèæàÿ èõ, ñî ñêðåáíÿ- T.arcticus ìè è êðóãëûìè ÷åðâÿìè. Îäíàêî óæå â 1952 ãîäó Halicrypto- H.spinulosus (Áåêëåìèøåâ, 1952) îí ñïðàâåäëèâî âûíåñ Halicryptidae Halicryptus ñêðåáíåé â îòäåëüíûé ïîäòèï. Ãàéìåí (Hyman, morpha H.higginsi Meiopriapulo- Meiopriapu- Meiopria- 1951) ïîâûñèëà ðàíã àñêãåëüìèíòîâ äî òèïà, M.fijiensis âêëþ÷èâ â íåãî êëàññû Rotifera, Gastrotricha, morpha lidae pulus Kinorhyncha, Priapulida, Nematoda, Nematomor- Chaetoste- M.tentaculatus Seticoronaria Maccabeus pha, ïðåäîñòàâèâ ñêðåáíÿì îòäåëüíûé òèï. Ïðè- phanidae M.cirratus íÿâ âî âíèìàíèå ýìáðèîíàëüíîå ðàçâèòèå ïðè- àïóëèä è äðóãèõ ÷åðâåé, Ëàíã (Lang, 1953) ñáëèæàë ïðèàïóëèä ñ Kinorhyncha è Acanthocephala â ïîäòèïå Rhynchohelminthes òèïà Aschelminthes. Ïî ìíåíèþ Ìåððèìàí (Merriman, 1985), íà îñ- íîâå ñðàâíèòåëüíî-àíàòîìè÷åñêèõ äàííûõ â îòäåëüíûé òèï Priapulida ñëåäóåò ââåñòè òðè êëàññà: Priapulida, Kinorhyncha è Archeopriapulida.

16 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.1. Âíåøíèé âèä Priapulus caudatus ñ áðþøíîé ñòîðîíû. CF — öèðêóìî- ðàëüíîå ïîëå; IN — èíòðîâåðò; PA — ïðåàíàëüíûé ïîäîòäåë; TA — õâîñò; TR — òóëîâèùå (= àáäîìåí); VN — áðþøíîé íåðâíûé ñòâîë. Fig. 3.1. Priapulus caudatus, ventral view. CF — circumoral field; IN — introvert; PA — preanal subregion; TA — tail; TR — trunk (=abdomen); VN — ventral nerve cord.

Ìû ðàññìàòðèâàåì Priapulida êàê îòäåëüíûé êëàññ â ñîñòàâå òèïà Cephalorhyncha Malakhov, 1980 âìåñòå ñ êëàññàìè Kinorhyncha, Loricifera è Nema- tomorpha (Ìàëàõîâ, 1980; Àäðèàíîâ, Ìàëàõîâ, 1995à–æ; 1996; Adrianov, Malakhov, 1995, 1996a; Ìàëàõîâ, Àäðèàíîâ, 1995). Ñîâðåìåííûå è èñêîïàåìûå ïðèàïóëèäû îáû÷- íî ðàññìàòðèâàþòñÿ îòäåëüíî. Îäíà èç ïîñëåäíèõ ñèñòåì ñîâðåìåííûõ ïðèàïóëèä ïðåäëîæåíà Ëàí- äîì (Land, 1970) (òàáë. 3.1). Èñêîïàåìûå ïðèàïó- ëèäû è äðóãèå êåìáðèéñêèå öåôàëîðèíõè (ïîä îá- ùèì íàçâàíèåì Archaeopriapulida) âîøëè â ñèñòå- ìó, ïðåäëîæåííóþ Êîíâåé Ìîððèñîì (Conway Morris, 1977) (òàáë. 3.2). Íåäàâíî âïåðâûå áûëà ïðåäëîæåíà îáùàÿ ñèñ- òåìà ïðèàïóëèä, ñîåäèíèâøàÿ êàê ñîâðåìåííûõ, òàê è èñêîïàåìûõ ôîðì (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996). Äîïîëíåííûé âàðèàíò ýòîé ñèñòåìû ïðåä- ñòàâëåí íà òàáë. 3.3.  ñèëó ñïåöèôèêè ðàáîòû ñ ïàëåîíòîëîãè÷åñ- êèì ìàòåðèàëîì, â íàñòîÿùåé ðàáîòå ìû îãðàíè- ÷èâàåìñÿ îïðåäåëèòåëåì ôàóíû òîëüêî íûíåæèâóùèõ ïðèàïóëèä è íå âêëþ÷àåì â îïðåäåëèòåëü- íûå òàáëèöû âûìåðøèå âèäû.

3.2. Ïëàí ñòðîåíèÿ è îñîáåííîñòè ìîðôîëîãèè ïðèàïóëèä, èñïîëüçóåìûå â ñèñòåìàòèêå

3.2.1. Ïëàí ñòðîåíèÿ, âíåøíÿÿ ìîðôîëîãèÿ è ñèììåòðèÿ ïðèàïóëèä

Ïðèàïóëèäû — ýòî ÷åðâåîáðàçíûå îðãàíèçìû, ðàçìåðû òåëà êîòîðûõ êîëåáëþòñÿ îò 40 ñì (Halicryptus higginsi) äî 1,5–2 ìì (Tubiluchus). Òåëî ïðèàïóëèä, êàê ó âñåõ ãîëîâîõîáîòíûõ ÷åðâåé, ðàçäåëÿåòñÿ íà äâà îñíîâíûõ îòäåëà: õîáîò (=ãîëîâà, èíòðîâåðò, ïðåñåïòóì) è òóëîâèùå (=àáäîìåí, ïîñòñåïòóì) (ðèñ. 3.1, 3.2). Íåêî- òîðûå ïðèàïóëèäû èìåþò òðåòèé îòäåë òåëà — õâîñò, ïðåäñòàâëåííûé 1 èëè 2 êàóäàëüíûìè ïðèäàòêàìè (ðèñ. 3.1–3.4). Õîáîò ñïîñîáåí öåëèêîì ââîðà÷èâàòüñÿ â òóëîâèùíûé îòäåë, êîòîðûé ó íåêîòîðûõ ïðèàïóëèä îòäåëåí îò èíòðîâåðòà êîðîòêîé øåéíîé îáëàñòüþ (ðèñ. 3.2). Ïåðåäíèé ó÷àñòîê õîáîòà ïðåäñòàâëåí öèðêóìîðàëüíûì ïîëåì (ó ìàêðîáåíòîñíûõ ïðèàïóëèä Priapulus, Priapulopsis, Acanthopriapulus è Halicryptus) èëè ðîòîâûì êîíóñîì (ó ìåéîáåíòîñíûõ ôîðì Tubiluchus, Meiopriapulus, Maccabeus) (ðèñ. 3.1–3.9). Çàäíÿÿ ãðàíèöà öèðêóìîðàëüíîãî ïîëÿ (èëè ðîòîâîãî êîíóñà) ìàðêèðóåòñÿ êðóãîì èç 8 öèðêóìîðàëüíûõ ñêàëèä è çàëåãàþùèì ïîä íèì íåðâíûì êîëüöîì (Land, 1970; Storch et al, 1990; Storch, 1991; Àäðèàíîâ, Ìàëàõîâ, 1995à–å; Àäðèàíîâ, Ìàëàõîâ, 1996). Ñïåðåäè îêîëîðîòîâîå ïîëå ìàêðîáåíòîñíûõ ïðèàïóëèä îãðàíè÷åíî ïåðåäíèìè êðóãàìè çóáîâ âûâîðà÷èâàþùåéñÿ ãëîòêè.  êàæäîì òàêîì êðóãå ïî 5 çóáîâ, ïðè÷åì â ñîñåäíèõ êðóãàõ çóáû ÷åðåäóþòñÿ äðóã ñ äðóãîì (ðèñ. 3.10–3.12). Îêîëîðîòîâîå ïîëå ìàêðîáåíòîñ- íûõ ïðèàïóëèä íåñåò ìíîãî÷èñëåííûå è õàîòè÷íî ðàñïîëîæåííûå ïðèäàòêè — áóêêàëüíûå ïà- ïèëëû è ìåëêèå ñêàëèäû. Ïî ñâîåé ìîðôîëîãèè ïàïèëëû è ñêàëèäû öèðêóìîðàëüíîãî ïîëÿ ñõîä- íû ñî ñêàëèäàìè öåíòðàëüíîé ÷àñòè èíòðîâåðòà, íî îòëè÷àþòñÿ ìåíüøèìè ðàçìåðàìè.

17 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

Ðèñ. 3.2. Ïðåäñòàâèòå- ëè èñêîïàåìûõ (1, 2) è ñîâðåìåííûõ ïðèàïóëèä (3–12) (1–2 ïî Conway Morris, 1977; 11 — ïî Morse, 1981; 12 — ïî Por, Bromley, 1974): 1 — Ottoia prolifica; 2 — Selkirkia co- lumbia; 3 — Priapulus cau- datus; 4 — P. abyssorum; 5 — P. tuberculatospinosus; 6 — Acanthopriapulus horri- dus; 7 — Priapulopsis bicau- datus; 8 — P. australis; 9 — Halicryptus spinulosus; 10 — Tubiluchus corallicola; 11 — Meiopriapulus fijiensis; 12 — Maccabeus tentaculatus. Fig. 3.2. Fossil (1–2) and recent (3–12) priapu- lids (1–2 after Conway Morris, 1977; 11 — after Morse, 1981; 12 — after Por & Bromley, 1974): 1 — Ottoia prolifica, 2 — Sel- kirkia columbia, 3 — Pria- pulus caudatus, 4 — P. abys- sorum, 5 — P. tuberculato- spinosus, 6 — Acanthopria- pulus horridus, 7 — Pria- pulopsis bicaudatus, 8 — P. australis, 9 — Halicryptus spinulosus, 10 — Tubiluchus corallicola; 11 — Meiopria- pulus fijiensis, 12 — Mac- cabeus tentaculatus.

Ó ìåëêèõ ñèäÿ÷èõ ïðèàïóëèä ðîäà Maccabeus, ó÷àñòîê èíòðîâåðòà, ãîìîëîãè÷íûé öèðêóìî- ðàëüíîìó ïîëþ, îáîñîáëåí â âèäå êîðîòêîãî ðîòîâîãî êîíóñà (Por, 1973; Por, Bromley, 1974; Salvini-Plawen, 1974). Ñçàäè îí òàêæå îãðàíè÷åí âåí÷èêîì èç 8 îïóøåííûõ öèðêóìîðàëüíûõ ñêà- ëèä, èëè “òðèããåðíûõ øèïîâ”, ìàðêèðóþùèõ ïîëîæåíèå íåðâíîãî êîëüöà (ðèñ. 3.7A–B). Ñðåäè âçðîñëûõ ïðèàïóëèä ðîòîâîé êîíóñ ëó÷øå âñåãî âûðàæåí ó ìåéîáåíòîñíûõ Tubiluchus (Land, 1968, 1970, 1982, 1985; Kirsteuer, Land, 1970; Àäðèàíîâ è äð, 1989; Àäðèàíîâ, Ìàëàõîâ, 1991) è

18 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.4. Priapulopsis australis, çàäíÿÿ ÷àñòü òóëîâèùà è õâîñòîâûå ïðèäàòêè. AN — àíóñ; CG — êàóäàëüíûé ãàí- ãëèé; LT — ëåâûé õâîñòîâîé ïðèäàòîê; NC — âåíòðàëü- Ðèñ. 3.3. Priapulopsis australis, âíåøíèé âèä ñ áðþø- íûé íåðâíûé ñòâîë; RP — êîëüöåâûå ïàïèëëû; RT — íîé ñòîðîíû. IN — èíòðîâåðò; NC — âåíòðàëüíûé íå- ðóäèìåíòàðíûé ïðàâûé õâîñòîâîé ïðèäàòîê; TV — õâîñ- ðâíûé ñòâîë; RP — êîëüöåâûå ïàïèëëû; TA — õâîñò; òîâûå âåçèêóëû. Ìàñøòàá 1 ìì. TR — òóëîâèùå. Ìàñøòàá 1 ìì. Fig. 3.4. Priapulopsis australis, posterior trunk and tail. AN Fig. 3.3. Priapulopsis australis, external morphology, — anal opening; CG — caudal ganglium; LT — left tail ventral view. IN — introvert; NC — ventral nerve cord; RP branch; NC — ventral nerve cord; RP — ring papillae; RT — — ring papillae; TA — tail; TR — trunk. Bar 1 mm. rudimental (right) tail branch; TV — tail vesicles. Bar 1 mm.

19 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

Ðèñ. 3.6. Îðãàíèçàöèÿ êàðëèêîâûõ ïðè- àïóëèä Tubiluchus: A — âíåøíèé âèä (ñáî- êó) # T. corallicola, ìàñøòàá 0,5 ìì; B — ñõåìà ïðîäîëüíîãî ðàçðåçà $ T. arcticus; C — ãðåáåí÷àòûå çóáû; D — ùåòèíêà ðîòî- âîãî êîíóñà; E — ÷óâñòâèòåëüíûå ïðèäàò- êè (1 — êëàâóëà, 2–4 ìîäèôèöèðîâàííûå òóáóëè) ãåíèòàëüíîãî ïîëÿ ñàìöà T. coral- licola (ïî Land, 1970); F — ïðèäàòêè (1–2) ïîëèòèðèäèóìà; G — ôëîñêóëÿ; H — êëà- âóëà T. remanei; I — òóáóëÿ ëè÷èíêè T. remanei. BC — êëåòêà ïîëîñòè òåëà, CM — ñêëàäêà êîëüöåâîé ìóñêóëàòóðû, CN — îêîëîãëîòî÷íîå íåðâíîå êîëüöî, CS — öèðêóìîðàëüíûå ñêàëèäû, F — ôëîñêóëÿ, G — ãîíàäà ñàìêè, I — èíòðîâåðò, L — ýêñòðàêëåòî÷íûé ëèãàìåíò. MC — ðîòîâîé êîíóñ, N — øåéíàÿ îáëàñòü, P — ïîëèòè- ðèäèóì, PH — ãëîòêà, PN — ïðîòîíåôðè- äèé, S — ñêàëèäû, R — ìûøöû-ðåòðàêòî- ðû èíòðîâåðòà, RS — ñåìåïðèåèíèê, Ò — õâîñò, TU — òóáóëÿ, U — óðîãåíèòàëüíûé îðãàí. Fig. 3.6. Morphology of the dwarf priapulid Tubiluchus: A — # of T. corallicola, lateral view, bar 0.5 mm; B — scheme of longitudinal section of $ of T. arcticus; C — pectinate teeth; D — bristle of mouth cone; E — sensory appendages (1 — clavula; 2–4 modified tubuli) of male genital field of T. corallicola (after Land, 1970); F — appendages (1–2) of polythyridium; G — flosculus; H — clavula of T. remanei; I — tubulus of lavral T. remanei. BC — cell of body cavity, CM — fold of circular musculature, CN — circumpharyngeal nerve ring, CS — circumoral scalids, F — flosculus, G — female gonad, I — introvert, L — extracellular ligament, MC — mouth cone, N — neck area, P — polythyridium, PH — pharynx, PN — protonephridium, S — scalids, R — retractor muscles of introvert, RS — receptaculum seminis, T — tail, TU — tubulus, U — urogenital organ.

Ðèñ. 3.5. Halicryptus spinulosus. À — èíòðîâåðò, âèä ñ áðþøíîé ñòîðîíû; co — ðîòîâîé êîíóñ (öèðêóìîðàëüíîå ïîëå); ds — äåíòîñêàëèäà; fl — ôëîñêóëÿðíàÿ ïàïèëëà; ng — øåéíûé ãàíãëèé; vn — áðþøíîé íåðâíûé ñòâîë; nr — îêîëîãëîòî÷íîå íåðâíîå êîëüöî; p — áóêêàëüíàÿ ïàïèëëà; ph — ãëîòêà; rs — “çà÷àòî÷íàÿ ñêàëèäà; ss1 — öèðêóìîðàëüíàÿ ñêàëèäà; ss — ñïèíîñêàëèäà; s — æåëåçèñòûé øèï; t1 — ïðèäàòîê 1-ãî ïåíòàãîíà; t2 — çóá 2-ãî ïåíòàãîíà; öèôðàìè 1–21 îáîçíà÷åíû êðóãè ñêàëèä.  — äèàãðàììà ïðèäàòêîâ èíòðîâåðòà; PR — ïåðâè÷íûå ðàäèóñû èíòðîâåòà; VS — âåíòðàëüíûé ñåêòîð; VR — âåíòðîëà- òåðàëüíûé ïðàâûé ñåêòîð; LR — ëàòåðàëüíûé ïðàâûé ñåêòîð; DR — äîðñîëàòåðàëüíûé ïðàâûé ñåêòîð; DS — äîðñàëüíûé ñåêòîð; DL — äîðñîëàòåðàëüíûé ëåâûé ñåêòîð; LL — ëàòåðàëüíûé ëåâûé ñåêòîð; VL — âåíòðîëàòåðàëüíûé ëåâûé ñåêòîð; öèôðàìè 1–25 íà äèàãðàììå îáîçíà÷åíû âòîðè÷íûå ðàäèóñû èíòðîâåðòà; 1 — öèðêóìîðàëüíàÿ ñêàëèäà; 2 — ñïèíîñêàëèäà; 3 — äâóçóáàÿ äåíòîñêàëèäà; 4 — îäíîçóáàÿ äåíòîñêàëèäà; 5 — “çà÷àòî÷íûå” ñêàëèäû; 6 — æåëåçèñòûé øèï. Ìàñøòàá 0,1 ìì. Fig. 3.5. Halicryptus spinulosus. À — introvert, ventral view; co — mouth cone (circumpharyngeal field); ds — dentoscalid; fl — floscular papilla; ng — neck ganglium; vn — ventral nerve cord; nr — circumpharyngeal nerve ring; p — buccal papilla; ph — pharynx; rs — developing scalid; ss1 — circumoral scalid; ss — spinoscalid; s — glandular spine; t1 — element of anteriormost pentagon; t2 — tooth of 2nd pentagon; numbers 1–21 indicate ordinal numbers of scalid circlets. B — diagrammatic representation of scalid arrangement; PR — primary radii of introvert; VS — ventral sector; VR — ventrolateral right sector; LR — lateral right sector; DR — dorsolateral right sector; DS — dorsal sector; DL — dorsolateral left sector; LL — lateral left sector; VL — ventrolateral left sector; numbers 1–25 on diagramm indicate secondary radii of introvert; 1 — circumoral scalid; 2 — spinoscalid; 3 — bifurcated dentoscalid; 4 — dentoscalid with single tooth; 5 — developing scalids; 6 — glandular spine. Bar 0.1 mm.

20 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.7. Maccabeus tentaculatus: A — ñõåìà ñòðîåíèÿ (ïî Por, 1983); B — âíåøíèé âèä $ (ïî Por, Bromley, 1974), ìàñøòàá 0,5 ìì; C — ãðåáåí÷àòûå çóáû âçðîñëîé îñîáè, ìàñøòàá 30 ìêì; D — èíòðîâåðò ëè÷èíêè; E — ëîðèêà ëè÷èíêè; F — ëè÷èíî÷íûå çóáû (C, D, E, F — ïî Salvini-Plaven, 1974). À — àíóñ,  — òóëîâèùå, CM — ÷åõîë êîëüöåâîé ìóñêóëàòóðû, CS — öèðêóìîðàëüíûå ñêàëèäû (òðèããåð- íûå øèïû), G — ãîíàäà, Í — ïðåàíàëüíûå êðþ÷üÿ, I — öåíòðàëüíàÿ ÷àñòü èíòðîâåðòà, L — ýêñòðàêëåòî÷íûé ëèãàìåíò. Ì — ðåòðàêòîð, MC — ââåðíóòûé ðîòîâîé êîíóñ, N — íåðâíîå êîëüöî, PN — ïðîòîíåôðèäèé, S — ñêàëèäû èíòðîâåðòà, Ò — çóáû. Fig. 3.7. Maccabeus tentaculatus: A — scheme of body structure (after Por, 1983); B — $ (after Por & Bromley, 1974), bar 0.5 mm; C — pectinate teeth of adult specimen, bar 30 mc; D — larval introvert; E — larval lorica; F — larval teeth (C, D, E, F — after Salvini-Plaven, 1974). A — anus, B — trunk, CM — muscular bell of circular fibres, CS — circumoral scalids (trigger spines), G — gonad, H — preanal hooks, I — central portion of introvert, L — extracellular ligament, M — retractor muscle, MC — with- drawn mouth cone, N — nerve ring, PN — protonephri- dium, S — scalids of introvert, T — teeth.

Ðèñ. 3.8. Âíåøíÿÿ ìîðôîëîãèÿ Maccabeus: À–G — M. tentaculatus (ïî Salvini-Plawen, 1974); H–O — M. cirratus (ïî Ìàëàõîâ, 1978); À–Í — âíåøíèé âèä;  — öèðêóìî- ðàëüíàÿ ñêàëèäà (òðèããåðíûé øèï); C–E, J–L — ñêàëè- äû öåíòðàëüíîé ÷àñòè èíòðîâåðòà; F, N — ïðåàíàëüíûé êðþê; G, O — àíàëüíàÿ òóáóëÿ. ah — ïðåàíàëüíûå êðþ- ÷üÿ; at — àíàëüíûå òóáóëè; fo — çîíà ïðîäîëüíî ñêëàä- ÷àòîé êóòèêóëû; ne — ïàðàçèòè÷åñêàÿ íåìàòîäà â ïîëî- ñòè òåëà; pa — ïðåàíàëüíàÿ îáëàñòü; sc — ñêàëèäû öåíò- ðàëüíîé çîíû èíòðîâåðòà; te — ïåðåäíÿÿ êîðîíà ùóïà- ëåö; tr — òóëîâèùå. Ìàñøòàá 0,5 ìì. Fig. 3.8. External morphology of Maccabeus: A–G — M. tentaculatus (after Salvini-Plawen, 1974); H–O — M. cirratus (after Malakhov, 1978); A–H — general view; B — circumoral scalid (Trigger spine); C–E, J–L — scalids of the central area of introvert; F, N — preanal hook; G, O — anal tubulus. ah — preanal hook; at — anal tubuli; fo — area of longitudinally folded cuticle; ne — parasitic nematode in the body cavity; pa — preanal area; sc — scalids of the central area of introvert; te — tentacles; tr — trunk. Bar 0.5 mm.

21 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

Meiopriapulus (Morse, 1981; Storch et al, 1989) (ðèñ. 3.6A– B; 3.9). Meiopriapulus èìåþò äëèííûé öèëèíäðè÷åñêèé ðîòîâîé êîíóñ, íåñóùèé ñîá- ñòâåííîå âîîðóæåíèå èç ìíî- ãî÷èñëåííûõ ãðåáåí÷àòûõ çó- áîâ (ðèñ. 3.9; 3.11B). Öåíòðàëüíàÿ ÷àñòü èíòðî- âåðòà ïðèàïóëèä íåñåò çàãíó- òûå íàçàä ñåíñîðíî-ëîêîìî- òîðíûå ïðèäàòêè — ñêàëèäû, ÷àñòü èç êîòîðûõ âûïîëíÿåò è æåëåçèñòóþ ôóíêöèþ (ðèñ. 3.1–3.9; 3.11B; 3.13–3.16). Ðàñ- ïîëîæåíèå ñêàëèä õàðàêòåðè- çóåòñÿ êîìáèíàöèåé ðàçëè÷- íûõ òèïîâ ñèììåòðèè è ïîä- ÷èíÿåòñÿ ñëîæíûì çàêîíî- ìåðíîñòÿì. Ó âñåõ ïðèàïóëèä ïåðâûé êðóã ñîäåðæèò 8 öèðêóìîðàëü- íûõ ñêàëèä, îïðåäåëÿþùèõ 8 ïåðâè÷íûõ ðàäèóñîâ è 8 ñåê- òîðîâ: äîðñàëüíûé, âåíòðàëü- íûé, ëåâûé è ïðàâûé ëàòå- ðàëüíûå, à òàêæå ëåâûé è ïðà- âûé ñóáäîðñàëüíûå (=äîðñî- ëàòåðàëüíûå) è ëåâûé è ïðà- âûé ñóáâåíòðàëüíûå (=âåíòðî- ëàòåðàëüíûå) (ðèñ. 3.5B; 3.9; 3.11B; 3.13; 3.16). Ïîñëåäóþùèå ñêàëèäû ó ìàêðîáåíòîñíûõ ïðèàïóëèä ðàñïîëîæåíû êðó- ãàìè, òàê ÷òî ñêàëèäû â ñîñåä- íèõ êðóãàõ ÷åðåäóþòñÿ, òî åñòü èìååò ìåñòî øàõìàòíûé ïîðÿ- Ðèñ. 3.9. Âíåøíÿÿ ìîðôîëîãèÿ Meiopriapulus fijiensis (ñïðàâà — ñõåìà ðàñïîëî- äîê ðàñïîëîæåíèÿ ïðèäàòêîâ. æåíèÿ ñêàëèä ïåðâûõ òðåõ êðóãîâ). (Ðèñóíîê ñäåëàí ñ ýêçåìïëÿðà, ëþáåçíî ïðå- 2-é, 5-é, 8-é, 11-é è ò.ä. êðó- äîñòàâëåííîãî äîêòîðîì Ð. Õèããèíñîì). an — àíóñ; em — ýìáðèîíû â ïðàâîì óðîãåíèòàëüíîì îðãàíå; hg — çàäíÿÿ êèøêà; ls — ëîêîìîòîðíàÿ ñêàëèäà; mc — ãè, ïîðÿäêîâûé íîìåð êîòîðûõ ðîòîâîé êîíóñ; mg — ñðåäíÿÿ êèøêà; ns — øåéíàÿ, èëè òóëîâèùíàÿ, ñêàëèäà; S1

îïðåäåëÿåòñÿ ôîðìóëîé 2+3n — ñêàëèäû 1-ãî êðóãà; S2 — ñêàëèäû 2-ãî êðóãà; S3 — ñêàëèäû 3-ãî êðóãà; so — (ãäå n=0, 1, 2, 3...), ñîäåðæàò ñîìàòè÷åñêèå ÷óâñòâèòåëüíûå îðãàíû; to — çóá; tu — òóìóëè; uo — ëåâûé óðîãå- êàæäûé ïî 9 ñêàëèä. 3-é, 6-é, íèòàëüíûé îðãàí; vn — âåíòðàëüíûé íåðâíûé ñòâîë. Ìàñøòàá 0.1 ìì. Fig. 3.9. External morphology of Meiopriapulus fijiensis (on the right — diagram of 9-é, 12-é è ò.ä. êðóãè, ïîðÿä- scalid arrangement (1st–3rd rings)). (The specimen was kindly provided by Dr. R. Higgins). êîâûé íîìåð êîòîðûõ çàäàåò- an — anal opening; em — embryons in the right urogenital organ; hg — hindgut; ls — locomotory scalid; mc — mouth cone; mg — midgut; ns — neck, or trunk, scalid; S ñÿ ôîðìóëîé 3+3n, ñîäåðæàò 1 — scalid of 1st ring; S — scalid of 2nd ring; S — scalid of 3rd ring; so — trunk sensory êàæäûé ïî 8 ñêàëèä. Òî÷íî òàê- 2 3 organs; to — tooth; tu — tumuli; uo — left urogenital organ; vn — ventral nerve cord. æå è 4-é, 7-é, 10-é, 13-é è ò.ä. Bar 0.1 mm. êðóãè, íîìåðà êîòîðûõ îïðå- äåëÿþòñÿ ôîðìóëîé 4+3n, ñîäåðæàò êàæäûé ïî 8 ñêàëèä. Ñêàëèäû 3+3n è 4+3n êðóãîâ ðàñïîëîæå- íû ïî îäíîé â êàæäîì èç 8 ñåêòîðîâ èíòðîâåðòà. ×òî êàñàåòñÿ ñêàëèä 2+3n êðóãîâ, òî â âåíòðàëü- íîì ñåêòîðå ðàñïîëîæåíî ïî 2 ñêàëèäû, à â êàæäîì èç îñòàëüíûõ ñåêòîðîâ — ïî 1. ×èñëî êðóãîâ

22 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.11. A — äèàãðàììà ãëîòî÷íûõ çóáîâ Priapulus caudatus. CS — öèðêóìîðàëü- íàÿ ñêàëèäà; öèôðàìè 1–7 îáîçíà÷åíû íîìåðà ïåíòàãî- íîâ.  — äèàãðàììà ïðèäàò- êîâ èíòðîâåðòà è çóáîâ Meio- priapulus fijiensis. R — ãëîòî÷- íûå ðàäèóñû; öèôðàìè 1–17 íà äèàãðàììå îáîçíà÷åíû âòîðè÷íûå ðàäèóñû èíòðî- âåðòà; öèôðàìè 1–7 íà îê- òîãðàììå îáîçíà÷åíû êðó- ãè îêòîãîíîâ; I — öèðêóìî- ðàëüíûå ñêàëèäû; II — ñêà- ëèäû 2-ãî êðóãà; III — ñêà- ëèäû 3-ãî êðóãà; IV — ëîêî- ìîòîðíûå ñêàëèäû; V — ôëîñêóëè ó îñíîâàíèé öèð- êóìîðàëüíûõ ñêàëèä; äðóãèå óñëîâíûå îáîçíà÷åíèÿ êàê íà ðèñ. 3.5Â. Fig. 3.11. A — diagram- matic representation of tooth arrangement in Priapulus cau- datus. CS — circumoral scalid; numbers 1–7 indicate ordinal numbers of pentagons. B — diagrammatic represerntation of scalid and tooth arrangements in Meiopriapulus fijiensis. R — pharyngeal radii; numbers 1– 17 on diagramm indicate secon- dary radii of introvert; numbers 1–7 on octogramm indicate ordinal numbers of octogons; I — circumoral scalids; II — sca- Ðèñ. 3.10. Äèàãðàì- lids of 2nd circlet; III — scalids ìû ðàñïîëîæåíèÿ ïå- of 3rd circlet; IV — locomotory ðåäíèõ çóáíûõ ïåíòàãî- scalids; V — flosculus. Other íîâ: À — Priapulopsis abbreviations as in fig. 3.5B. australis; B — P. bicau- datus; C — Priapulus tuberculatospinosus; D — P. abyssorum. CS — öèð- êóìîðàëüíàÿ ñêàëèäà; GA — ïðîìåæóòîê íå- âîîðóæåííîé êóòèêóëû ìåæäó ïÿòüþ ïåðâûìè è ïîñëåäóþùèìè ïåí- òàãîíàìè. Öèôðàìè îáîçíà÷åíû ñîîòâåòñò- âóþùèå ïåíòàãîíû. Fig. 3.10. Diagrams of anterior tooth pentagons: À — Priapulopsis australis; B — P. bicaudatus; C — Priapulus tuberculatospi- nosus; D — P. abyssorum. CS — circumoral scalids; GA — gap between 5 anterior pentagons and following circlets. Num- bers indicate correspon- ding pentagons.

23 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

Ðèñ. 3.12. Halicryptus spinulosus: A — äèàãðàììà ãëîòî÷íûõ çóáîâ, ñî- âìåùåííàÿ ñ êîëüöåâîé äèàãðàììîé 4 ïåðâûõ êðóãîâ ñêàëèä; R — ãëî- òî÷íûå ðàäèóñû; öèôðàìè 1–7 íà ïåíòîãðàììå îáîçíà÷åíû íîìåðà ïåíòàãîíîâ; âñå äðóãèå óñëîâíûå îáîçíà÷åíèÿ êàê íà ðèñ. 3.5Â.  — âû- âåðíóòîå ãëîòî÷íîå âîîðóæåíèå; öèôðàìè 1–17 îáîçíà÷åíû íîìåðà ïåíòàãîíîâ; ñâåòëûì ïîêàçàíû çóáû íà íå÷åòíûõ ãëîòî÷íûõ ðàäèóñàõ; òåìíûì ïîêàçàíû çóáû íà ÷åòíûõ ãëîòî÷íûõ ðàäèóñàõ. Fig. 3.12. Halicryptus spinulosus: A — diagrammatic representation of tooth arrangement and circular diagramm of 4 anterior circlets of scalid; R — pha- ryngeal radii; numbers 1–7 on penta- gramm indicate ordinal numbers of pen- tagons. Other abbreviations as in fig. 3.5B. B — everted pharyngeal armament: numbers 1–17 indicate ordinal numbers of pentagons; light elements — teeth on odd pharyngeal radii; dark elements — teeth on even pharyngeal radii.

Ðèñ. 3.13. Tubiluchus vanuatensis: À — èíòðîâåðò, âèä ñ áðþøíîé ñòîðîíû (ìàñøòàá 30 ìêì); BS — áóêêàëüíàÿ ñêàëèäà; CF — öèðêóìîðàëüíîå ïîëå; CS — öèðêóìîðàëüíàÿ ñêàëèäà; FL — ôëîñêóëÿ; IN — öåíòðàëüíàÿ ÷àñòü èíòðî- âåðòà; NE — øåéíàÿ îáëàñòü; NR — îêîëîãëîòî÷íîå íåðâíîå êîëüöî; VN — áðþøíîé íåðâíûé ñòâîë; öèôðàìè 1–23 îáîçíà÷åíû êðóãè ñêàëèä.  — äèàãðàììà ïðèäàòêîâ èíòðîâåðòà; öèôðîé 23 îáîçíà÷åíî ÷èñëî êðóãîâ ñêàëèä; 1 — öèðêóìîðàëüíàÿ ñêàëèäà; 2 — ñêàëèäà; âñå äðóãèå óñëîâíûå îáîçíà÷åíèÿ êàê íà ðèñ. 3.5Â. Fig. 3.13. Tubiluchus vanuatensis: À — introvert, ventral view (bar 30 microns). BS — buccal scalid; CF — circumpharyngeal field; CS — circumoral scalid; FL — flosculus; IN — central portion of introvert; NE — neck area; NR — circumpharyngeal nerve ring; VN — ventral nerve cord; numbers 1–23 indicate ordinal numbers of scalid circlets. B — diagrammatic representation of scalid; number 23 indicates total number of scalid circlets; 1 — circumoral scalid; 2 — scalid. Other abbreviations as in fig. 3.5B.

24 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.14. Ïàïèëëà öèðêóìîðàëüíîãî ïîëÿ (P) è ñêàëèäû ìîëîäîãî H. spinulosus. Öèôðû îçíà÷àþò íîìåð êðóãà, â êîòî- ðîì ðàñïîëàãàþòñÿ èçîáðàæåííûå ñêàëèäû. Ìàñøòàá 50 ìêì. Fig. 3.14. Papilla of circumoral field (P) and scalids of a young H. spinulosus. Numerals indicate the corresponding scalid rings. Bar 0.05 mm. Ðèñ. 3.15. Ñîìàòè÷åñêèå îðãàíû ÷óâñòâ ìîëîäîãî H. spinulosus: À,  — îäèíî÷íàÿ ôëîñêóëÿ, Ñ — äâîéíàÿ ôëîñêó- ëÿ, D — ôëîñêóëÿ ñ òóáóëåé, Å — ÷óâñòâèòåëüíûé øèï, F — ÷óâñòâèòåëüíàÿ ïàïèëëà, G — êàóäàëüíûé øèï. fl — ôëîñ- êóëÿ, tu — òóáóëÿ. Màñøòàá: À–D — 5 ìêì, Å–G — 50 ìêì. Fig. 3.15. Somatic sensory organ of a young H. spinulosus: A, B — solitary floscula, C — double floscula, D — floscula and tubula, e — sensory spine, F — sensory papilla, G — caudal spine. fl — floscula, tu — tubula. Bar: A–D — 0.005 mm, E–G — 0.05 mm.

Ðèñ. 3.16. Ðàñïîëîæåíèå ñêàëèä íà èíòðîâåðòå Priapulus caudatus. Öèôðàìè îáîçíà÷åíû ãëàâíûå (ïåðâûå) ñêàëèäû ñåðèé, ñîîòâåòñòâóþùèå êðóãàì ñêàëèä ó ëè÷èíîê (1 — öèðêóìîðàëüíûå ñêàëèäû; 2–10 — ñïèíîñêàëèäû). Ìàñø- òàá 150 ìèêðîí. Fig. 3.16. Arrangement of scalids in Priapulus caudatus. Numbers indicate primary scalids of series corresponding to larval rings (1 — circumoral scalids; 2–10 — spinoscalids). Bar 150 microns.

óâåëè÷èâàåòñÿ ñ âîçðàñòîì è ïîñòåïåííî çà ñ÷åò êîìáèíàöèè 9+8+8 âîçíèêàþò 25 ïðîäîëüíûõ ðÿäîâ, ïðè÷åì ïåðâè÷íûå ðàäèóñû, ðàçäåëÿþùèå ñåêòîðà, íå ñîâïàäàþò íè ñ îäíèì èç 25 ðÿäîâ.  âåíòðàëüíîì ñåêòîðå îêàçûâàþòñÿ 4 ðÿäà ñêàëèä, òîãäà êàê â äðóãèõ — òîëüêî ïî 3 ðÿäà. Êîìáèíà- öèÿ 8 ñêàëèä 1-ãî êðóãà è 25 ðÿäîâ ñêàëèä, îáðàçîâàííûõ ïîñëåäóþùèìè êðóãàìè, à òàêæå ðàçëè÷- íîå êîëè÷åñòâî ðÿäîâ â âåíòðàëüíîì è äîðñàëüíîì ñåêòîðàõ ïðèäàþò èíòðîâåðòó â öåëîì áèëàòå- ðàëüíóþ ñèììåòðèþ. Äâà ñáëèæåííûõ ðÿäà, ïðîõîäÿùèõ ïîñðåäèíå âåíòðàëüíîãî ñåêòîðà è ñîñòî- ÿùèõ èç ñêàëèä 2+3n êðóãîâ, ìàðêèðóþò ïîëîæåíèå áðþøíîãî íåðâíîãî ñòâîëà. Èíîãäà (êàê íàïðèìåð, ó Halicryptus spinulosus) íåñêîëüêî çàäíèõ êðóãîâ èç 2+3n ïîñëåäîâà- òåëüíîñòè ñîäåðæàò íå ïî 9, à âñåãî ïî 7 ñêàëèä çà ñ÷åò èõ îòñóòñòâèÿ â âåíòðàëüíîì ñåêòîðå (ðèñ. 3.5).  ðåçóëüòàòå ýòîãî 2 ðÿäà, îêàéìëÿþùèõ áðþøíîé íåðâíûé ñòâîë, îáû÷íî íåñêîëüêî êîðî÷å îñòàëüíûõ. Íàäî îòìåòèòü, ÷òî óæå ñ 5-ãî êðóãà äâå ñáëèæåííûå âåíòðàëüíûå ñêàëèäû âñåõ êðóãîâ èç 9 ïðèäàòêîâ îêàçûâàþòñÿ íåñêîëüêî ñìåùåííûìè äðóã îòíîñèòåëüíî äðóãà — â êàæäîé ïàðå ëåâàÿ ñêàëèäà ñäâèíóòà ÷óòü âïåðåä ïî îòíîøåíèþ ê ïðàâîé (ðèñ. 3.5; 3.13).

25 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

Ðèñ. 3.17. Èíòðîâåðò Priapulopsis australis: À — ðàñïîëîæåíèå ñêàëèä;  — ïåðåäíèå ñêàëèäû; Ñ — òåëåñêîïè÷åñêàÿ ñïèíîñêàëèäà ïåðåäíåé ñåðèè; D — ïåðåäíÿÿ ñåðèÿ ñêàëèä; Å — çàäíèå çà÷àòî÷íûå ñêàëèäû. BS — áóêêàëüíàÿ ñêàëèäà öèðêóìîðàëüíîãî ïîëÿ; CF — öèðêóìîðàëüíîå ïîëå; CS — öèðêóìîðàëüíàÿ ñêàëèäà; SS — ñïèíîñêàëèäà ïåðåäíåé ñåðèè. Ìàñøòàá: À — 0,4 ìì; Â, Ñ, Å — 60 ìêì, D — 30 ìêì. Fig. 3.17. Introvert of Priapulopsis australis: A — arrangements of scalids; B — anterior scalids; C — telescopic spinoscalid of anterior series; D — anterior series of scalids; E — posterior developing scalids. BS — buccal scalid of circumoral field; CF — circumoral field; CS — circumoral scalid; SS — spinoscalid of anterior series. Bar: A — 0.4 mm; B, C, E — 60 mc; D — 30 mc.

Ðàñïîëîæåíèå è ôîðìà ñêàëèä ÿâëÿþòñÿ âàæíûìè ñèñòåìàòè÷åñêèìè ïðèçíàêàìè (3.14–3.18). Ñêàëèäû èíòðîâåðòà Halicryptus äèôôåðåíöèðîâàíû íà íåñêîëüêî ìîðôîëîãè÷åñêèõ òèïîâ: (1) êîíè÷åñêèå äâó÷ëåíèñòûå öèðêóìîðàëüíûå ñêàëèäû, ïîäîáíî áóêêàëüíûì ïàïèëëàì, ãóñòî îïóøåííûå êóòèêóëÿðíûìè âîëîñêàìè; (2) äëèííûå êîíè÷åñêèå ñïèíîñêàëèäû íåñêîëüêèõ ïåð- âûõ êðóãîâ, ñ ìåëêîçóá÷àòûì êðàåì ó òðåóãîëüíîãî îñíîâàíèÿ è öèëèíäðè÷åñêîé äèñòàëüíîé ÷àñòüþ; (3) óïëîùåííûå òðåóãîëüíûå ïðèäàòêè ñ ñòóïåí÷àòûìè è çàçóáðåííûìè êðàÿìè; (4) äåíòîñêàëèäû 2–3 çàäíèõ êðóãîâ, íåñóùèå êóòèêóëÿðèçîâàííûå ðàçäâîåííûå íàêîíå÷íèêè; (5) äåíòîñêàëèäû ñ îäíîçóáûì íàêîíå÷íèêîì, âñòðå÷àþùèåñÿ ó ìîëîäûõ ôîðì (ðèñ. 3.5; 3.14–3.15;

26 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.18. Ñêàëèäû âçðîñëîãî Halicryptus spinulosus: A — ñïèíîñêàëèäà;  — äåíòîñêàëèäà áåç òåðìèíàëüíûõ çóáöîâ; Ñ — äâóçóáàÿ äåíòîñêàëèäà; D — âåðøèíà ñïèíîñêàëèäû ñ òåðìèíàëüíûìè ïàïèëëàìè. Çâåçäî÷êàìè îáîçíà÷åíû òåðìè- íàëüíûå ïàïèëëû ñïèíîñêàëèäû. Ìàñøòàá: À–Ñ — 30 ìêì, D — 6 ìêì. Fig. 3.18. Scalids of adult Halicryptus spinulosus: A — spinoscalid; B — dentoscalid without terminal teeth; C — bifurcated dentoscalid; D — top of spinoscalid with terminal papillae. Asterisks indicate terminal papillae of spinoscalids. Bar: A–C — 30 mc, D — 6 mc. 3.18) (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996). Ó ìîëîäûõ îñîáåé èìåþòñÿ åùå äâóõ ïîñëåäóþùèõ çà äåí- òîñêàëèäàìè êðóãà ðóäèìåíòàðíûõ ïàïèëëîáðàçíûõ ñêàëèä. Äâà ïîñëåäíèõ êðóãà ó ìîëîäûõ Halicryptus îáðàçîâàíû ÷óâñòâèòåëüíûìè øèïàìè, íè÷åì íå îòëè÷àþùèìèñÿ îò ÷óâñòâèòåëüíûõ øèïîâ, áåñïîðÿäî÷íî ðàñïîëîæåííûõ ïî âñåìó òóëîâèùó. Ó ïðåäñòàâèòåëåé ñåìåéñòâà Priapulidae ïåðâûé êðóã ïðèäàòêîâ ïðåäñòàâëåí 8 öèðêóìîðàëü- íûìè ñêàëèäàìè, çà êîòîðûìè ñëåäóþò 25 ðÿäîâ ñêàëèä. Êàæäûé ðÿä ìîæåò ñîäåðæàòü íåñêîëüêî äåñÿòêîâ ïðèäàòêîâ (äî 50 ó ïðåäñòàâèòåëåé ðîäà Priapulus) è ïîäåëåí íà âûðàæåííûå ñåðèè (ðèñ. 3.16). Ñåðèÿ íà÷èíàåòñÿ êðóïíîé ñêàëèäîé, çà êîòîðîé ñëåäóþò íåñêîëüêî ïîñòåïåííî óìåíüøà- þùèõñÿ ïðèäàòêîâ. ×èñëî ñêàëèä â îäíîé ñåðèè ó ðàçíûõ âèäîâ ðàçëè÷àåòñÿ, ïðè ýòîì ïåðåäíèå ñåðèè âñåãäà ñîäåðæàò áîëüøåå ÷èñëî ñêàëèä, ÷åì çàäíèå. Òàê, ó Priapulus abyssorum ïåðåäíèå ñåðèè ñîäåðæàò îêîëî 10 ñêàëèä (Menzies, 1959). Ó Priapulopsis australis ïåðåäíèå ñåðèè ñîäåðæàò 3–4 ñêàëèäû, à çàäíèå ñîñòîÿò èç 2 èëè äàæå 1 ñêàëèäû (ðèñ. 3.2(8); 3.3; 3.17). Ó ïðåäñòàâèòåëåé ñåìåéñòâà Priapulidae îòìå÷åííûå âûøå çàêîíîìåðíîñòè ðàñïîëîæåíèÿ ðå- àëèçóþòñÿ íå íà óðîâíå îòäåëüíûõ ïðèäàòêîâ, à íà óðîâíå ñåðèé ñêàëèä, êîòîðûå òàêæå ðàñïîëà- ãàþòñÿ êðóãàìè ïî 9 (â 2+3n êðóãàõ) èëè ïî 8 (â 3+3n è 4+3n êðóãàõ) ñåðèé â êàæäîì (ðèñ. 3.1; 3.16–3.17). Îñîáåííî î÷åâèäíûì øàõìàòíîå ðàñïîëîæåíèå ñåðèé ñêàëèä ñòàíîâèòñÿ òàì, ãäå ñî- ñåäíèå ñåðèè â ðÿäó îòäåëåíû äðóã îò äðóãà áîëüøèìè ïðîìåæóòêàìè, êàê, íàïðèìåð, ó Priapulopsis australis (ðèñ. 3.2(8); 3.3; 3.17). Âñå ñêàëèäû Priapulus è Acanthopriapulus ïðåäñòàâëÿþò ñîáîé îäíîòèïíûå âûñîêîêîíè÷åñêèå ïðèäàòêè ñ òåðìèíàëüíûìè è ñóáòåðìèíàëüíûìè ìèêðîïàïèëëàìè (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996).

27 Àäðèàíîâ À.Â., Ìàëàõîâ Â.Â. Ãîëîâîõîáîòíûå ÷åðâè Ìèðîâîãî îêåàíà

Ó Priapulopsis öèðêóìîðàëüíûå ñêàëèäû è ñêàëèäû ïåðâûõ ñåðèé ãóñòî îïóøåíû êóòèêóëÿðíûìè âîëîñêàìè è íàïîìèíàþò öèðêóìîðàëüíûå ñêàëèäû Halicryptus (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996) (ðèñ. 3.17). Ñêàëèäû â ïåðåäíèõ ñåðèÿõ ó Priapulopsis australis è P. bicaudatus òåëåñêîïè÷åñêèå è ñîñòîÿò èç 2 ÷ëåíèêîâ (ðèñ. 3.17) (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996).  çàäíåé ÷àñòè èíòðîâåðòà ïðåä- ñòàâèòåëåé ñåìåéñòâà Priapulidae ñêàëèäû î÷åíü ìåëêèå (“çà÷àòî÷íûå”) è ïðåäñòàâëåíû áóãîð÷à- òûìè îáðàçîâàíèÿìè ñ íåñêîëüêèìè òåðìèíàëüíûìè ìèêðîïàïèëëàìè.  ïðîìåæóòêàõ ìåæäó ðÿ- äàìè è ñêàëèäàìè êóòèêóëà èíòðîâåðòà íåñåò ìèêðîðåëüåô â âèäå òîíêèõ êîëüöåâûõ ñêëàäîê, ìèêðîñêîïè÷åñêèõ òóìóëåé èëè òîíêîãî ñåò÷àòîãî ðèñóíêà (ðèñ. 3.16–3.17). Ñðåäè ìåéîáåíòîñíûõ ïðèàïóëèä íàèìåíåå äèôôåðåíöèðîâàíû ñêàëèäû êàðëèêîâûõ Tubiluchus, íî èõ ðàñïîëîæåíèå ïîä÷èíÿåòñÿ îáùèì äëÿ ïðèàïóëèä çàêîíîìåðíîñòÿì (ðèñ. 3.13) (Land, 1968; 1970; Àäðèàíîâ è äð., 1989; Àäðèàíîâ, Ìàëàõîâ, 1991; 1996). Ïåðâûé êðóã ñîäåðæèò 8 öèðêóìî- ðàëüíûõ ñêàëèä. Ñêàëèäû ïîñëåäóþùèõ êðóãîâ ÷åðåäóþòñÿ è êàæäûé êðóã ñ ïîðÿäêîâûì íîìåðîì 2+3n (ãäå n=0, 1, 2, 3...) ñîäåðæèò ïî 9 ñêàëèä, à êðóãè ïîä íîìåðàìè 3+3n è 4+3n ñîäåðæàò ïî 8 ñêàëèä. Âñåãî ó Tubiluchus íàñ÷èòûâàåòñÿ 23–24 êðóãà (ðèñ. 3.13), ñêàëèäû êîòîðûõ ñêëàäûâàþòñÿ â õàðàêòåðíûå äëÿ ïðèàïóëèä 25 ðÿäîâ. Ñêàëèäû Tubiluchus ïîñòðîåíû ïî îáùåìó ïëàíó è ïðåä- ñòàâëÿþò ñîáîé äâó÷ëåíèñòûå çàãíóòûå íàçàä ïðèäàòêè (ðèñ. 3.13). Äèñòàëüíûå ÷ëåíèêè íåñóò 1– 3 òåðìèíàëüíûõ “ñîñî÷êà” è ùåòèíêó. Ó Meiopriapulus ïåðâûé êðóã ñîäåðæèò 8 íå÷ëåíèñòûõ ñêàëèä, ïðè îñíîâàíèè êàæäîé èç êîòîðûõ ðàñïîëàãàåòñÿ ïàðà ôëîñêóëåé (ðèñ. 3.9). Âòîðîé êðóã ñîñòîèò èç 9 ÷ëåíèñòûõ ñêàëèä (2 ñêàëèäû â âåíòðàëüíîì ñåêòîðå è ïî îäíîé — â êàæäîì èç äðóãèõ). Òðåòèé êðóã ñîäåðæèò 8 ÷ëåíèñòûõ ñêàëèä (ðèñ. 3.9). Ïîçàäè íèõ ðàñïîëàãàåòñÿ øèðîêàÿ çîíà ìåëêèõ ëîêîìîòîðíûõ ñêàëèä, ïðåäñòàâëÿþùèõ ñîáîé ãðåáåí÷àòóþ ïëàñòèíêó ñ óäëèíåííûì ñêëåðîòèçîâàííûì ìàíóáðèóìîì (ðèñ. 3.9). Òî÷íîå ÷èñëî èõ îïðåäåëèòü çàòðóäíèòåëüíî. Ïî äàííûì Ìîðñ (Morse, 1981), Meiopriapulus fijiensis èìååò 150 ïðîäîëüíûõ ðÿäîâ ïî 8–9 ãðåáåí÷àòûõ ñêàëèä â êàæäîì. Òàêèì îáðàçîì, M. fijiensis — ýòî åäèíñòâåííàÿ ïðèàïóëèäà, ó êîòîðîé íå óäàåòñÿ âûäåëèòü 25 ïðîäîëüíûõ ðÿäîâ ñêàëèä. Íàèáîëüøàÿ ñòåïåíü â äèôôåðåíöèðîâêå ïðèäàòêîâ èíòðîâåðòà ó âçðîñëûõ ïðèàïóëèä èìååò ìåñòî ó Maccabeus (ðèñ. 3.8). 8 öèðêóìîðàëüíûõ ñêàëèä ïðåäñòàâëåíû çäåñü äëèííûìè ïåðèñòûìè ùóïàëüöàìè — òðèããåðíûìè øèïàìè (Por, 1973; Por, Bromley, 1974; Salvini-Plawen, 1974; Ìàëà- õîâ, 1978; Wolter, 1987). Çà íèìè ñëåäóåò êðîíà èç 25 åùå áîëåå äëèííûõ äâóâåòâèñòûõ ñêàëèä, òàêæå êàê è ïðåäûäóùèå, ïðèñïîñîáëåííûõ ê ôèëüòðàöèè ïèùè. Ñëåäóþùèå êðóãè ÷óâñòâèòåëü- íûõ, òðèôôèäíûõ è æåëåçèñòûõ ñêàëèä òàêæå ñîäåðæàò ïî 25 ïðèäàòêîâ (ðèñ. 3.8). Ïåðâûé èç íèõ ïðåäñòàâëåí êîðîòêèìè ÷óâñòâèòåëüíûìè ïàïèëëàìè, òîãäà êàê ïîñëåäóþùèå êðóãè ñîäåðæàò ñêàëèäû â âèäå èçîãíóòûõ êðþ÷üåâ è øèïîâ è âûïîëíÿþò ëîêîìîòîðíûå ôóíêöèè. Øåéíàÿ îáëàñòü ïðåäñòàâëÿåò ñîáîé êîðîòêóþ çîíó ìåæäó âîîðóæåííîé ñêàëèäàìè ÷àñòüþ õîáîòà è òóëîâèùåì. Ó âçðîñëûõ ìàêðîáåíòîñíûõ ïðèàïóëèä, à òàêæå ó Maccabeus, øåÿ ïðàêòè- ÷åñêè íå âûðàæåíà. Ó ìåéîáåíòîñíûõ Tubiluchus øåéíàÿ îáëàñòü õàðàêòåðèçóåòñÿ âûðàæåííûì ñóæåíèåì, îäåòà ñêëàä÷àòîé êóòèêóëîé è íåñåò îäèí èëè íåñêîëüêî êðóãîâ ìèêðîñêîïè÷åñêèõ ÷óâñòâèòåëüíûõ îáðàçîâàíèé — ôëîñêóëåé (ðèñ. 3.6A) (Land, 1968, 1970). Êîëüöåâûå ñêëàäêè ñîñòàâëåíû ìèêðî- ñêîïè÷åñêèìè êóòèêóëÿðíûìè áóãîðêàìè — òóìóëÿìè. Ó T. vanuatensis øåéíàÿ îáëàñòü íåñåò 6–7 êðóãîâ ïðÿìîóãîëüíûõ êóòèêóëÿðíûõ ïëàñòèíîê (Àäðèàíîâ, Ìàëàõîâ, 1991) (ðèñ. 3.13A). Ó Meiopriapulus ýòà îáëàñòü òàêæå íåñêîëüêî ñóæåíà, íåñåò êîëüöåâûå êóòèêóëÿðíûå ñêëàäêè è 7–9 êðóãîâ øèðîêèõ òðåóãîëüíûõ ñêàëèä (Morse, 1981). Òóëîâèùå ìàêðîáåíòîñíûõ ïðèàïóëèä ïîêðûòî ìíîãî÷èñëåííûìè êîëüöåâûìè ñêëàäêàìè — êóòèêóëÿðíûìè àííóëàìè (ðèñ. 3.1–3.2). Ó Halicryptus íàñ÷èòûâàåòñÿ äî 80–100 òîíêèõ àííóë, ãóñòî óñàæåííûõ òîíêèìè êóòèêóëÿðíûìè øèïàìè ñ âîëîñîâèäíûìè äèñòàëüíûìè ÷ëåíèêàìè (ðèñ. 3.2(9)). Ó ìîëîäûõ îñîáåé ïîìèìî øèïîâ èìåþòñÿ òóëîâèùíûå ïàïèëëû, íåñóùèå òóáóëè è/ èëè ôëîñêóëè. Àííóëû â ñåìåéñòâå Priapulidae áîëåå øèðîêèå, òàê ÷òî íà òóëîâèùå P. caudatus óêëàäûâàåòñÿ äî 50 àííóë (ðèñ. 3.1). Ó P. australis êîëüöåâûå àííóëû óñàæåíû ñôåðè÷åñêèìè òóìó- ëÿìè, ÷òî ïðèäàåò òóëîâèùó ñïåöèôè÷åñêèé êóòèêóëÿðíûé ðèñóíîê (ðèñ. 3.17E) (ñì. Àäðèàíîâ, Ìàëàõîâ, 1996). Àííóëû ýòèõ âèäîâ íåñóò õàîòè÷íî ðàñïîëîæåííûå òóëîâèùíûå ïàïèëëû — êî- íè÷åñêèå îáðàçîâàíèÿ, óñàæåííûå ðàçíîîáðàçíûìè æåëåçèñòî-÷óâñòâèòåëüíûìè ïðèäàòêàìè. Ó

28 Ãëàâà 3. Ïðèàïóëèäû

Ðèñ. 3.19. Ïðèäàòêè ïðåàíàëüíîãî ïîäîòäåëà ìîëîäîãî Priapulus caudatus: À — îáùèé âèä ïðåàíàëüíîãî ïîäîòäåëà;  — “áîðîäàâêè” è æåëåçèñòûå ïàïèëëû; Ñ — “áîðîäàâêà” ñ ìóòîâ÷àòîé ïàïèëëîé; D — “áîðîäàâêà”; Å — âåðøèíà ìóòîâ÷àòîé ïàïèëëû ñ òóáóëÿìè; F — æåëåçèñòàÿ ïàïèëëà. an — àíóñ; go — ãîíîïîð; pa — ïðåàíàëüíûå àííóëû; rp — êîëüöåâûå ïàïèëëû; sp — ñåêðåòîðíàÿ ïàïèëëà; ta — õâîñò; tp — ìóòîâ÷àòàÿ, èëè òóáóëÿðíàÿ, ïàïèëëà; tu — òóáóëè ìóòîâ÷àòîé ïàïèëëû; wa — “áîðîäàâêà”. Ìàñøòàá: À — 0,6 ìì;  — 80 ìêì; Ñ — 30 ìêì; D, F — 20 ìêì; Å — 10 ìêì. Fig. 3.19. Preanal subregion of young Priapulus caudatus: A — general view from posterior side; B — warts and glandular papillae; C — wart with pedicillate papilla; D — wart; E — top of pedicillate papilla with tubuli; F — glandular papilla. an — anal opening; go — gonopore; pa — preanal annuli; rp — ringpapillae; sp — glandular papilla; ta — tail; tp — pedicillate papilla; tu — tubuli of pedicillate papilla; wa — wart. Bar: A — 0.6 mm; B — 80 mc; C — 30 mc; D, F — 20 mc, E — 10 mc.

Priapulus, Priapulopsis è Acanthopriapulus â òóëîâèùíîì îòäåëå âûäåëÿåòñÿ ïðåàíàëüíûé ïîäîòäåë, îòäåëåííûé îò öåíòðàëüíîé îáëàñòè íàèáîëåå øèðîêîé è âûñòóïàþùåé àííóëîé èëè êîëüöîì èç ïàïèëë (“ringpapilla”) (ðèñ. 3.2). Ïîñëåäóþùèå ïðåàíàëüíûå àííóëû íåñóò íåñêîëüêî ìîäèôèöè- ðîâàííûõ ïàïèëë è áóãîð÷àòûõ “áîðîäàâîê” (“warts”) (ðèñ. 3.1). Ïîñëåäíèå ñíàáæåíû òåðìèíàëü- íûìè òðóáî÷êàìè, ïîðîâûìè ïîëÿìè è, ïî-âèäèìîìó, ÿâëÿþòñÿ æåëåçèñòî-÷óâñòâèòåëüíûìè îðãàíàìè. Íàèáîëåå ðàçíîîáðàçíû ýòè îðãàíû ó ìîëîäûõ îñîáåé (ðèñ. 3.19). Êóòèêóëà ïðåàíàëüíî-

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