Головохоботные черви (Cephalorhyncha) Мирового Океана (Определитель морской фауны)

А.В. АДРИАНОВ, В.В. МАЛАХОВ
ГОЛОВОХОБОТНЫЕ ЧЕРВИ
(CEPHALORHYNCHA)
МИРОВОГО ОКЕАНА
(Определитель морской фауны)
A.V. ADRIANOV & V.V. MALAKHOV
CEPHALORHYNCHA
OF THE WORLD OCEAN
KMK Scientific Press Ltd.
Москва 



 1999

УДК 595.13:591.4:575.86
А.В. Адрианов, В.В. Малахов. Головохоботные черви (Cephalorhyncha) Мирового Океана (Определитель морской фауны). Москва: Товарищество научных изданий КМК. 1999. 328 стр.
A.V. Adrianov & V.V. Malakhov. Cephalorhyncha of the World Ocean. Moscow: KMK Scientific Press Ltd.
1999. 328 pp.
Монография представляет собой первую в мировой литературе полную сводку по морфологии
и систематике морских головохоботных червей 
— представителей типа Cephalorhyncha: приапулид, лорицифер, киноринх, морских нематоморф (волосатиков). Приведены подробные определительные таблицы как надвидовых таксонов, так и видов; для каждого таксона даны подробные
диагнозы; для каждого вида приведены подробные иллюстрации определительных признаков, экологические и биогеографические характеристики. 1 семейство, 1 род и 14 видов описаны как
новые для науки. Подробно описаны находки ряда ранее известных видов. Монография богато
иллюстрирована рисунками и микрофотографиями.
Для зоологов, гидробиологов и экологов, аспирантов и студентов.
Таб. 8, Ил. 193, Библ. 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
 А.В. Адрианов, В.В. Малахов, текст, иллюстрации
 К.Г. Михайлов, оригинал-макет
 KMK Scientific Press Ltd., издание

Лодейников склонился над листами,
И в этот миг привиделся ему
Огромный червь, железными зубами
Схвативший лист и прянувший во тьму.
Так вот она, гармония природы,
Так вот они, ночные голоса!
Так вот о чем шумят во мраке воды...!
Н. Заболоцкий, “Лодейников”
PRIAPULIDA: STRUCTURE, DEVELOPMENT,
PHYLOGENY, AND CLASSIFICATION
3
KMK LTD.
МОСКВА
1996

СОДЕРЖАНИЕ
1. ПРЕДИСЛОВИЕ ..............................................................................................................................12
2. ВВЕДЕНИЕ. План строения головохоботных червей ...................................................................13
3. ПРИАПУЛИДЫ................................................................................................................................14
3.1. Исторический обзор системы приапулид .............................................................................14
3.2. План строения и особенности морфологии, используемые в систематике ......................17
3.2.1. План строения, внешняя морфология и симметрия ..................................................17
3.2.2. Микроскопическая анатомия ........................................................................................32
3.2.3. Личинки приапулид ........................................................................................................32
3.3. Филогения приапулид .............................................................................................................40
3.4. Система, определительные таблицы и диагнозы таксонов .................................................40
Класс Priapulida............................................................................................................40
Ключ для определения надвидовых таксонов по взрослым формам ......................41
Ключ для определения личинок.................................................................................41
Отряд Priapulomorpha ............................................................................................42
Семейство Priapulidae ......................................................................................42
Род Priapulus ................................................................................................42
Род Acanthopriapulus ....................................................................................47
Род Priapulopsis ............................................................................................48
Семейство Tubiluchidae ....................................................................................49
Род Tubiluchus .............................................................................................49
Отряд Meiopriapulomorpha ....................................................................................52
Семейство Meiopriapulidae ..............................................................................52
Род Meiopriapulus.........................................................................................52
Отряд Halicryptomorpha .........................................................................................52
Семейство Halicryptidae ...................................................................................52
Род Halicryptus .............................................................................................52
Отряд Seticoronaria .................................................................................................53
Семейство Chaetostephanidae ..........................................................................53
Род Maccabeus ...................................................................................................53
3.5. Зоогеография ............................................................................................................................54
3.5.1. Макробентосные приапулиды .......................................................................................54
3.5.2. Мейобентосные приапулиды .........................................................................................57
3.5.3. Гипотеза гляциальных реликтов ....................................................................................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. ЛОРИЦИФЕРЫ ...............................................................................................................................67
4.1. История систематики и современное состояние системы ..................................................67
4.2. План строения и особенности морфологии, используемые в систематике ......................67
4.2.1. Внешняя морфология .....................................................................................................67
4.2.2. Микроскопическая анатомия ........................................................................................75
4.2.3. Развитие лорицифер .......................................................................................................75
4.3. Система, определительные таблицы и диагнозы таксонов .................................................76
Класс Loricifera ............................................................................................................76
Отряд Nanaloricida .................................................................................................76
Ключ для определения таксонов по взрослым формам.....................................76
Ключ для определения таксонов по личинкам...................................................77
Семейство Nanaloricidae ..................................................................................77
Род Nanaloricus ............................................................................................77
Семейство Pliciloricidae....................................................................................78
Род Pliciloricus .............................................................................................78
Род Rugiloricus .............................................................................................80
4.4. Зоогеография ............................................................................................................................81
4.5. Summary of chapter 4 ...............................................................................................................81
5. КИНОРИНХИ ..................................................................................................................................82
5.1. История систематики и современное состояние системы ..................................................82
5.2. План строения и особенности морфологии, используемые в систематике ......................85
5.2.1. Внешняя морфология .....................................................................................................85
5.2.2. Микроскопическая анатомия ........................................................................................87
5.2.2.1. Покровы тела ..........................................................................................................87
5.2.2.2. Мускулатура ............................................................................................................88
5.2.2.3. Нервная система .....................................................................................................90
5.2.2.4. Органы чувств .........................................................................................................91
5.2.2.5. Пищеварительная система ....................................................................................92
5.2.2.6. Полость тела............................................................................................................93
5.2.2.7. Выделительная система .........................................................................................93
5.2.2.8. Половая система .....................................................................................................93
5.2.3. Развитие и стадии жизненного цикла ..........................................................................94
5.3. Филогения киноринх ..............................................................................................................96
5.4. Система, определительные таблицы и диагнозы таксонов ...............................................104
Класс Kinorhyncha ..................................................................................................... 104
Ключ для определения надвидовых таксонов .........................................................104
Отряд Cyclorhagida ...............................................................................................105
Подотряд Protorhagae ...........................................................................................105
Семейство Zelinkaderidae ...............................................................................106
Род Zelinkaderes ......................................................................................... 106
Семейство Antigomonidae ..............................................................................107
Род Antigomonas ......................................................................................... 108
Подотряд Cryptorhagae......................................................................................... 108
Семейство Cateriidae ......................................................................................108
Род Cateria................................................................................................. 108
Подотряд Conchorhagae .......................................................................................109
Семейство Semnoderidae ................................................................................109
5

Род Semnoderes .......................................................................................... 109
Род Sphenoderes .........................................................................................111
Подотряд Cyclorhagae ..........................................................................................112
Семейство Centroderidae ................................................................................112
Род Centroderes .......................................................................................... 112
Род Condyloderes ........................................................................................114
Род Campyloderes .......................................................................................115
Семейство Echinoderidae ...............................................................................116
Род Echinoderes .......................................................................................... 116
Семейство Dracoderidae ................................................................................. 157
Род Dracoderes ........................................................................................... 158
Семейство Cephalorhynchidae........................................................................161
Род Cephalorhyncha ....................................................................................161
Отряд Homalorhagida ...........................................................................................164
Подотряд Homalorhagae.......................................................................................164
Семейство Pycnophyidae ................................................................................164
Род Pycnophyes...........................................................................................164
Род Kinorhynchus .......................................................................................211
Семейство Neocentrophyidae .........................................................................228
Род Neocentrophyes ....................................................................................228
Род Paracentrophyes ...................................................................................229
5.5. Зоогеография ..........................................................................................................................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. МОРСКИЕ НЕМАТОМОРФЫ ....................................................................................................306
6.1. История систематики и современное состояние системы ................................................306
6.2. План строения и особенности морфологии, используемые в систематике ....................307
6.3. Филогения волосатиков ........................................................................................................310
6.4. Система, определительные таблицы и диагнозы таксонов ...............................................311
Класс Nematomorpha ................................................................................................. 311
Подкласс Nectonematoidea ........................................................................................311
Отряд Nectonematida............................................................................................311
Семейство Nectonematidae ............................................................................311
Род Nectonema ...........................................................................................311
6.5. Зоогеография ..........................................................................................................................313
6.6. Summary of chapter 6 ............................................................................................................. 314
Class Nematomorpha ...................................................................................................314
Subclass Nectonematoidea ...........................................................................................314
Order Nectonematida ............................................................................................314
Family Nectonematidae ...................................................................................314
Genus Nectonema .......................................................................................314
7. ЛИТЕРАТУРА ................................................................................................................................. 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