Revision of the Glyceridae Grube 1850 (Annelida: Polychaeta)

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Language: English

This paper comprises the first revision of all worldwide described Glyceridae. Under the application of light and scanning electron microscopy most of the 172 species and subspecies of the glycerids are examined by their type material. Three of these species (†Glycera baltica (Eisenack 1939), †G. glaucopsammensis Charletta & Boyer 1974, †G. pilicae Szaniawski 1974) represent fossil taxa and are therefore not further investigated. The descriptions and drawings of three other taxa (Lumbricus capitarus O. Fabricius 1780, Nereis filigera Blainville 1825, Nereis (Nephtys) hirsuta Dalyell 1853), which have been referred to as glycerids, are clearly shown to belong to other families (Capitellidae, Cirratulidae, Nephtyidae). Glycera viridescens Stimpson 1854 seems to be a goniadid, whereas the taxon Pseudolacydonia caeca Rullier 1964 is a synonym for Glycerella magellanica (McIntosh 1885). Altogether, only 42 species (36 Glycera, 1 Glycerella and 5 Hemipodia) of the 166 recent taxa are considered to be valid species, the others being synonyms or nomina dubia and nomina nuda.

More than 4000 specimens from all over the world were studied and measured, and the 42 valid species are described and illustrated in detail. A list of all taxa together with their synonyms and an identification key for all Glyceridae known to this date are also presented. Addionally all available biological and ecological data from the literature are listed.

The results of the cladistic analysis support a sister group relationship between Goniadidae and Glyceridae. Both taxa are monophyletic as well as the three genera Glycera, Glycerella and Hemipodia within the glycerids.

Contents

Introduction 1
General characteristics 1
Taxonomic history 1
Biological notes 2
Distribution, ecology and feeding behaviour 2
Anatomy 4
Head appendages 4
Sensory elements 4
Jaws and venom glands 5
Coelom condition and osmoregulation 5
Digestive system and excretion 7
Reproduction and regeneration 7
Epitokous stages and their spawning 7
Larval development 8
Regenerative capacity 10
Parasitism and commensalism 10
Collecting, observing and culturing 10

Taxonomic features 11
Colour 11
Prostomium 11
Jaws 13
Chaetae 14
Parapodia 14
Branchiae 14
Segmental annotation 16
Ailerons 17
Proboscidial papillae 17

Materials and methods 20
Documentation of the examined material 20
Observations, drawings and measurements 20
Systematic section 20
Distribution charts 21
SEM and X-ray microanalysis 21
Cladistic analysis 21

List of original described glycerids 21

Systematic section 31
Diagnosis of the family Glyceridae GRUBE 1850 31
Key to the genera 31

Diagnosis of the genus Glycera SAVIGNY 1818 31
Key to the species of Glycera 32
Description of the valid species of the genus Glycera
Glycera capitata ØRSTED 1842 34
Glycera lapidum QUATREFAGES 1866 37
Glycera oxycephala EHLERS 1887 40
Glycera branchiopoda MOORE 1911 41
Glycera sphyrabrancha SCHMARDA 1861 42
Glycera guatemalensis BÖGGEMANN & FIEGE 2001 43
Glycera brevicirris GRUBE 1870 44
Glycera tesselata GRUBE 1863 47
Glycera benhami BÖGGEMANN & FIEGE 2001 48
Glycera russa GRUBE 1870 49
Glycera bassensis BÖGGEMANN & FIEGE 2001 50
Glycera celtica O'CONNOR 1987 51
Glycera robusta EHLERS 1868 52
Glycera pseudorobusta BÖGGEMANN & FIEGE 2001 53
Glycera dibranchiata EHLERS 1868 53
Glycera knoxi KIRKEGAARD 1995 54
Glycera macintoshi GRUBE 1877 55
Glycera nicobarica GRUBE 1868 57
Glycera madagascariensis BÖGGEMANN & FIEGE 2001 58
Glycera fallax QUATREFAGES 1850 58
Glycera sagittariae MCINTOSH 1885 60
Glycera unicornis SAVIGNY 1818 60
Glycera ovigera SCHMARDA 1861 62
Glycera pacifica KTNBERG 1865 64
Glycera americana LEIDY 1855 65
Glycera lamelliformis MCINTOSH 1885 66
Glycera onomichiensis IZUKA 1912 68
Glycera cinnamomea GRUBE 1874 69
Glycera gilbertae BÖGGEMANN & FIEGE 2001 70
Glycera posterobranchia HOAGLAND 1920 70
Glycera macrobranchia MOORE 1911 71
Glycera alba (O.F. MÜLLER 1776) 72
Glycera natalensis DAY 1957 73
Glycera africana ARWIDSSON 1899 74
Glycera tridactyla SCHMARDA 1861 75
Glycera prosobranchia BÖGGEMANN & FIEGE 2001 77

Diagnosis of the genus Glycerella ARWIDSSON 1899 78
Glycerella magellanica (MCINTOSH 1885) 78

Diagnosis Of the genus Hemipodia KINBERG 1865 79
Key to the species Of Hemipodia 79
Description Of the valid species Of the genus Hemipodia
Hemipodia simplex (GRUBE 1857) 79
Hemipodia yenourensis (IZUKA 1912) 81
Hemipodia californiensis (HARTMAN 1938) 82
Hemipodia pustatula (FRTEDRICH 1956) 83
Hemipodia armata (HARTMAN 1950) 84

Glyceridae incertae sedis and nomina nude 84
Non Glyceridae 87

Drawings, SEM pictures and distribution maps 89
Glycera capitata ØRSTED 1842 90
Glycera lapidum QUATREFAGES 1866 92
Glycera oxycephala EHLERS 1887 94
Glycera branchiopoda MOORE 1911 96
Glycera sphyrabrancha SCHMARDA 1861 98
Glycera guatemalensis BÖGGEMANN & FIEGE 2001 100
Glycera brevicirris GRUBE 1870 102
Glycera tesselata GRUBE 1863 104
Glycera benhami BÖGGEMANN & FIEGE 2001 106
Glycera russa GRUBE 1870 108
Glycera bassensis BÖGGEMANN & FIEGE 2001 110
Glycera celtica O'CONNOR 1987 112
Glycera robusta EHLERS 1868 114
Glycera pseudorobusta BÖGGEMANN & FIEGE 2001 116
Glycera dibranchiata EHLERS 1868 118
Glycera knoxi KIRKEGAARD 1995 120
Glycera macintoshi GRUBE 1877 122
Glycera nicobarica GRUBE 1868 124
Glycera madagascariensis BÖGGEMANN & FIEGE 2001 126
Glycera fallax QUATREFAGES 1850 128
Glycera sagittariae MCINTOSH 1885 130
Glycera unicornis SAVIGNY 1818 132
Glycera ovigera SCHMARDA 1861 134
Glycera pacifica KINBERG 1865 136
Glycera americana LEIDY 1855 138
Glycera lamelliformis MCINTOSH 1885 140
Glycera onomichiensis IZUKA 1912 142
Glycera cinnamomea GRUBE 1874 144
Glycera gilbertae BÖGGEMANN & FIEGE 2001 146
Glycera posterobranchia HOAGLAND 1920 148
Glycera macrobranchia MOORE 1911 150
Glycera alba (O.F. MÜLLER 1776) 152
Glycera natalensis DAY 1957 154
Glycera africana ARWIDSSON 1899 156
Glycera tridactyla SCHMARDA 1861 158
Glycera prosobranchia BÖGGEMANN & FIEGE 2001 160
Glycerella magellanica (MCINTOSH 1885) 162
Hemipodia simplex (GRUBE 1857) 164
Hemipodia yenourensis (IZUKA 1912) 166
Hemipodia californiensis (HARTMAN 1938) 168
Hemipodia pustatula (FRIEDRICH 1956) 170
Hemipodia armata (HARTMAN 1950) 172
Glyceridae incertae sedis and nomina nuda 173

Cladistic analysis 175
Selection of taxa 175
Characters 175
Coding 183
Analysis 183
Application of the cladistic analysis 183

Acknowledgements 186
References 187

Appendix 207
Additional registrations of Glyceridae from the literature 207
Abbreviations of the museums and institutions 245
Abbreviations in the text 245
Index 246

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