Biodiversity and Zoogeography of the Polychaeta (Annelida) in
the deep Weddell Sea (Southern Ocean, Antarctica) and adjacent
deep-sea basins
Dissertation zur Erlangung des Grades eines Doktors der Naturwissenschaften der
Fakultät für Biologie und Biotechnologie der Ruhr-Universität Bochum
angefertigt im Lehrstuhl für
Evolutionsökologie und Biodiversität der Tiere
vorgelegt von
Myriam Schüller
aus
Aachen
Bochum 2007
Biodiversität und Zoogeographie der Polychaeta (Annelida) des
Weddell Meeres (Süd Ozean, Antarktis) und angrenzender
Tiefseebecken
The most exciting phrase to hear in science, the one that heralds new discoveries, is not
„EUREKA!“ (I found it!) but “THAT’S FUNNY…”
Isaak Asimov US science fiction novelist & scholar (1920-1992)
photo: Ampharetidae from the Southern Ocean, © S. Kaiser
Title photo: Polychaete samples from the expedition DIVA II, © Schüller & Brenke, 2005
Erklärung
Hiermit erkläre ich, dass ich die Arbeit selbständig verfasst und bei keiner anderen
Fakultät eingereicht und dass ich keine anderen als die angegebenen Hilfsmittel
verwendet habe. Es handelt sich bei der heute von mir eingereichten Dissertation um
fünf in Wort und Bild völlig übereinstimmende Exemplare.
Weiterhin erkläre ich, dass digitale Abbildungen nur die originalen Daten enthalten und
in keinem Fall inhaltsverändernde Bildbearbeitung vorgenommen wurde.
Bochum, den 28.06.2007
____________________________________
Myriam Schüller
INDEX
iv
Index 1 Introduction 1 1.1 The Southern Ocean 2
1.2 Introduction to the Polychaeta 4
1.2.1 Polychaete morphology 5
1.2.1.1 Ampharetidae MALMGREN 1866 7
1.2.1.2 Glyceridae GRUBE 1850 7
1.2.1.3 Goniadidae KINBERG 1866 8
1.2.1.4 Hesionidae GRUBE 1850 8
1.2.1.5 Nephtyidae GRUBE 1850 8
1.2.1.6 Nereididae JOHNSTON 1865 9
1.2.1.7 Opheliidae MALMGREN 1867 9
1.2.1.8 Sabellariidae JOHNSTON 1865 10
1.2.1.9 Scalibregmatidae MALMGREN 1867 10
1.2.1.10 Sphaerodoridae MALMGREN 1867 11
1.2.1.11 Syllidae GRUBE 1850 11
1.2.1.12 Terebellidae MALMGREN 1867 12
1.2.1.13 Trichobranchidae MALMGREN 1866 12
1.2.2 Polychaete ecology, reproduction and feeding strategies 13
1.2.2.1 Ampharetidae MALMGREN 1866 14
1.2.2.2 Glyceridae GRUBE 1850 15
1.2.2.3 Goniadidae KINBERG 1866 15
1.2.2.4 Hesionidae GRUBE 1850 16
1.2.2.5 Nephtyidae GRUBE 1850 16
1.2.2.6 Nereididae JOHNSTON 1865 16
1.2.2.7 Opheliidae MALMGREN 1867 17
1.2.2.8 Sabellariidae JOHNSTON 1865 17
1.2.2.9 Scalibregmatidae MALMGREN 1867 17
1.2.2.10 Sphaerodoridae MALMGREN 1867 18
1.2.2.11 Syllidae GRUBE 1850 18
1.2.2.12 Terebellidae MALMGREN 1867 19
INDEX
v
1.2.2.13 Trichobranchidae MALMGREN 1866 19
1.2.3 Polychaete systematics 19
2 Material and methods 21 2.1 The expeditions ANDEEP I, II, and III: sampling and on-board treatment 21
2.2 Taxonomic analyses 24
2.2.1 Final sorting, identification, and labeling of species 24
2.2.2 Description of new species 24
2.2.3 Construction of identification keys 25
2.3 Univariate and multivariate community analyses 25
2.3.1 Species accumulation plots 25
2.3.2 Standardization and transformation of sampling data 25
2.3.3 Univariate biodiversity measures 26
2.3.4 Similarity measures 27
2.3.5 Comparison of EBS epi- and supranets 27
2.3.6 Cluster analysis and MDS plotting 28
2.3.7 Environmental factors and species sets explaining
station similarities 29
2.3.7.1 BIO-ENV 29
2.3.7.2 BV Step 30
2.3.8 Average Taxonomic Distinctness (AvTD) and Variations
in Taxonomic Distinctness (VarTD) 31
2.4 Reconstruction of vertical and global distribution patterns 32
2.4.1 Vertical distribution patterns 32
2.4.2 Global distribution patterns 32
3 Results 33 3.1 Composition of polychaete communities 33
3.2 Identification keys to selected species found during ANDEEP I-III 34
3.2.1 Key to the Ampharetidae MALMGREN 1866 34
3.2.2 Glyceridae GRUBE 1850 41
3.2.3 Key to the Goniadidae KINBERG 1866 42
INDEX
vi
3.2.4 Key to the Hesionidae GRUBE 1850 43
3.2.5 Key to the Nephtyidae GRUBE 1850 45
3.2.6 Key to the Nereididae JOHNSTON 1865 46
3.2.7 Key to the Opheliidae MALMGREN 1867 47
3.2.8 Sabellariidae JOHNSTON 1865 51
3.2.9 Key to the Scalibregmatidae MALMGREN 1867 52
3.2.10 Key to the Sphaerodoridae MALMGREN 1867 56
3.2.11 Key to the Syllidae GRUBE 1850 58
3.2.12 Key to the Terebellidae MALMGREN 1867 64
3.2.13 Key to the Trichobranchidae MALMGREN 1866 71
3.3 Descriptions of new species 73
3.3.1 Ampharetidae MALMGREN 1866 73
3.3.1.1 Anobothrus pseudoampharete sp.n. 73
3.3.2 Hesionidae GRUBE 1850 76
3.3.2.1 Amphiduros serratus sp.n. 76
3.3.2.2 Micropodarke cylindripalpata sp.n. 77
3.3.2.3 Ophiodromus calligocervix sp.n. 78
3.3.2.4 Parasyllidea delicata sp.n. 79
3.3.3 Opheliidae MALMGREN 1867 83
3.3.3.1 Ammotrypanella MCINTOSH 1879 83
3.3.3.1.1 Ammotrypanella arctica MCINTOSH 1879 83
3.3.3.1.2 Ammotrypanella cirrosa sp.n. 84
3.3.3.1.3 Ammotrypanella mcintoshi sp.n. 86
3.3.3.1.4 Ammotrypanella princessa sp.n. 87
3.3.3.2 Ophelina ÖRSTED 1843 88
3.3.3.2.1 Ophelina ammotrypanella sp.n. 88
3.3.3.2.2 Ophelina robusta sp.n. 89
3.3.4 Scalibregmatidae MALMGREN 1867 93
3.3.4.1 Pseudoscalibregma papilia sp.n. 93
3.3.5 Sphaerodoridae MALMGREN 1867 94
3.3.5.1 Ephesiella hartmanae sp.n. 94
3.3.5.2 Sphaerodoropsis HARTMAN & FAUCHALD 1971 95
INDEX
vii
3.3.5.2.1 Sphaerodoropsis distincta sp.n. 95
3.3.5.2.2 Sphaerodoropsis maculata sp.n. 96
3.3.5.2.3 Sphaerodoropsis simplex sp.n. 98
3.4 Univariate and multivariate community analyses 101
3.4.1 Analysis of the epi- and supranet 101
3.4.2 Species richness and biodiversity 102
3.4.2.1 Species richness (Margalef’s Index) 103
3.4.2.2 Biodiversity and Evenness 105
3.4.3 Clustering and Multi Dimensional Scaling (MDS) 107
3.4.3.1 ANDEEP I-III 107
3.4.3.2 ANDEEP I/II 110
3.4.3.3 ANDEEP III 112
3.4.4 Correlation of polychaete communities to environmental
data (BIO-ENV) 113
3.4.5 Correlation of species to stations similarities (BV Step) 115
3.4.6 Average Taxonomic Distinctness (AvTD) and Variation
in Taxonomic Distinctness (VarTD) 115
3.5 Zoogeography 119
3.5.1 Vertical distribution patterns in the Southern Ocean 119
3.5.2 Global distribution patterns 119
4 Discussion 124 4.1 Efficiency of the gear and methods used for sampling 124
4.2 Polychaete abundance and family composition 126
4.3 Species identification, composition and descriptions of new species 129
4.3.1 Species composition of families 130
4.3.2 Descriptions of new species 131
4.3.2.1 Ampharetidae MALMGREN 1866 131
4.3.2.2 Hesionidae GRUBE 1850 132
4.3.2.3 Opheliidae MALMGREN 1867 133
4.3.2.4 Scalibregmatidae MALMGREN 1867 135
4.3.2.5 Sphaerodoridae MALMGREN 1867 136
INDEX
viii
4.4 Polychaete diversity in the Southern Ocean 139
4.5 Similarities between sampling areas 143
4.6 Influence of environmental factors and species on similarities 146
4.7 Taxonomic diversity 148
4.8 Zoogeography and vertical distribution 150
4.8.1 Origin of the Southern Ocean deep-sea fauna 150
4.8.2 Vertical distribution 151
4.8.3 Global distribution patterns 152
5 Summary 158 5.1 Zusammenfassung 160
6 References 162
7 Appendix 180 7.1 Figures 180
7.2 Tables 204
8 Acknowledges 245
9 Lebenslauf 247
INTRODUCTION 1
1 Introduction During the austral summers of 2002 and 2005 the expeditions ANDEEP I-III took place
to conduct one of the first thorough surveys concerning the faunal composition of the
deep Weddell and Scotia Seas. Transects starting south of South Africa, crossing the
Weddell Sea and the Drake Passage, and ending at the western coast of the Antarctic
Peninsula were sampled with several gear gathering faunal as well as geological data.
Since then specialists of different scientific backgrounds have analyzed the samples and
put together a complex puzzle of taxonomy, systematics, zoogeography, and ecology of
the epi- and infaunal communities in the deep Southern Ocean.
As part of this approach, the polychaetes sampled with an epibenthic sledge (EBS)
during ANDEEP I-III are focus of this study. Based on taxonomic analysis, a
characterization of the polychaete communities in the deep Southern Ocean and the
global zoogeography of the sampled species is achieved on a scale that is unique in its
outline to date. Due to the high number of individuals in the samples, an analysis on a
taxonomic level lower than family is only done for thirteen families, belonging to four
different orders, respectively suborders, which are all common representatives of the
deep-sea polychaete fauna world wide. These are the Ampharetidae, Sabellariidae,
Terebellidae, Trichobranchidae (Terebellida), Glyceridae, Goniadidae, Nephtyidae,
Sphaerodoridae (Glyceriformia), Hesionidae, Nereididae, Syllidae (Nereidiformia),
Opheliidae, and Scalibregmatidae (Opheliida). The study includes a detailed taxonomic
analysis of these families, analyses of their community structure, and descriptions of
new species. In combination with biodiversity measures and ecological analyses, this
approach will substantially contribute to our knowledge about the deep-sea benthos in
the Southern Ocean. In addition, the global distribution patterns of the polychaete
species sampled are reconstructed. The results are used to give insight into possible
ways of distribution of polychaetes in the Southern Ocean and adjacent deep-sea basins,
and contribute to the answers about the colonization history of the Southern Ocean deep
sea.
INTRODUCTION 2
1.1 The Southern Ocean
The Southern Ocean markedly differs from other oceans in that it lacks continental
borders. Rather, its water masses are isolated from the other world oceans by circulatory
systems and water fronts resulting in steep physical gradients at the transition zones.
The uniqueness of its oceanography (Dayton et al., 1994) and the continent of
Antarctica have been subject to extensive research during the last centuries. Antarctica
is permanently covered with ice. While the ice is restricted to the landmasses and
coastal waters during the austral summer, great parts of the Southern Ocean are covered
with thick masses of ice during the austral winter.
Constant low temperatures, the almost circular outline of the continent and prevailing
westerly winds lead to the formation of an unique ocean system. The air temperatures
around Antarctica seldomly rise above 2°C and water temperature near the continent lie
between -2° and -1°C throughout the year (Brey & Clarke, 1993; Knox & Lowry, 1977;
Pearse et al., 1991).
The Southern Ocean undergoes a steady eastwardly moving circumpolar current (White
& Peterson, 1996; Knox & Lowry, 1977) resulting from prevailing westerly winds
(West-Wind Drift) (Knox & Lowry, 1977) (Fig. 1). The Circumpolar Current has been
stable in position since its formation in the Cenozoic (Barker & Thomas, 2004) and is
today the strongest ocean current known (Barker & Thomas, 2004; Gnanadesikan &
Hallberg, 2000). Near the Antarctic continent east and south-east winds result in a
westerly water flow (East-Wind-Drift) that, when meeting the east coast of the Antarctic
Peninsula flows north to the Scotia Ridge and then eastwards across the South Atlantic
(Weddell-Sea-Drift) (Knox & Lowry, 1977). In addition, cold and dense surface water
flows northwards where it meets less dense southerly flowing subantarctic water. At
around 50°-60°S these two water masses collide and the denser Antarctic water sinks
below to form the Antarctic Convergence. The Antarctic Convergence is characterized
by steep gradients in salinity and temperature (Deacon, 1933, 1937; Mackintosh, 1946;
Moore et al., 1999). Close to the continent the sea ice formation results in high density
water that sinks below 4000 m and forms the Antarctic bottom water (Adkins et al.,
2002; Stössel & Kim, 1998). Meanwhile, deep water from higher latitudes flows south
INTRODUCTION 3
to rise to the surface at around 60°S (Antarctic Divergence) (Knox & Lowry, 1977;
Matsumoto et al., 2001).
Fig. 1: Current systems in the Southern Ocean (modified after Berkman, 1992)
While the cold temperatures and water currents lead to an isolation of the Southern
Ocean that is also partly reflected in the Antarctic shelf fauna (Brandt, 1992; Dayton et
al., 1994; Hilbig, 2004), the Antarctic deep sea shows ecological characters similar to
other deep-sea basins of the world. It is characterized by the total lack of light, very
slow bottom currents, and a constant temperature of about 0-0.5 °C (Fütterer et al.,
2003). Nevertheless some unusual characters can be found. Because of the heavy ice
cover of Antarctica the continent is pushed downwards. Therefore the shelf, though
narrow at most sites, reaches down to 600 m, even 800 m in the Ross Sea. This is about
twice as deep as in other oceans. Abyssal plains are first found below 3700 m (Knox &
Lowry, 1977). Still the ecological similarities between the Southern Ocean deep sea and
lower latitude deep seas imply that the deep-sea basins around Antarctica might not be
INTRODUCTION 4
as faunally isolated as the Antarctic shelf. Some invertebrate genera and species found
in the Southern Ocean have also been found in deep-sea basins world wide (e.g.,
Brandt, 1991; Hilbig, 2004).
1.2 Introduction to the Polychaeta
The polychaetes are one of the most common macroinfaunal invertebrate taxa in oceans
world wide. Aside from the benthos, polychaetes are also found in the interstitial, in the
water column (holopelagic forms and pelagic larvae), and as parasites. They occur in
shallow waters as well as in the deep sea. This unique variability and flexibility in life
strategies, together with their high abundance makes polychaetes one of the most
important invertebrate groups in marine biodiversity and monitoring studies.
The taxon Polychaeta is very large. Over 85 different families are distinguished to date
(Fauchald & Rouse, 1997), only a small percentage of them being well studied. The
different polychaete families are characterized by a high variability in body shape.
While some families can easily be classified to lower taxonomic levels (e.g.,
Scalibregmatidae, Sphaerodoridae), others show only a low number of distinct
characters (e.g., Cirratulidae, Maldanidae). In addition, many taxa are very speciose
(e.g., Spionidae). Thus species identifications can be problematic, sometimes resulting
in an insufficient resolution of the taxonomy of some taxa. As complex and confusing
as the taxonomy is the phylogeny. Many monospecific genera exist and species are
often shifted between genera and even families.The group of the polychaetes is too large
for a thorough molecular study of the whole taxon. Single attempts have been made to
date to resolve the relationships within single genera or families. A cladistic analysis
based on morphologic characters has been presented by Rouse and Fauchald (1997).
But still the dataset used for this study is not complete – some information on characters
for several families is missing – so that the result only gives a trend in polychaete
cladistics (Rouse & Fauchald, 1997).
INTRODUCTION 5
1.2.1 Polychaete morphology
Fauchald (1977), Hartmann-Schröder (1996), and Fauchald & Rouse (1997) give
excellent summaries of polychaete morphology and taxonomy. All subsequent
information about the morphology of polychaetes in general and selected families is,
unless stated otherwise, taken from these studies.
The polychaete worms follow the annelid “bauplan”. They are characterized by either
homonomous or heteronomous segmentation along the whole body. The presegmental
region consists of the prostomium and the peristomium. The prostomium can bear
several antennae, palps (which can also derive from the peristomium, e.g., buccal
tentacles), eyes, and nuchal organs (chemosensory organs). The peristomium, which, as
well as the prostomium, always lacks chaetae, often bears peristomial cirri. Ventrally,
between the prostomium and the peristomium, the mouth is located. Subsequent to the
presegmental region is the segmented trunk. Each segment usually carries parapodia and
chaetae (therefore called chaetiger) as well as segmentally arranged internal organs. The
first chaetigers often carry tentacular cirri arising from reduced parapodia. Those
segments can either be recognized as separate segments or are fused with each other and
sometimes the peristomium (cephalization) so that the original number is hard to
determine. The overall number of chaetigers usually differs within members of one
species. The terminal segment (postsegmental region), which is always achaetous, is the
pygidium with the anus and possible anal cirri. The pygidium includes the growth zone
where new segments derive.
The final body shape of each polychaete species is very variable. Many species look
highly complex due to the presence of more than one body region with differently
shaped chaetigers (heteronomous segmentation). Also the head region can be strongly
modified as adaptation to different life forms (e.g., life in tubes, in the sediment,
pelagic) and food sources.
The plesiomorphic parapodial form consists of two rami (biramous parapodium), the
neuropodium (ventral) and the notopodium (dorsal). The two rami can be similar or
distinctly different in size and shape. If one ramus is reduced in size (subbiramous
parapodium) or lacking (uniramous parapodium) it is always the notopodium. The
parapodia are used for crawling and swimming. In burrowing and tube dwelling forms
INTRODUCTION 6
they are often reduced to rudiments that do not hinder the animal’s movement through
the sediment, or function as anchors in tubes. Each neuro-/ notopodium is classically
characterized by the presence of a neuro-/ notopodial lobe that carries the chaetae and a
ventral/ dorsal cirrus. The lobes are often supported by one or more internal strong
chaetae, the aciculae. If lobes are formed as welts or ridges without aciculae they are
often regarded as tori. Most tori are characterized by the presence of hooks or unicini. In
addition to the parapodial lobes, pre- or postchaetal lobes and additional ligules, as well
as variably shaped branchiae, can be present. Ventral and dorsal cirri are very variable
in shape in different taxa and on different regions of the body. They can also be lacking.
An important character in polychaete taxonomy is the shape of the chaetae. They can be
simple or compound, smooth or serrated, falcigerous or spinigerous, internally with or
without structure, capillaries or spines, and much more. Some chaetae are so strongly
modified that they are hardly recognized as such, e.g., uncini (chaetae modified to
hooks) or palae in the Terebellomorpha. Within one individual, several different chaetae
types can occur, either in different body regions or within one parapodium. The
presence or absence of chaetae can be an important taxonomic character as well.
The polychaeta operate skin respiration. In order to enlarge the respiratory surface,
branchiae can be present. These are often found dorsally on the anterior body region or
attached to the parapodia.
The internal structure of the Polychaeta is characterized by the presence of longitudinal,
ring and parapodial muscles, a rope-ladder nervous system, a more or less closed
circulatory system, a gut consisting of pharynx (often eversible and armed with jaws)
and sometimes proventricle, middle gut and end gut, segmental proto- or metanephridia,
and gonads. Reductions and variations in structure and number of these features are
common.
In the following short summaries of the “bauplan” for the thirteen families focused on
in this study are given. As for the general polychaete morphology, the information is
mainly taken from Fauchald (1977), Hartmann-Schröder (1996), and Fauchald & Rouse
(1997), additional citations are marked as such. The summaries contribute to the
understanding of the taxonomic classification of the species sampled and the
descriptions of new species.
INTRODUCTION 7
1.2.1.1 Ampharetidae MALMGREN 1866
The prostomium of the Ampharetidae is rather small but variable in shape, the
peristomium is limited to the lips and mouth. Antennae are missing, palps are present as
well developed buccal tentacles. These are retractable. They are everted through an
eversion of the lip-like structure on which they are located. Nuchal organs are small and
indistinct. The first and second segments are fused to the head and achaetous. The third
segment can bear large, robust chaetae, called paleae (Day, 1964, Holthe, 1986). The
segmented trunk consists of two regions, the thorax and the abdomen. The parapodia of
the thorax bear chaetae in the notopodia and uncini (chaetae modified to hooks) in the
neuropodia. The notopodia of the abdomen are reduced, notopodial lobes can still be
present. Dorsal and ventral cirri are lacking throughout. The pygidium sometimes bears
abdominal cirri.
An important taxonomic character of the Ampharetidae is the number and arrangement
of the branchiae on the anterior segments. Up to four pairs can be present that are
arranged in transverse rows across the dorsum of the anterior segments. Additionally,
the number of thoracic setigers is species specific.
1.2.1.2 Glyceridae GRUBE 1850
The prostomium is ringed externally, strongly prolonged and tapering, and bears a pair
of antennae and palps at its tip. The peristomium is limited to the lips. The glycerids are
characterized by homonomous segmentation with well developed parapodia. The
parapodia are either biramous or uniramous, often with highly differentiated post- and
prechaetal lobes. Dorsal and ventral cirri, as well as one pair of anal cirri are present.
Branchiae can be present.
The eversible pharynx bears a multitude of papillae of one or two kinds and is tipped by
four jaws supported by ailerons.
INTRODUCTION 8
1.2.1.3 Goniadidae KINBERG 1866
The prostomium is externally ringed and tapers to a blunt tip with two antennae and
palps. The peristomium is limited to the lips. The anterior parapodia of the
homonomous trunk consist of well developed neuropodia, the notopodia are reduced to
the dorsal cirri. In median and posterior parapodia both rami are of similar development.
One pair of anal cirri is present, branchiae are lacking.
The pharynx always bears various pharyngeal organs. These can either be of few kinds
or characteristically differentiated in variety and size. The arrangement of the
pharyngeal papillae is considered a species character. At the tip of the pharynx a circlet
of two macrognaths and two arcs of micrognaths are found forming a complex jaw
apparatus (Hartman, 1950).
1.2.1.4 Hesionidae GRUBE 1850
The prostomium is well developed and bears 2-3 antennae and usually one pair of
ventral palps. Large eyes, consisting of a disinct lense are often present as well as
apparent nuchal organs on the posterior margin of the prostomium. The peristomium is
limited to the lips. The hesionids are homonomously segmented, several anterior
segments are fused and cephalized. The dorsal and ventral cirri of these segments are
prolonged to form up to eight pairs of tentacular cirri. Chaetae are usually missing in
these segments. The following segments are uniramous or biramous, subbiramous
parapodia are also reported. Ventral and dorsal cirri are usually well developed. While
the notochaetae are simple throughout, the neurochaetae are composite. The pygidium
bears one pair of anal cirri. Jaws may be present, branchiae are always lacking.
1.2.1.5 Nephtyidae GRUBE 1850
The prostomium is quadrangular or pentagonal with one pair of lateral antennae and one
pair of ventrolateral palps. The peristomium is limited to the lips. Nuchal organs are
INTRODUCTION 9
present, often indistinct. The first segment is smaller than the subsequent with smaller
parapodia. One or two pairs of tentacular cirri are present. All parapodia are biramous,
well developed, with dorsal and ventral cirri and complex pre- and postchaetal lobes.
The two rami are often widely separated from each other, resulting in a square cross
section that is characteristical for the Nephtyidae. Ventrally on the notopodia branchiae
are present that reach into the space between the noto- and neuropodia, curled inward
(e.g., Aglaophamus KINBERG 1866) or outward (e.g., Nephtys CUVIER 1817) The
chaetae are simple, but characteristically ornamented. One single anal cirrus is present
medially on the pygidium. The pharynx is armed with buccal papillae and one pair of
lateral jaws, the papillae of which are of major importance in taxonomic studies.
1.2.1.6 Nereididae JOHNSTON 1865
The prostomium is inversely T-shaped, and in small species often diamond shaped. It
bears a pair of frontal antennae and a pair of usually large articulated palps. The
peristomium is limited to the lips. Nuchal organs are present. The first indistinct
segment carries tentactular cirri (usually four pairs, sometimes only two or three). The
biramous parapodia are well developed with both dorsal and ventral cirri. The notopodia
additionally have flattened ligules. The chaetae are exclusively composite. For Hediste
MALMGREN 1867 heavily fused chaetae are reported that appear simple at first view
(Breton et al., 2003). The pygidium bears a pair of anal cirri. Branchiae are absent. The
pharynx is armed with prominent lateral jaws. Terminal papillae are lacking while the
outer surface of the pharynx is covered with papillae. Additionally, the presence and
shape of paragnaths is an important distinguishing character for the Nereididae.
1.2.1.7 Opheliidae MALMGREN 1867
Opheliidae are fusi-form polychaetes, the body might be prolonged with a ventral
groove. The prostomium is usually conical and sometimes carries a papilliform distal
palpode. Antennae and palps are absent. The peristomium is limited to the lips. A pair
INTRODUCTION 10
of distinct eversible nuchal organs is present laterally on the prostomium. The mouth is
present as a transverse slit ventrally on the first chaetiger rather than between the
prostomium and the peristomium. All chaetigers are similar in shape, the parapodia are
biramous but reduced in size. The chaetae are exclusively capillaries with various
ornamentations. Branchiae, when present, are associated with the upper end of the
parapodia. They consist of single filaments only. Dorsal and ventral cirri are lacking.
The pygidium bears multiple cirri, or may be hood-shaped with internal or marginal
cirri. The presence of anal tubes is common. The pharynx is unarmed.
1.2.1.8 Sabellariidae JOHNSTON 1865
The prostomium is fused to the peristomium and often only visible as a median keel.
The peristomium is visible as lips, usually covered by the first two chaetigers (the first
of which is fused to the head). The chaetae of these chaetigers are modified to form an
operculum. Antennae are missing while palps and nuchal organs are present. The
notopodia are short and cylindrical, the neuropodia are tori. Dorsal and ventral cirri are
absent, flattened branchiae are present dorsally. The Sabellariidae are characterized by
chaetal inversion, abdominal uncini are notopodial while thoracic ones are neuropodial.
Chaetae types are variously ornamented capillaries, spines and uncini.
1.2.1.9 Scalibregmatidae MALMGREN 1867
The prostomium of the Scalibregmatidae is truncate or t-shaped. Antennae and palps are
absent. The peristomium is well developed, forming a ring around the prostomium.
Anterior segments might be characteristically expanded, or the body is maggot-like. The
parapodia are biramous, fully developed with rather short rami. Dorsal and ventral cirri
are reported for some species (Blake, 1981; Schüller & Hilbig, 2007). The presence of
branchiae is a common trait. The chaetae observed are capillaries, furcate chaetae and
sometimes anterior acicular spines.
INTRODUCTION 11
1.2.1.10 Sphaerodoridae MALMGREN 1867
The prostomium is either free or fused to peristomium. The peristomium is limited to
the lips. Paired lateral antennae and a median antenna are present in addition to a ventral
pair of palps. Nuchal organs are also observed. The first segment of the homonomously
segmented trunk is indistinct. The uniramous parapodia are well developed with distinct
neuropodia and ventral cirri. Branchiae are absent. The dorsum of the body is densely
covered by small (microtubercles) and numerous rows of large tubercles
(macrotubercle). The most lateral rows of these marcotubercles are assumed to be
modified dorsal cirri. The tubercles can be sessil or with a short stem; their number and
shape are strong taxonomic characters. The chaetae are either compound falcigers or
ornamented capillaries. Some taxa bear anterior spines.
1.2.1.11 Syllidae GRUBE 1850
The prostomium is truncate bearing three antennae and a pair of unarticulated, more or
less fused palps. The peristomium is limited to the lips. Nuchal organs are present. The
first segment differs from the following by lacking chaetae, two pairs of tentacular cirri
are present instead. The parapodia are usually biramous, the notopodia being less
developed than the neuropodia. Dorsal and ventral cirri are present in most subfamilies,
the Autolytinae lack ventral cirri. The cirri vary in length and shape, they are either
smooth or articulated. One pair of anal cirri is present. Branchiae are absent. The
chaetae are composite with variously structured appendages. In addition, one or two
simple chaetae can be present per parapodium. The anterior digestive tract is equipped
with a muscular proventricle that is unique for this family in its form. The cells of the
proventricle are arranged in distinct rows whose approximal number is fixed for each
species. The pharynx is often armed with one or more teeth.
INTRODUCTION 12
1.2.1.12 Terebellidae MALMGREN 1867
The prostomium, peristomium and anterior segments are fused. The peristomium and
the anterior segments form an extended upper lip. Antennae are lacking and numerous
grooved tentacular palps emerge at the margin between pro- and peristomium. These
palps are not retractable into the mouth, in contrast to that of the Ampharetidae. Nuchal
organs can be present or absent. The body is separated into two regions. The thorax is
characterized by biramous parapodia of which the notopodia carry ornamented
capillaries, the neuropodia uncini. In the abdominal segments the notopodia are strongly
reduced. Tentacular, dorsal and ventral cirri are absent. Branchiae of various types can
be present dorsally in anterior chaetigers.
1.2.1.13 Trichobranchidae MALMGREN 1866
The pro- and peristomium are fused, the peristomium forms an extended lip similar to
that of the Terebellidae. Antennae are absent, palps are present as multiple buccal
tentacles. These are grooved and originate from the prostomial edge. Nuchal organs
have been observed in some taxa. The first segment is fused to the head and achaetous.
The following thoracic segments have biramous parapodia with chaetae carrying
notopodia and uncinal neuropodia. One or several of the anterior segments can bear
neuropodial bent spines. In abdominal segments, the notopodia are strongly reduced.
While the thoracic uncini are long-handled, that of the abdomen are short-handled.
Ventral, dorsal and anal cirri are lacking. Branchiae are either present as two to three
groups of single filaments or as a single branchia which may be cirriform or consist of
four lamellate lobes dorsally on anterior chaetigers.
INTRODUCTION 13
1.2.2 Polychaete ecology, reproduction and feeding strategies
As mentioned above the Polychaeta have occupied most ecological niches found in
marine environments (Ushakov, 1974). Most species are benthic forms (epi- or infauna)
but also pelagic (e.g., Alciopidae, Tomopteridae) (Ushakov, 1974; Fauchald, 1977),
parasitic (e.g., some Eunicidae) (Clark, 1956; Pettibone, 1957), and commensalist forms
(e.g., some Syllidae, Nereididae, or Polynoidae associated with clams, sponges, star
fish, or tunicates) (Hartman, 1936; Licher, 1999; Rosbaczylo & Canete, 1993) are
known. As benthic samples are the base for the presented study the focus is laid upon
benthic forms in the following.
Benthic polychaetes have evolved in high variability. They cover a size range from few
micrometers (e.g., some Syllidae) to several centimeters (e.g., some Ampharetidae) or
even meters (e.g., some Eunicidae) in length. As variable as form and size are the
ecological niches they occupy. E.g., interstitial forms, burrowers (e.g., Opheliidae,
Scalibregmatidae, Capitellidae), tube dwellers (e.g., Ampharetidae, Sabellariidae, many
Terebellidae and Trichobranchidae), and epibenthic vagile forms (e.g., Eunicidae,
Glyceridae, Hesionidae, Nereididae, Phyllodocidae) are observed. This flexibility is
obviously not achieved by variability in form and size alone, but also by adaptations in
physiology, such as reproduction and feeding strategies.
Feeding strategies range from omnivorous and carnivorous hunters, selective and non-
selective deposit feeders, to suspension feeders. Even in chemoautotrophic
environments such as hydrothermal vents, cold seeps, and whale falls, polychaetes play
a dominant role. Many forms have been found living in symbiosis with chemoautotroph
bacteria (e.g., Alvinella pompejana DESBRUYÈRES AND LAUBIER 1980 living in
symbiosis with Proteobacteria spec. (Campbell et al., 2001)).
The reproduction in polychaetes is also widely variable. Though most polychaetes have
separate sexes, parthenogenesis and hermaphroditism have been reported in some
species (e.g., the sphaerodorid species Ephesiella mixta (FAUCHALD 1974) (Ushakov,
1974)). Many species can additionally switch between generative and vegetative
reproduction. During vegetative reproduction the polychaetes abandon certain parts of
their body and replace those lost segments by regeneration of new segments.
Fragmentation and collateral budding are also very common. Vegetative reproduction
INTRODUCTION 14
enables the polychaetes to react fast to changes in environment (e.g., sudden food
input).
Generative reproduction can occur in various ways, too. The most common way is that
the female releases the eggs into the water column (either through the nephridium or by
rupture of the body wall) where they are fertilised by the sperms of the male. A second
way is that only one part of the body becomes fertile (epitokous) and is released from
the otherwise sterile (atokous) polychaete (schizogamy). The epitokous parts of both
sexes then ascend to the water surface, mate and ‘die’ afterwards while the atokous
parts live on as before. Sometimes the whole animals becomes epitokous (epigamy). A
well known family for epitoky are the Syllidae. They are characterized by the formation
of numerous fertile epitokous parts in the posterior derivation zone, the so called stolons
(e.g., Rouse & Pleijel, 2006). The separation of atokous infertile stages and epitokous
fertile stages is consequently limited to free living forms, it is not reported for tube-
dwellers.
Originally polychaetes have pelagic larvae (trochophorae). These are either
planktotrophic or lecithotrophic. For some species both larval forms have been reported
(e.g., Tharyx marioni (ST. JOSEPH 1894) (Cazaux, 1972; Dales, 1951; Gibbs, 1971) or
Nereis pelagica LINNÉ 1761 (Herpin, 1925; Wilson, 1932)). Vivipary (Ushakov, 1974),
external gestation (e.g., some Syllidae (Licher, 1999)) and brooding have also been
observed.
Although a general idea of the variability in reproduction and feeding of the Polychaeta
exists little is known about actual strategies of the different families. In the following a
short overview of the knowledge to date concerning the families treated in detail in this
study is given.
1.2.2.1 Ampharetidae MALMGREN 1866
The Ampharetidae are tube-building suspension feeders. They are often rather large
forms (few millimeters up to several centimeters) that are common on soft bottom and
hard substrates of all depths, especially in the deep sea (Cosson-Saradin et al., 1998;
INTRODUCTION 15
Fauchald, 1977; Hartman, 1966; 1967; 1978; Hilbig, 2004). Forms living in symbiosis
with chemoautotrophic bacteria have been observed.
Ampharetidae have pelagic larvae (e.g., Melinna cristata (SARS 1851)), free-swimming
juveniles (e.g., Ampharete grubei MALMGREN 1865) or are active brooders (such as
Hobsonia florida (HARTMAN 1951)) (Wilson, 1991). Their reproduction is mainly
generative.
1.2.2.2 Glyceridae GRUBE 1850
The Glyceridae are vagile forms that live on and in soft, sandy and muddy substrate.
They are suspectively carnivorous hunters (Fauchald, 1977). Their well developed jaw
apparatus enables them to feed on smaller invertebrates.
Little is known about their reproduction. It is probably generative with pelagic (species
of Glycera SAVIGNY 1818) or non-swimming larvae. Also epitoky is reported for
species of Glycera with species bearing prolonged epitokous setae (Arwidsson, 1899;
Ehlers, 1868; Fage & Legendre, 1927; Hartman, 1950; Wilson, 1991).
1.2.2.3 Goniadidae KINBERG 1866
The feeding strategies of the Goniadidae are still unresolved. While species of this
family have been regarded carnivorous predators (Ehlers, 1868; McIntosh, 1910), Stolte
(1932) suspected them to be detritus feeders. Their reproductive strategies are very
variable. Epitoky is reported (Hartman, 1950). The presence of epitokous setae is thus
still questionable. They are not reported in Ophioglycera VERRILL 1885 but might be
present in some species of Goniada AUDOUIN & EDWARDS 1834 (Hartman, 1950).
Glycinde armigera MOORE 1911 has lecithotrophic larvae, Goniada emerita AUDOUIN
& EDWARDS 1834 planktotrophic (Blake, 1975; Cazaux, 1972; Wilson, 1991).
INTRODUCTION 16
1.2.2.4 Hesionidae GRUBE 1850
The Hesionidae are of medium size, usually around a few millimeters long. They are
mainly epibenthic and vagile. Most species prefer primary and secondary hard
substrates (Fauchald, 1977), but also several pelagic or commensalist forms are known.
The presence of papillae and small chitinized jaws in some species suggests omnivorous
feeding. Their reproduction strategies include free-swimming planktotrophic and
lecitotrophic larvae (such as species of Gyptis MARION & BOBRETZKY 1875 and
Ophiodromus SARS 1862), free-swimming juveniles (species of Hesionides FRIEDRICH
1937), and encapsulation of planktotrophic larvae and juveniles in gelatinous capsules
(Microphthalmus MECZNIKOW 1865) (Blake, 1975; Haaland & Schram, 1983;
Treadwell, 1898; Westheide, 1967; 1970; Wilson, 1991).
1.2.2.5 Nephtyidae GRUBE 1850
The Nephtyidae are vagile forms of the benthic community. They prefer soft or sandy
substrates that they can crawl on or burrow in. They mainly occur in shallower depths;
deep-sea species have also been reported (Hartman, 1950). Their feeding strategy is
propably similar to that of the Hesionidae since only some horny papillae are present
instead of jaws.
Their reproduction strategies include epitoky. Prolonged epitokous chaetae and enlarged
parapodial lobes are reported for some species (Augener, 1912; Fage & Legendre,
1927). The larval stages are mostly planktotrophic (Hartman, 1950; Wilson, 1991).
1.2.2.6 Nereididae JOHNSTON 1865
The Nereididae are easily the best known polychaetes. They are most common in
marine environments of all depths but also freshwater penetration is reported (Ushakov,
1974; Fauchald, 1977). Their size ranges from few millimeters to several centimeters.
As omnivorous vagile forms they feed on algae and hunt small invertebrates.
INTRODUCTION 17
Their reproduction is mainly generative with pelagic larvae. These can be
planktotrophic (e.g., Nereis grubei (KINBERG 1866), Perinereis cultrifera (GRUBE
1840)) or lecithotrophic (e.g., Nereis diversicolor O. F. MÜLLER 1776, Platynereis
bicanaliculata (BAIRD 1863)) (Blake, 1975; Cazaux, 1969; Dales, 1950; Reish, 1954;
1957). Free-swimming juvelines and brooders with tubes and direct development are
also known (Wilson, 1991). For some species several reproduction strategies are
reported (e.g., Platynereis dumerilii (AUDOUIN & MILNE EDWARDS 1833), Nereis
pelagica LINNÉ 1761) (Wilson, 1991). In addition, epitoky is common for the
Nereididae, the epitokous forms are then called Heteronereis (e.g., Clark, 1961; Rouse
& Pleijel, 2006).
1.2.2.7 Opheliidae MALMGREN 1867
The Ophelilidae are burrowing deposit feeders. They are very common in sandy and
muddy bottoms of all depths (Fauchald, 1977). During reproduction pelagic larvae are
produced that are either planktotrophic or lecithotrophic (Wilson, 1991). For Ophelina
bicornis SAVIGNY 1818 both variants are reported (Riser, 1987; Wilson, 1948).
1.2.2.8 Sabellariidae JOHNSTON 1865
The body of the Sabellariidae is completely adapted to life in a tube (Fauchald, 1977).
They are the only known reef-builders (Schäfer, 1972) among polychaetes. Exclusively
plantotrophic larvae are known to date. Reproductive strategies are, however, only
reported for few genera (Rouse & Pleijel, 2006; Wilson, 1991).
1.2.2.9 Scalibregmatidae MALMGREN 1867
Their similarity in body shape to the Opheliidae suggests similar feeding and
reproduction strategies. The Scalibregmatidae are deposit feeders that burrow in the
INTRODUCTION 18
sediment. Studies on their reproduction strategies are lacking, information about their
larval stages is not available.
1.2.2.10 Sphaerodoridae MALMGREN 1867
The Sphaerodoridae are small, vagile hunters, probably omnivorous. They are
exclusively epibenthic and are most common on sandy and muddy bottoms in deep
waters. Some records from shallow hard bottoms exist. Generative reproduction with
demersal larvae seems to be the most common reproduction strategy, but
hermaphroditism and the occurance of lecithotrophic, suprabenthic larvae are suggested
in some species (Fauchald, 1974; 1977).
1.2.2.11 Syllidae GRUBE 1850
Similar to the Sphaerodoridae the Syllidae are also relatively small, fragile, and vagile
polychaetes. Their food supposedly consists of small algae and protozoans, although
larger forms might be able to feed on meiobenthic invertebrates. They occupy a great
variety of ecological niches including commensalism with sponges and other
invertebrates offering cavern-like structures, and parasitism (Licher, 1999), but an
interstitial and benthic way of living in waters of all depths is but the most common.
The Syllidae are known for a huge flexibility in reproductive strategies (Wilson, 1991).
Besides generative reproduction (schizogamy, and epigamy with stolonization), the
reproduction by collateral budding is reported. Brooding, vivipary and external
gestation are very common for the Syllidae (e.g, Exogone OERSTED 1845, Sphaerosyllis
CLAPAREDE 1863) (Cazaux, 1972; Westheide, 1974). In addition, cases of
hermaphroditism are reported (Licher, 1999).
INTRODUCTION 19
1.2.2.12 Terebellidae MALMGREN 1867
The Terebellidae are tube-dwelling suspension feeders found in all environments
(Fauchald, 1977). They are not completely restricted to living in their tubes and can
leave them occasionally. The reproduction strategies of the Terebellidae are mainly
generative reproduction with lecitotrophic pelagic larvae (e.g., Artacama proboscidea
MALMGREN 1866 (Thorson, 1946)) or brooding. Direct development is also very
common (Wilson, 1991).
1.2.2.13 Trichobranchidae MALMGREN 1866
The Trichobranchidae are very similar to the Terebellidae and Ampharetidae. It is
therefore proposed that their ecology, including feeding and reproduction strategies, are
similar. Most species are represented in cold-water soft bottoms, especially in greater
depths (Fauchald, 1977). Terebellides stroemi SARS 835 is reported to have gelatinous
capsules and direct development (Thorson, 1946).
1.2.3 Polychaete systematics
Systematics in polychaetes is still unresolved and problematic. There is still controversy
as to whether the taxon Polychaeta is monophyletic or paraphyletic within the Annelida
(Kojima, 1998; Mc Hugh, 1997; Rouse & Fauchald, 1995; Westheide et al., 1999).
There is evidence from molecular studies that the polychaetes are paraphyletic and that
the Echiura, Pogonophora and Clitellata nest among them (Bleidorn et al., 2003a;
2003b; Jördens et al., 2004; Struck et al., 2002). In addition, the monophyly of the
Annelida is in question to some extent. Westheide et al. (1999) give the most recent
summary about the different approaches to solve the systematics of the Annelida and
the position of the polychaetes.
To date it is unknown what the basal and most ancient annelid looks like. Knowledge of
this stem species would strongly benefit to determining the systematics of the
INTRODUCTION 20
Polychaeta in general (Rouse & Pleijel, 2003; Westheide et al. 1999). One hypothesis is
that the most basal forms are simple-bodied taxa. This would lead to the assumption that
the ancient annelids were mud-dwelling burrowers (Clark, 1964; 1969; Fauchald, 1974;
Kojima, 1998) feeding on sediment. A second hypothesis expresses the idea of an
epifaunal annelid with homonomous segmentation and biramous parapodia as the stem
species (Conway Morris & Peel, 1995; Mc Hugh, 1997; Westheide, 1997).
Within the Polychaeta, the position of the separate families is also largely unknown.
This is due to the great size of this taxon (a particular problem for molecular studies),
and also the fact that many families are only poorly studied. Many characters are not
determined so that there is great lack of information for morphological systematics. As
for the Annelida the most basal form of the polychaetes is unknown. It is again either a
burrowing form (similar to Opheliidae or Questidae) or an aciculate, epifaunal form
(Rouse & Pleijel, 2003).
The most complete approach to date to bring some light into the relationships between
the different families of the Polychaeta was presented in 1997 by Rouse and Fauchald.
They presented a cladistic analysis on morphological data covering almost all
polychaete groups then distinguished. Rouse and Pleijel (2006) give an excellent
summary of this analysis including a discussion of the monophyly of different groups
based on autapomorphies. Due to a lack of appropriate alternatives based either on
morphological or molecular data sets, the tree originating from that study is used here as
systematic background where needed (App.-Fig. 1). The dendrogram suggests a
monophyletic taxon Polychaeta, with a sister taxon Clitellata, within the monophyletic
Annelida. The most basal polychaetes here are the simple-bodied Scolecida such as
Orbinidae, Opheliidae, and Maldanidae. Two sister taxa of the Scolecida are proposed,
the vagile Aciculata consisting of the Phyllodocida and Eunicida, and the less motile,
often tube-dwelling Canalipalpata with the Sabellida, Terebellida, and Spionida. In
contrast to former classifications, such as that of Fauchald, 1977, the Sabellariidae are
considered to be Sabellida instead of Terebellida (Rouse & Fauchald, 1997).
MATERIAL & METHODS
21
2 Material and methods
2.1 The expeditions ANDEEP I, II and III: sampling and on-board treatment
The expeditions ANDEEP I and II took place from January 23rd, 2002 to April 1st, 2002.
Starting from Punta Arenas, Chile, samples were taken in the Drake Passage, off
Elephant Island, the Bransfield Strait, across the Weddell Sea, and off the South
Sandwich Islands. In early 2005 (January 21st to April 6th) the expedition ANDEEP III
started in Cape Town, South Africa, to take further samples on a transect from South
Africa to the eastern Weddell Sea, crossing the Weddell Sea to the Antarctic Peninsula,
sampling also some sites near the South Orkney Islands, in the Drake Passage, and in
the Bransfield Strait (Fig. 2).
Fig. 2: Position of stations: ANDEEP I/II (white stations) and ANDEEP III (black stations). Map by A.
Brandt
Four different gears were deployed to sample the epifauna. This study is based on the
samples taken by an epibenthic sledge (EBS) at 29 different stations (Tab. 1). The EBS
was constructed at the Ruhr-University of Bochum, a detailed description of
MATERIAL & METHODS
22
construction and deployment is given by Brenke, 2005. The EBS consists of two nets,
the epi- and the supranet, each 1 m wide (Fig. 3). During deployment, the EBS was
trawled on the bottom for 10 minutes steaming time (speed approximately 1.0 Kn) as
soon as it touched the ground. Then the ship was stopped and the EBS was hauled in
with 0.5 m/s winch speed. The exact times when the EBS touched the ground, started to
trawl, and left the ground again was determined by curve changes in the tension
recorder protocol. This way the trawling distance and sampling area (trawling distance x
EBS width [1.0 m]) could be determined with minimum error (Tab. 1).
For optimal conservation for molecular methods the samples were immediately fixed in
precooled 96 % ethanol and gradually transferred to 70 % ethanol within the subsequent
48 hours. After final fixation in 70 % ethanol, samples were partly sorted to higher
taxonomic levels on board. After the end of the expeditions samples were shipped to the
German Center for Marine Biodiversity Research (DZMB) in Wilhelmshaven and to the
Zoological Museum, University of Hamburg where the sorting to higher taxa was
completed. Then, the Polychaeta were transferred to the Ruhr-University of Bochum for
final analyses.
Fig. 3: Epibenthic sledge: model deployed during the expeditions ANDEEP I-III. Photo by M. Schüller &
N. Brenke
MATERIAL & METHODS
23
Tab. 1: Station coordinates and trawling distances of the expeditions ANDEEP I-III
station date latitude longitude location depth (m) trawling distance (m)
41-3 26.01.2002 59°22.24'S- 59°22.55'S
60°4.06'W- 60°4.01'W Ona Basin/ Drake Passage 2360 1298
42-2 27.01.2002 59°40.29'S- 59°40.32'S
57°35.43'W-57°35.64'W Ona Basin/ Drake Passage 3690 2318
46-7 30.01.2002 60°38.35'S- 60°38.06'S
53°57.36'W-53°57.51'W Elephant Island/ Drake Passage 2889 2232
131-3 05.03.2002 65°19.83'S- 65°19.99'S
51°31.62'W-51°31.23'W Central Weddell Sea 3050 1898
132-2 06.03.2002 65°17.74'S- 65°17.62'S
53°22.82'W-53°22.86'W Central Weddell Sea 2086 1328
133-3 07.03.2002 65°20.15'S- 65°20.09'S
54°14.35'W-54°14.36'W Central Weddell Sea 1123 908
134-4 09.03.2002 65°19.20'S- 65°19.05'S
48°3.81'W- 48°2.92'W Central Weddell Sea 4069 2378
135-4 11.03.2002 65°0.06'S- 48°2.92'S
43°1.19'W- 43°0.83'W Central Weddell Sea 4678 2708
141-10 23.03.2002 58°25.08'S- 58°24.63'S
25°0.77'W- 25°0.74'W South Sandwich Islands 2313 1508
143-1 25.03.2002 58°44.69'S- 58°44.45'S
25°10.27'W-25°10.66'W South Sandwich Islands 774 819
16-10 26.01.2005 41°07.55'S-41°07.02'S
09°55.94'E-09°54.85'E South Africa/ South Atlantic 4720 3198
21-7 29.01.2005 47°39.87'S-47°38.52'S
04°15.79'E-04°14.94'E South Africa/ South Atlantic 4574 2923
59-5 14.02.2005 67°30.75'S-67°29.81'S
00°00.23'W-00°01.94'E Atlantic Sector of SO 4651 2878
74-6 20.02.2005 71°18.42'S-71°18.33'S
13°58.21'W-13°57.65'W Eastern Weddell Sea 1047 711
78-9 22.02.2005 71°09.52'S-71°09.34'S
14°00.76'W-13°58.85'W Eastern Weddell Sea 2182 2376
80-9 23.02.2005 70°38.45'S-70°39.18'S
14°42.86'W-14°43.43'W Eastern Weddell Sea 3138 1778
81-8 24.02.2005 70°31.08'S-70°32.23'S
14°34.82'W-14°34.90'W Eastern Weddell Sea 4419 2935
88-8 27.02.2005 68°03.84'S-68°03.64'S
20°31.39'W-20°27.49'W
Transect eastern to central Weddell Sea 4928 3488
94-14 02.03.2005 66°39.08'S-66°37.16'S
27°09.26'W-27°10.13'W
Transect eastern to central Weddell Sea 4889 3476
102-13 06.03.2005 65°33.18'S-65°34.32'S
36°33.24'W-36°31.05'W
Transect eastern to central Weddell Sea 4817 3283
110-8 10.03.2005 64°59.20'S-64°00.91'S
43°02.05'W-43°02.10'W
Transect eastern to central Weddell Sea 4698 2904
121-11 14.03.2005 63°38.27'S-63°37.31'S
50°37.16'W-50°38.04'W Central Weddell Sea 2668 1945
133-2 16.03.2005 62°46.73'S-62°46.33'S
53°02.57'W-53°04.14'W Central Weddell Sea 1582 1164
142-5 18.03.2005 62°11.36'S-62°11.36'S
49°27.62'W-49°29.57'W Central Weddell Sea 3403 2251
150-6 20.03.2005 61°49.13'S-61°48.52'S
47°27.51'W-47°28.16'W South Orkney Islands 1970 1567
151-7 21.03.2005 61°45.67'S-61°45.42'S
47°07.19'W-47°08.07'W South Orkney Islands 1178 1383
152-6 23.03.2005 62°20.64'S-62°19.91'S
57°53.12'W-57°53.68'W Elephant Island/ Drake Passage 2002 2113
153-7 29.03.2005 63°19.82'S-63°19.18'S
64°36.44'W-64°37.53'W King George Island 2014 1954
154-9 30.03.2005 62°32.52'S-62°31.31'S
64°39.45'W-64°38.66'W King George Island 3784 2525
MATERIAL & METHODS
24
2.2 Taxonomic analyses
2.2.1 Final sorting, identification, and labeling of species
Polychaete specimens were sorted to family level, the number of individuals per family
was documented. The families Ampharetidae, Glyceridae, Goniadidae, Hesionidae,
Nephtyidae, Nereididae, Opheliidae, Sabellariidae, Scalibregmatidae, Sphaeodoridae,
Syllidae, Terebellidae, and Trichobranchidae were determined to species level.
Specimens of different stations and different EBS nets were documented separately in
species assemblage matrices. Specimens are stored in single vials in 70 % ethanol at the
Ruhr-University of Bochum and will be transferred to the Zoological Museum,
University of Hamburg after completion of the ANDEEP project.
Identification of species took place mainly with help of the monographs and studies of
Hartman, 1964, 1966, 1967, 1978, Fauchald, 1977, and Hartmann-Schröder &
Rosenfeldt, 1988, 1989, 1990, 1991, 1992. For species not found in these works,
original species descriptions were consulted. Species of questionable identity and new
species were named by the lowest taxonomic level known plus chronological numbers.
New species also found by Dr. B. Ebbe during former studies were named with Ebbe’s
denotation (pers. comments) plus the appendix BH to achieve constant labeling.
2.2.2 Description of new species
Fifteen new species were described. The type material is stored at the Ruhr-University
Bochum and will be transferred to Zoological Museum, Hamburg after publication of
descriptions. For magnification of specimens a stereomicroscope type Olympus SZH 10
and a microscope type Olympus BX40 were used. Drawings were prepared with
Rotring Rapidograph pens, sizes 0.25 mm and 0.18 mm. After completion, drawings
were scanned (600 dpi, tiff files), digitally optimized, and labeled with Adobe
Photoshop 7.0.
MATERIAL & METHODS
25
2.2.3 Construction of identification keys
Based on the sampled material and information from literature (e.g. Fauchald, 1977;
Fauchald & Rouse, 1997; Hartmann-Schröder, 1996) identification keys were prepared.
The keys include species found in the ANDEEP I-III material only. During construction
of keys emphasis was laid upon identification by simple distinct characters that do not
require complex preparation.
2.3 Univariate and multivariate community analyses
Analyses of species data were made with Microsoft Office Excel and PRIMER v 6.1.6,
a program designed for the analysis of biological data in biodiversity and monitoring
studies (Clarke & Warwick, 2001).
2.3.1 Species accumulation plots
To find out if the sampling volume covers the maximum of expected species for the
sampled area a species accumulation plot is drawn for all species of ANDEEP I-III. By
means of UGE analyses (after Ugland, Gray & Ellingsen, 2003) a smoothed S curve
based on 999 permutations is achieved. A rising curve indicates an undersampling of
the area, meaning that not all species present have been collected. A graph converging
to maximum values indicates that all species of the area are sampled.
2.3.2 Standardization and transformation of sampling data
The sampling volume of all stations is different due to different trawling distances.
Additionally, samples of ANDEEP I/II are incomplete, not all net samples are available
for this study. Before similarity analyses, the species assemblage matrix of ANDEEP
MATERIAL & METHODS
26
I/II is therefore transformed into a presence/ absence matrix. The same applies to the
combined matrix of ANDEEP I-III.
The sample data of ANDEEP III are complete. Species abundance is standardized to
1000 m trawling distance and quantitative data are accomplished. Before the calculation
of similarities fourth root transformation is applied to reduce the influence of the few
very abundant species and increase that of the numerous rare species.
2.3.3 Univariate biodiversity measures
For the data of ANDEEP III the Shannon Index was chosen as biodiversity measure:
H’ = -∑i pi log (pi), with log- base = e
pi = Ni/Ntotal
Ni = number of individuals of species i
Ntotal = number of individuals per station
This index relates the number of individuals of each species to the number of
individuals of all species from one sample. It is the one most used in biodiversity
studies and allows the comparison of this study to data from literature. Due to its high
dependence on the sample volume it is not suitable for non-quantitative data.
In addition, the Pielou’s Evenness Index is calculated for ANDEEP III-samples:
J = H’/ log S, with S = number of species per station
The index describes how evenly individuals between the species of one station are
distributed. It is based on the Shannon Index and underlies the same dependences.
For the non-quantitative data of ANDEEP I-III the Margalef’s Index for species
richness is chosen:
d = (S-1) / log N, with S = number of species per station
N = number of individuals per station
MATERIAL & METHODS
27
2.3.4 Similarity measures
The similarities between different stations were measured with the Bray-Curtis Index:
Sjk = 100 (1 – [∑pi=1⏐yij - yik⏐] / [∑p
i=1(yij + yik)]), with yij/k = ith species in the
jth/ kth sample
i = 1, 2, 3, …, p
It is the most used similarity measure in ecological studies. Its main characteristics are:
1.) S = 0 when there are no similarities and S = 100 when samples are identical
2.) Changes in scales (e.g., g mg) do not change the result
3.) The lack of a species in both samples compared does not change the result
which is thus exclusively based on similarities
4.) For a presence/ absence matrix the Bray-Curtis Index equals the Sörensen
Coefficient
The index was applied to all ANDEEP matrices.
For comparison of ecological data, differences between stations instead of similarities
are measured by means of Euclidean distance:
djk = √∑pi=1(yij - yik)2, with yij/k = ith value in the jth/ kth
sample
i = 1, 2, 3, …, p
2.3.5 Comparison of EBS epi- and supranets
To determine if samples of the epi- and supranet of each station are to be treated as
different stations or as pseudoreplicates (results in a summation of the nets to a single
abundance value per station) a one-way ANOSIM (ANalysis Of SIMilarities) with 999
permutations was carried out on the species resemblance matrix of ANDEEP III
(abundances standardized to 1000 m trawling distance, 4th root transformation). The
MATERIAL & METHODS
28
method compares the similarities (R) between nets of one station with that of different
stations:
R = (rB-rW)/(1/2M), with rB = average similarities of nets between stations
rW = average similarities of nets within stations
M = n(n-1)/2, with n = number of samples
R = 1 expresses that nets of one station are more similar to each other than nets of
different stations. R = 0 expresses the null hypothesis that no differences in similarities
between the nets of one station and that of different stations exist. During permutation
the sample labels are altered and random data matrices are constructed. The R values for
these test-matrices are determined and compared to that of the original data. If the R
value of the original data lies outside the range of all random values, the null hypothesis
can be rejected with a significance level (p) of 1 to 999 (in case of 999 permutations),
and p < 0.1 %.
2.3.6 Cluster analysis and MDS plotting
Based on the Bray-Curtis resemblance matrices similarities and relations between
different stations are visualized by clustering and multi dimensional plotting (MDS)
(Clarke & Warwick, 2001).
Cluster analysis was done hierarchically with group-average linkage, resulting in a
dendrogram. This is done by first grouping the two most similar stations. A new
resemblance matrix is now created. The similarity of the formed group to other stations
equals the average of the similarities of the single stations of this group to other stations.
Then again the most similar stations are grouped together, and so on. The x-axis of the
resulting dendrogram presents the different groupings, the y-axis the percental
similarities. Statistical significance of the groups is tested by a SIMPROF (similarity
profile permutation) test and visualized by the shape of the branches. Drawn through
branches are well, dotted branches weakly supported.
MATERIAL & METHODS
29
During MDS plotting, station similarities are visualized by relative distances in a multi
dimensional space. A two dimensional space is chosen for this study. MDS plots are
achieved by the following steps (compare Kruskal, 1964):
1. Construction of a random starting configuration on the dimensional space
chosen
2. Construction of a Shepard diagram comparing the actual similarities of stations
with that in the random diagram
3. Construction of a regression line in the Shepard diagram giving the line of
minimal stress level between the actual similarities and the diagram distances
4. Calculation of the present stress level (expresses the differences between the
actual similarities and diagram ordination)
5. Movement of ordination points towards the direction of decreasing stress
6. Repetition of steps 2 to 5 until no lower stress level can be achieved
The stress level shows how well the similarities are mirrored by the ordination in the
end. Stress levels below 0.1 are very good values, below 0.2 show potentially good
results. Higher stress levels have to be treated with caution, they might indicate random
ordinations.
2.3.7 Environmental factors and species sets explaining station similarities
2.3.7.1 BIO-ENV
Based on the assumption that stations with similar environments have similar species
compositions, the resemblance matrices of these two descriptive factors can be
correlated to each other by rank correlation coefficients. Coefficients are based on
rankings of stations instead of total values because the resemblance matrices are
calculated by different coefficients (species data: Bray-Curtis Index, ecological data:
Euclidean distances).
MATERIAL & METHODS
30
For this study the Spearman coefficient is chosen:
ρ = 1 – 6 / (N (N2-1)) ∑Ni=1 (ri - -si)2, with N = n (n-1) / 2 (n = number of samples)
ri / si = elements of resemblence matrices
The Primer function BIO-ENV uses this coefficient to compare every possible
combination of environmental factor sets to the species matrix. The sets resulting in the
highest ρ-values present a likely explanation for found species community structures.
Significance of the result is tested by a permutation test (RELATE with 99
permutations) based on the same procedure as the ANOSIM test explained above.
The BIO-ENV method is applied to the data of ANDEEP I/II comparing species
composition with sediment grain size, O2 depth, and water depth.
2.3.7.2 BV Step
On the lines of the BIO-ENV procedure species matrices can be compared to
themselves to find the set of species dominating station similarities. Due to the huge
data size the BV Step method is used. This method is also based on the Spearman
coefficient. It first searches for the species resulting in the highest ρ-value; then
stepwise adds further species, always in search of the maximum ρ-value for the sets,
until no higher values can be achieved.
The method is applied to the ANDEEP I-III species matrix reduced to species
contributing a minimum of 4 % to any station. The starting point of the procedure was
determined with six random variables and ten repetitions, the significance of the result
is tested by a 99 permutation RELATE test.
MATERIAL & METHODS
31
2.3.8 Average Taxonomic Distinctness (AvTD) and Variations in Taxonomic
Distinctness (VarTD)
The AvTD expresses the expected taxonomic distance between two randomly chosen
different species. For presence/ absence matrices it is defined as:
Δ+ = [∑∑i<j ωij] / [S (S-1) / 2], with ωij = distance between the species i and j
S = total number of species in the sample
The VarTD [Λ+] describes the expected variation of the actual taxonomic distances
between two randomly chosen species and the AvTD:
Λ+ = [∑∑i<j (ωij-Δ+)2] / [S (S-1) / 2], with ωij = distance between the species i and j
S = total number of species in the sample
In this study the AvTD and VarTD of the stations of ANDEEP I-III (presence/ absence
matrix) are compared with a master list including all species found during the
expeditions. The master list gives the expected values for the sampling area. It also
functions as a systematic tree for the analysis. It assigns species to their according
genera (with a taxonomic distance of 1) and genera to their according families
(taxonomic distance of 2). The AvTDs and VarTDs of different stations are separately
compared to those of the master list in funnel plots. Also shown is the 95 % significance
range for the values. A combined analysis of both two values is shown in elliptic
graphs. This comparison is advisable in case negative correlations between AvTD and
VarTD at some stations exist.
MATERIAL & METHODS
32
2.4 Reconstruction of vertical and global distribution patterns
2.4.1 Vertical distribution patterns
Vertical distribution patterns are reconstructed by dividing water depths into depth
ranges of 1000 m each and marking the ranges in which the different species were
found.
2.4.2 Global distribution patterns
The global distribution of named species is reconstructed based on former records of
species. Charts were created with the programs GEBCO and Adobe Photo Shop 7.0. For
comparison of the distribution patterns within the families the global distribution was
split into six categories: LR- locally restricted to certain areas within the Southern
Ocean, SU- Subantarctic (south of 45°S), SA- also found in the southern Atlantic, SP-
also found in the southern Pacific, SH- southern hemisphere, C- cosmopolitan.
All records used for the construction of the global patterns are taken from the literature
(Augener, 1912, 1932; Benham, 1921, 1927; Blake, 1981; Blankensteyn & Lana, 1986;
Ehlers, 1897, 1900, 1901, 1908, 1912, 1913; Fauchald, 1972; Fauvel, 1916, 1936, 1941,
1951; Gillet & Dauvin, 2000; Gravier, 1906a, 1906b, 1906c, 1907a, 1907b, 1911a,
1911b; Grube, 1877; Hartley, 1985; Hartman, 1952, 1953, 1964, 1966, 1967, 1971,
1978; Hartman & Fauchald, 1971; Hartmann-Schröder, 1965, 1996; Hartmann-Schröder
& Rosenfeldt, 1988, 1989, 1990, 1991, 1992; Hessle, 1917; Hilbig & Blake, 2006;
Holthe, 1986; Kinberg, 1866, 1858-1910; Knox, 1962; Levenstein, 1964; Licher, 1999;
Monro, 1930, 1936, 1939; McIntosh, 1879, 1885; Perrson & Pleijel, 2005; Pocklington
& Fournier, 1987; Ramsay, 1914; San Martín & Parapar, 1997; Uschakov, 1952;
Wesenberg-Lund, 1949, 1961; Willey, 1902, unpublished data).
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 33
3 Results 3.1 Composition of polychaete communities
During the expeditions a total of 14,176 specimens were collected belonging to 47
families. 3541 specimens from 38 families originated from the expeditions ANDEEP I-
II and 10635 specimens from 46 families from ANDEEP III. Eighty-two specimens
could not be identified to family level due to poor condition. The complete assemblage
matrices are given in App.-Tab. 1-6.
App.-Fig. 2 summarizes the family composition of the different stations focusing on the
most abundant families. Spionidae, Polynoidae, Opheliidae, Hesionidae, Ampharetidae,
and Cirratulidae are among the dominant families at most stations. Spionidae,
Polynoidae, Opheliidae, and Hesionidae are also among the most abundant families
overall, together with the Syllidae, Pholoididae, Sphaerodoridae, and Terebellidae. The
latter are only seldomly dominant at one station, they have a more even distribution
over all stations.
Only the families Ampharetidae, Glyceridae, Goniadidae, Hesionidae, Nephtyidae,
Nereididae, Opheliidae, Sabellariidae, Scalibregmatidae, Sphaerodoridae, Syllidae,
Terebellidae, and Trichobranchidae were determined to species level. These included
6669 specimens from 155 species (ANDEEP I-II: 1308 specimens/ 89 species,
ANDEEP III: 5361 specimens/ 130 species). App.-Tab. 4-6 give a detailed overview of
the species composition at each station.
Among the 155 discriminated species a total of 46 species are new to science, 17
species could not be determined to species level without doubt (labeled as aff. or cf.).
Twenty species could only be determined to higher taxonomic levels due to low
abundance and poor condition. It is, therefore, unclear whether these specimens are
already named species or new to science. In course of this study 18 new species were
described, three of these descriptions are already published (Schüller & Hilbig, 2007);
the remaining 15 descriptions are presented in the following chapters. Additionally,
species identification keys are presented considering only the species found and
determined in this study.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 34
3.2 Identification keys to selected species found during ANDEEP I-III
3.2.1 Key to the Ampharetidae MALMGREN 1866
A total of 464 Ampharetidae have been collected belonging to 34 species. One
specimen was a juvenile of the genus Amphicteis and could not be determined to any
species. Ten species (Ampharetidae ssp. 1 – 8, Eusamythella sp. 1, and Samytha sp. 1)
were only present with few individuals of poor condition, a determination of these
species was not possible. The determination of three further species, respectively
genera, is in question. Eight species new to science were collected.
1 Anterior chaetigers with needle-like uncini. Segment four with nuchal hooks.
Segment six with a pectinate postbranchial membrane (App.-Fig. 3A)...Melinna
only M. cristata
# Uncini never needle-like. Nuchal hooks and postbranchial membrane absent
…………………………………………………………………………………...2
2 Four pairs of branchiae…………………………………………………………..3
# Three pairs of branchiae………………………………………………………..10
3 Palae present (App.-Fig. 3B)…………………………………………………….4
# Palae absent………………………………………………………………………7
4 Twelve thoracic uncinigers………………………………………………………5
# Fourteen thoracic uncinigers………………………………………….Amphicteis
Palae short, few. Abdominal segments number 10. Pygidium with 2 dorsal cirri……A. gunneri Palae fine, of median length. 10 abdominal segments. Last thoracic segment dorsally with 2
wing-like appendages (App.-Fig. 3C)…………………………………………………A. sp. 1 With eyes. Prostomium anteriorly tapering, projecting upwards. Dorsal ridge on segment 3.
Wing-like appendages on last thoracic segment. Palae fine, of median length………….A. sp. 2 Prostomium with 2 button-shaped anterior projections. Palae long and coarse. Abdominal
notopodial rudiments button-shaped to spherical, posterior segments inflated………….A. sp. 3
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 35
5 Fourth or fifth to last thoracic chaetiger with elevated parapodia or dorsal ridge.
Uncini from third or fourth thoracic chaetiger…………………………………..6
# Dorsal ridge absent. Parapodia not elevated. Uncini from third thoracic
chaetiger. Abdomen without notopodial rudiments…………………..Ampharete
only A. kerguelensis
6 Fifth-to-last parapodia elevated (App.-Fig. 3D) with hirsute chaetae. Branchiae
and buccal tentacles papillose………………………………………Anobothrella
Palae slender, arranged in a whirl………………………………………………A. antarctica
Palae slender, arranged in a straight line………………………………………………….A. sp. 1
# Fourth or fifth to last thoracic chaetiger with dorsal ridge. Branchiae and buccal
tentacles smooth……………………………………………………...Anobothrus
Palae long and slender. Uncini from fourth segment after palae……………………...A. gracilis Palae stout. Suddenly tapering to a fine tip. Uncini from third segment after palae (Fig. 4A)
……………………………………………………………………..A. pseudoampharete sp.n.
7 Twelve thoracic uncinigers………………………………………………………8
# Fourteen thoracic uncinigers……………………………………...Amphisamytha
# Eleven thoracic uncinigers………………………………………………...Amage
only A. sculpta
8 Parapodia not elevated…………………………………………………………...9
# Third to last thoracic parapodia elevated (App.-Fig. 3D)……………..Sosanopsis Prostomium distinctly scoop shaped, trilobed (App.-Fig. 3E-F), branchiae all of similar width
…………………………………………………………………………………...S. kerguelensis
Prostomium not scoop shaped. Median branchiae distinctly broader than lateral ones…..S. sp. 1
9 Branchial position nearly segmental. Branchiae and buccal tentacles smooth
……………………………………………………………………….Grubianella Prostomium anterior with two spherical processes. Posterior end inflated with two long cirri
……………………………………………………………………………….G. antarctica Prostomium anteriorly slightly incised. 12 abdominal segments. Posterior end not inflated
……………………………………………………………………………………………G. sp. 1
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 36
# Branchial position one in front of the other three. Branchiae lamellate
………………………………………………………………………Phyllocomus
only P. crocea
10 With palae (App.-Fig. 3B)……...………………………………………………11
# Without palae…………………………………………………………………...13
11 Twelve thoracic uncinigers……………………………………………………..12
# Nine thoracic uncinigers. Last notopodia elevated, chaetae crossing over dorsum
(App.-Fig. 3G)……………………………………………………………..Mugga
only M. sp. 1BH
12 Uncini from third thoracic chaetiger. Abdomen without notopodial rudiments
……………………………………………………………………...Neosabellides
only N. elongatus
# Uncini from fourth thoracic chaetiger. Abdomen with notopodial rudiments
……………………………………………………………………...Eusamythella
13 More than ten thoracic uncinigers……………………………………………...14
# Ten thoracic uncinigers……………………………………………….Muggoides
Last notopodia slightly elevated……………………………………………..M. cinctus
Last notopodia not elevated……………………………………………………...M. sp. 1BH
14 Fourteen thoracic uncinigers……………………………………………………15
# Eleven thoracic uncinigers. Prostomium scoop-shaped. Abdomen without
notopodial rudiments…………………………………………..Glyphanostomum
only G. scotiarum
15 Uncini from fourth thoracic chaetiger…………………………………...Samytha
# Uncini from third thoracic chaetiger. Buccal tentacles short, on thick membrane.
Outermost branchiae grooved…………………………………………...Amythas
only A. membranifera
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 37
Annotated species list
Amage sculpta EHLERS 1908
Stations (No of specimens): 42-2 (1), 21-7 (8), 74-6 (1), 80-9 (3), 121-11 (1), 133-2 (2)
Distribution: subantarctic, 73 – 4574 m
Records: Benham, 1927; Ehlers, 1908; Hartman, 1966; 1967; 1978; Hartmann-Schröder
& Rosenfeldt, 1989; 1991; Monro, 1930; 1936
Ampharete kerguelensis MCINTOSH 1885
Stations (No of specimens): 41-3 (2), 46-7 (7), 16-10 (13), 59-5 (1), 78-9 (7), 81-8 (6),
133-2 (1)
Distribution: subantarctic and Southern Atlantic, 0 – 4720 m
Records: Augener, 1932; Ehlers, 1913; Hartman, 1966; 1967; 1978; Hartmann-Schröder
& Rosenfeldt, 1989; 1991; Hessle, 1917; McIntosh, 1885; Monro, 1936; 1939
Ampharetidae sp. 1
Stations (No of specimens): 141-10 (1)
Ampharetidae sp. 2
Stations (No of specimens): 134-4 (2), 21-7 (1), 78-9 (2), 121-11 (2), 142-5 (1),
153-7 (5)
Ampharetidae sp. 3
Stations (No of specimens): 134-4 (2)
Ampharetidae sp. 4
Stations (No of specimens): 151-7 (1)
Ampharetidae sp. 5
Stations (No of specimens): 78-9 (2)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 38
Ampharetidae sp. 6
Stations (No of specimens): 80-9 (1), 81-1 (1)
Ampharetidae sp. 7
Stations (No of specimens): 81-8 (2)
Ampharetidae sp. 8
Stations (No of specimens): 151-7
Amphicteis gunneri (SARS 1835)
Stations (No of specimens): 46-7 (1), 78-9 (1), 80-9 (1), 121-11 (1)
Distribution: cosmopolitan, 5 – 3138 m
Records: Augener, 1932; Hartley, 1985; Hartman, 1952; 1953; 1966; Hartmann-
Schröder, 1996; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Hessle, 1917;
Holthe, 1986; Monro, 1930; 1936; 1939
Amphicteis juvenile
Stations (No of specimens): 78-9 (1)
Amphicteis sp. 1
Stations (No of specimens): 41-3 (1), 78-9 (15), 81-8 (1), 121-11 (32), 133-2 (1),
150-6 (8), 153-7 (3)
Amphicteis sp. 2
Stations (No of specimens): 78-9 (7)
Amphicteis sp. 3
Stations (No of specimens): 74-6 (5), 80-9 (1), 142-5 (1), 150-6 (2), 154-9 (1)
cf. Amphisamytha
Stations (No of specimens): 74-6 (1)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 39
Amythas membranifera BENHAM 1921
Stations (No of specimens): 133-2 (1), 142-5 (3)
Distribution: subantarctic, 437 – 3403 m
Records: Benham, 1921; Hartman, 1966; Monro, 1939
Anobothrella antarctica (MONRO 1939)
Stations (No of specimens): 143-1 (1), 74-6 (3), 81-8 (1), 150-6 (1)
Distribution: subantarctic, 40 – 4419 m
Records: Hartman, 1967; 1978; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Monro,
1939
Anobothrella sp. 1
Stations (No of specimens): 81-8 (1)
Anobothrus gracilis (MALMGREN 1866)
Stations (No of specimens): 81-8 (2), 102-13 (1), 133-2 (7), 154-9 (2)
Distribution: cosmopolitan, shelf down to 4817 m
Records: Hartmann-Schröder, 1996; Hessle, 1917; Hilbig & Blake, 2006
Anobothrus pseudoampharete sp.n.
Stations (No of specimens): 42-2 (1), 133-3 (1), 143-1 (7), 21-7 (1), 74-6 (104),
81-8 (3), 121-11 (9), 13-2 (4), 142-5 (1), 150-6 (12), 154-9 (1)
cf. Eusamythella
Stations (No of specimens): 42-2 (4), 74-6 (1), 81-8 (2)
Glyphanostomum scotiarum HARTMAN 1978
Stations (No of specimens): 42-2 (3), 135-4 (1), 74-6 (3), 78-9 (1)
Distribution: Drake Passage to Weddell Sea, 298 – 4678 m
Records: Hartman, 1978
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 40
Grubianella antarctica MCINTOSH 1885
Stations (No of specimens): 42-2 (2), 16-10 (2), 153-7 (1)
Distribution: subantarctic to Southern Atlantic, 412 –4720 m
Grubianella sp. 1
Stations (No of specimens): 74-6 (1)
Melinna cristata (SARS 1851)
Stations (No of specimens): 80-9 (4), 150-6 (4)
Distribution: cosmopolitan, 0 – 3803 m
Records: Ehlers, 1908; Hartman, 1966; 1967; Hartmann-Schröder, 1996; Hartmann-
Schröder & Rosenfeldt, 1989; Hessle, 1917; Holthe, 1986; Monro, 1930
Mugga sp. 1BH
Stations (No of specimens): 141-10 (1), 16-10 (3), 154-9 (1)
Muggoides cf. cinctus HARTMAN 1965
Stations (No of specimens): 81-8 (11)
Distribution: cosmopolitan, down to 4419 m
Records: Ebbe (pers. comment)
Muggoides sp. 1BH
Stations (No of specimens): 42-2 (1), 16-10 (2), 102-13 (4)
Neosabellides elongatus (EHLERS 1912)
Stations (No of specimens): 46-7 (2), 134-4 (1), 21-7 (1), 74-6 (4), 78-9 (2), 80-9 (1),
81-8 (1), 133-2 (1), 153-7 (2)
Distribution: subantarctic, 0 – 4574 m
Records: Benham, 1927; Ehlers, 1912; 1913; Hartman, 1953; 1966; 1967; 1978;
Hartmann-Schröder & Rosenfeldt, 1989; 1991; Monro, 1930; 1936; 1939
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 41
cf. Neosabellides
Stations (No of specimens): 74-6 (1)
Phyllocomus crocea GRUBE 1877
Stations (No of specimens): 59-5 (2), 74-6 (2), 81-8 (2), 151-7 (2)
Distribution: subantarctic, 68 – 4651 m
Records: Augener, 1932; Benham, 1921; Grube, 1877; Hartman, 1966; 1967;
Hartmann-Schröder & Rosenfeldt, 1989; Hessle, 1917; Monro, 1930; 1936;
1939
Samytha sp. 1
Stations (No of specimens): 74-6 (1)
Sosanopsis kerguelensis MONRO 1939
Stations (No of specimens): 131-3 (3), 134-4 (2), 16-10 (4), 21-7 (6), 59-5 (4), 74-6 (3),
78-9 (1), 80-9 (2), 88-8 (1), 94-14 (3), 102-13 (1), 110-8 (2), 121-11 (3),
133-2 (8), 150-6 (2), 153-7 (6), 154-9 (1)
Distribution: subantarctic and Soutern Atlantic
Records: Hartman, 1966; 1978, Monro, 1939, 20 – 4928 m
Sosanopsis sp. 1
Stations (No of specimens): 150-6 (5), 153-7 (2)
3.2.2 Glyceridae GRUBE 1850
The Glyceridae are represented in the samples by 353 specimens that all belong to one
species.
Glycera kerguelensis MCINTOSH 1885
Stations (No of specimens): 41-3 (1), 42-2 (2), 46-7 (3), 135-4 (17), 141-10 (23),
143-1 (31), 16-10 (4), 21-7 (2), 59-5 (14), 74-6 (43), 78-9 (47), 80-9 (45),
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 42
81-8 (2), 94-14 (19), 102-13 (8), 110-8 (29), 133-2 (19), 150-6 (6), 153-7 (4), 1
54-9 (5)
Distribution: subantarctic and Southern Atlantic, 40 – 4889 m
Records: Hartman, 1964; 1967; 1978; Hartmann-Schröder & Rosenfeldt, 1988; 1990;
1992; McIntosh, 1885
3.2.3 Key to the Goniadidae KINBERG 1866
Six goniadid specimens from three species have been collected, two of them are new to
science.
1 Proboscis lateral with chevrons (App.-Fig. 3H). Neurochaetae only spinigers
…………………………………………………………………………...Goniada
only G. maculata
# Proboscis without chevrons. Neurochaetae variable………………Bathyglycinde
Prechaetal lobes bilobate……………………………………………...Bathyglycinde sp. 1BH
Prechaetal lobes simple……………………………………………………Bathyglycinde sp. 2
Annotated species list
Bathyglycinde sp. 1BH
Stations (No of specimens): 42-2 (3), 16-10 (1)
Bathyglycinde sp. 2
Stations (No of specimens): 42-2 (1)
Goniada maculata ÖRSTEDT 1843
Stations (No of specimens): 16-10 (1)
Distribution: cosmopolitan, 1550 – 4720 m
Records: Hartman, 1950; Hilbig & Blake, 2006
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 43
3.2.4 Key to the Hesionidae GRUBE 1850
Ten species of at least six genera are found for the Hesionidae (1420 specimens).
Fourteen specimens from two species could not be identified (Hesionidae sp. 1, aff.
Hesionides). Five species are new to science, four of them are described in this study.
1 Four pairs of tentacular cirri, three antennae………………………….Hesionides
# Six or eight pairs of tentacular cirri……………………………………………...2
2 Six pairs of tentacular cirri……………………………………………………….3
# Eight pairs of tentacular cirri…………………………………………………….5
3 Two antennae. Parapodia uniramous…………………………………………….4
# Three antennae. Parapodia biramous. Eversible pharynx distally with numerous
fine cirri…………………………………………………………….Ophiodromus
Dorsal cirri long, articulated. Two pairs of eyes…………………………………….O. comatus Dorsal and ventral cirri smooth. Eyes absent. Posterior margin of prostomium light brown in
ethanol (Fig. 5E-H)……………………………………………O. calligocervix sp.n.
4 Eversible pharynx distally with numerous fine cirri (Fig. 6D)……...Parasyllidea
only P. delicata sp.n.
# Eversible pharynx with eleven distal papillae (Fig. 6A)……...…...Micropodarke
M. cylindripalpata sp.n.
5 Three antennae. Parapodia biramous…………………………………………….6
# Two antennae. Parapodia uniramous. Eversible pharynx distally with numerous
fine cirri………………………………………………………………Kefersteinia
only K. fauveli
6 Median antenna attached medially (Fig. 5A). Pharynx terminally with ring of
numerous cirri………………………………………………………..Amphiduros
Notochaetae chambered and serrated (Fig. 5C)……………………………….A. serratus sp.n.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 44
Notochaetae not serrated………………………………………………………………….A. sp. 1
# Median antenna attached frontally (App.-Fig. 3J). Eversible pharynx with 40
distal papillae……………………………………………………………….Gyptis
only G. incompta
Annotated species list
Amphiduros serratus sp.n.
Stations (No of specimens): 132-2 (2), 134-4 (1), 152-6 (2)
Amphiduros sp. 1
Stations (No of specimens): 46-7 (12), 131-3 (1), 141-10 (1)
Gyptis incompta EHLERS 1912
Stations (No of specimens): 74-6 (48), 133-2 (736)
Distribution: subantarctic and Southern Pacific, 18 – 1582 m
Records: Ehlers, 1912; 1913; Wesenberg-Lund, 1961
Hesionidae sp. 1
Stations (No of specimens): 131-3 (3)
aff. Hesionides
Stations (No of specimens): 16-10 (4), 21-7 (3), 59-5 (1), 88-8 (1), 133-2 (2)
Kefersteinia fauveli AVERNICEV 1972
Stations (No of specimens): 141-10 (35), 16-10 (3), 59-5 (6), 74-6 (70), 133-2 (203),
151-7 (11)
Distribution: South Atlantic and Atlantic sector of the Southern Ocean, 40 – 4720 m
Records: Hartman, 1978; Hartmann-Schröder & Rosenfeldt 1988; 1990; 1992
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 45
Micropodarke cylindripalpata sp.n.
Stations (No of specimens): 42-2 (9), 46-7 (4), 131-3 (2), 141-10 (2), 16-10 (1),
21-7 (2), 59-5 (5), 80-9 (5), 81-8 (4), 88-8 (6), 94-14 (1), 110-8 (2), 121-11 (1),
150-6 (3), 154-9 (2)
Ophiodromus calligocervix sp.n.
Stations (No of specimens): 42-2 (28), 131-3 (3), 132-2 (1), 134-4 (6), 135-4 (1),
141-10 (1), 143-1 (1), 16-10 (2), 21-7 (3), 59-5 (2), 78-9 (14), 80-9 (12),
88-8 (8), 94-14 (4), 102-13 (1), 110-8 (3), 121-11 (1), 153-7 (7), 154-9 (2)
Ophiodromus comatus (EHLERS 1912)
Stations (No of specimens): 74-6 (25), 80-9 (7), 81-8 (3), 121-11 (4), 133-2 (43),
150-6 (3), 151-7 (17), 153-7 (1)
Distribution: subantarctic, 325 – 4419 m
Records: Ehlers, 1912; 1913; Hartman, 1964
Parasyllidea delicata sp.n.
Stations (No of specimens): 131-3 (1), 16-10 (2), 21-7 (3), 80-9 (4), 110-8 (2),
142-5 (3), 150-6 (7), 53-7 (1)
3.2.5 Key to the Nephtyidae GRUBE 1850
A total of 177 specimens from three species (two genera) were found for the
Nephtyidae. The species of the genus Aglaophamus are named, that of the genus
Micronephtys is new to science.
1 Interramal cirri strongly reduced or absent………………………...Micronephtys
only M. sp. 1
# Interramal cirri involute (App.-Fig. 3I)…………………...………..Aglaophamus
Postchaetal lobes always shorter than parapodial lobes, broadly rounded….A. paramalmgreni
Postchaetal lobes large, longer than wide………………………………………A. trissophyllus
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 46
Annotated species list
Aglaophamus paramalmgreni HARTMANN-SCHRÖDER & ROSENFELDT 1992
Stations (No of specimens): 42-2 (5), 46-7 (6), 59-5 (14), 78-9 (1), 110-8 (2), 150-6 (1),
151-7 (2), 152-6 (1), 153-7 (15)
Distribution: Weddell Sea, Antarctic Peninsula, 546 – 4698 m
Records: Hartmann-Schröder & Rosenfeldt, 1992
Aglaophamus trissophyllus (GRUBE 1866)
Stations (No of specimens): 42-2 (2), 46-7 (27), 74-6 (73), 78-9 (18), 80-9 (2),
110-8 (1), 150-6 (2), 154-9 (4)
Distribution: Weddell Sea, Antarctic Peninsula, 400 – 4698 m
Records: Hartman, 1978
Micronephtys sp. 1
Stations (No of specimens): 46-7 (1)
3.2.6 Key to the Nereididae JOHNSTON 1865
The Nereididae are represented by ten specimens from three different species and
genera. Only one species is identified without doubt, one species might be new to
science.
1 Eversible pharynx without paragnaths and papillae………………………..Nicon
# Eversible pharnyx with pharyngeal processes…………………………………...2
2 Eversible pharynx with soft papillae. Ventral cirri partially double
…..…………………………………………………………………Ceratocephale
only C. sp. 1BH
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 47
# Eversible pharynx with conical paragnaths. Ventral cirri simple throughout.
Falcigers and spinigers in posterior notopodia…………………………….Nereis
only N. eugeniae
Annotated species list
Ceratocephale sp. 1BH
Stations (No of specimens): 78-9 (1), 88-8 (1), 110-8 (1), 153-7 (1), 154-9 (1)
Nereis eugeniae (KINBERG 1866)
Stations (No of specimens): 42-2 (1), 46-7 (1), 132-2 (1)
Distribution: Southern hemisphere, 0 – 3690 m
Records: Ehlers, 1897; 1900; 1901; Fauvel, 1941; Hartman, 1964; Hartmann-Schröder
& Rosenfeldt, 1992; Kinberg, 1866; Monro, 1930; 1936; 1939; Ramsay, 1914;
Wesenberg-Lund, 1961
cf. Nicon KINBERG 1866
Stations (No of specimens): 154-9 (1)
3.2.7 Key to the Opheliidae MALMGREN 1867
A total of 479 specimens from two genera and 14 species of Opheliidae were found. Six
of these species were already known to science, the remaining eight are new. In this
study, two species of the genus Ophelina are described, as well as three species of
Ammotrypanella. In addition, the genus Ammotrypanella is redefined and the type
species redescribed.
1 Body elongated with a ventral groove along whole body length. Branchiae
present on anterior and posterior segments……………………………..Ophelina
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 48
Without branchiae on last four chaetigers. These distinctly reduced in length (App.-Fig. 3L)
………………………………………………………………………………………..O. breviata Without anal tube. Without branchiae on last three to four chaetigers. These not distinctly
reduced (App.-Fig. 3K)…………………………………………………………..O. gymnopyge
Without branchiae and distinct segmental sutures. Reminiscent of Nematoda….O. nematoides Without branchiae at posterior segments. These ventrally concave with a pair of cirriform
processes……………………………………………………………………..O. scaphigera Anterior chaetigers numerous, prolonged, and bent. Analtube marginally with circlet of fine
cirri…………………………………………………..……………………………….O. setigera Branchiae enlarged in third charter of body and missing in last. Anal tube without ventral cirrus,
margin with numerous fine cirri (Fig. 8F)………………………..O. ammotrypanella sp.n. Anterior branchiae of median size, posterior ones enlarged. Anal tube seemingly articulated, with
robust ventral cirrus and marginally with fine cirri (Fig. 8D)…………………O. robusta sp.n. Anterior branchiae prolonged. Without branchiae on last three chaetigers. These not reduced in
length. Anal tube with ventral cirrus……………………………………………………...O. sp. 3
Branchiae only present from chaetiger 5 – 13. Segmental sutures very indistinct………O. sp. 4 Anterior branchiae of median size. Anal tube strongly reduced in length, without ventral cirrus
………………………………………………………………………………………O. sp. 5
# Body elongated with a ventral groove along whole body length. Branchiae
limited to posterior half of the body (Figs. 7)…………………..Ammotrypanella
Anal tube without ventral cirrus, margin with short cirri (Fig. 7E)……………………A. arctica
Anal tube with thick ventral cirrus, margin with short cirri (Fig. 7B)………….A. cirrosa sp.n.
Anal tube with ventral cirrus, margin smooth (Fig. 7F)……………………..A. princessa sp.n.
Without anal tube (Fig. 8B)………………………………………………….A. mcintoshi sp.n.
Annotated species list
Ammotrypanella arctica MCINTOSH 1879
Stations (No of specimens): 42-2 (2), 131-3 (7), 134-4 (1), 16-10 (6), 59-5 (4), 78-9 (1),
102-13 (2), 110-8 (14), 121-11 (22), 153-7 (2), 154-9 (3)
Distribution: cosmopolitan, 2014 – 5023 m
Records: Ebbe (pers. comment); Hartman & Fauchald, 1971; McIntosh, 1879
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 49
Ammotrypanella cirrosa sp.n.
Stations (No of specimens): 42-2 (4), 131-3 (64), 134-4 (2), 141-10 (1), 16-10 (6),
59-5 (2), 78-9 (2), 102-13 (4), 110-8 (1), 121-11 (40), 153-7 (4), 154-9 (1)
Ammotrypanella princessa sp.n.
Stations (No of specimens): 42-2 (1), 134-4 (1), 134-5 (1), 16-10 (1), 78-9 (1), 81-8 (1),
121-11 (1), 153-7 (1)
Ammotrypanella mcintoshi sp.n.
Stations (No of specimens): 16-10 (3), 21-7 (1), 59-5 (1), 74-6 (2), 110-8 (3), 121-11 (1)
Ophelina breviata (EHLERS 1913)
Stations (No of specimens): 42-2 (1), 46-7 (2), 131-3 (16), 16-10 (1), 74-6 (16),
78-9 (1), 102-13 (1), 110-8 (1), 121-11 (6), 133-2 (19), 150-6 (1), 153-7 (1),
154-9 (1)
Distribution: subantarctic and Southern Atlantic, 20 – 4720 m
Records: Augener, 1932; Hartman, 1953; 1966; 1978; Hartmann-Schröder &
Rosenfeldt, 1989; 1991; Monro, 1930; 1936; 1939
Ophelina gymnopyge (EHLERS 1908)
Stations (No of specimens): 42-2 (3), 131-3 (15), 16-10 (2), 59-5 (1), 81-8 (3),
110-8 (3), 121-11 (2), 133-2 (14), 154-9 (4)
Distribution: South Atlantic and Atlantic sector of the Southern Ocean, 10 – 4720 m
Records: Ehlers, 1908; Hartman, 1952; 1953; 1966; Hartmann-Schröder & Rosenfeldt,
1989; 1991
Ophelina nematoides (EHLERS 1913)
Stations (No of specimens): 41-3 (1), 42-2 (14), 46-7 (7), 131-3 (2), 141-10 (5),
16-10 (2), 74-6 (18), 80-9 (1), 153-7 (1)
Distribution: subantarctic and Southern Atlantic, 208 – 4758 m
Records: Ehlers, 1913; Hartman, 1966; 1967; 1978; Hartmann-Schröder & Rosenfeldt,
1989; 1991
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 50
Ophelina scaphigera (EHLERS 1901)
Stations (No of specimens): 16-10 (4), 59-5 (1), 133-2 (2)
Distribution: South Atlantic and Atlantic sector of the Southern Ocean, 18 – 4720 m
Records: Ehlers, 1900; 1901; Hartman, 1953; 1966; Monro, 1936
Ophelina setigera HARTMAN 1978
Stations (No of specimens): 46-7 (1), 131-3 (1), 121-11 (1)
Distribution: South Atlantic and Atlantic sector of the Southern Ocean, 1622 – 5575 m
Records: Ebbe (pers. comment); Hartman, 1978
Ophelina ammotrypanella sp.n.
Stations (No of specimens): 131-3 (2), 16-10 (4), 78-9 (10), 81-8 (4), 110-8 (4),
121-11 (7), 154-9 (8)
Ophelina robusta sp.n.
Stations (No of specimens): 131-3 (1), 78-9 (10), 80-9 (2), 121-11 (4), 150-6 (5),
151-7 (2), 153-7 (7)
Ophelina sp. 3
Stations (No of specimens): 131-3 (2)
Ophelina sp. 4
Stations (No of specimens): 42-2 (1)
Ophelina sp. 5
Stations (No of specimens): 133-2 (10), 150-6 (2), 154-9 (5)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 51
3.2.8 Sabellariidae JOHNSTON 1865
Only two specimens have been sampled for the Sabellariidae, belonging to one species.
Phalacrostemma elegans FAUVEL 1911
Stations (No of specimens): 46-7 (1), 80-9 (1)
Distribution: cosmopolitan, 330 – 5490 m
Records: Gillet & Dauvin, 2000; Hartman, 1971
3.2.9 Key to the Scalibregmatidae MALMGREN 1867
The Scalibregmatidae are represented by 642 specimens from 19 species. Nine different
genera were found. Six species were new to science, three of them have already been
described in course of this study, an additional description is presented here. The
identity of two species is not completely confirmed.
1 Body grub-, maggot- or barrel-shaped …………………………………………..2
# Body more elongated, sometimes anteriorly inflated, prostomium bifid or T-
shaped (Fig. 9A, App.-Fig. 3M)…………………………………………………4
2 Prostomium entire, cone-shaped…………………………………………………3
# Prostomium with two lateral processes or incised…………………...Axiokebuita
Notopodia with postchaetal lobes………………………………………………………A. millsia
Notopodia without postchaetal lobes…………………………………………………..A. minuta
3 Branchiae absent……………………………………………………………Kesun
only K. abyssorum
# Branchiae present………………………………………………………...Travisia
Segments number 23 – 27. Pygidium with anal cylinder...............................T. kerguelensis
Segements number 52 – 53, body prolonged. Pygidium with anal disc………….T. lithophila
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 52
4 Posterior parapodia reduced……………………………………………………...5
# Posterior parapodia not reduced…………………………………………………6
5 With acicular spines (App.-Fig. 3M)……………………………...Asclerocheilus
only A. ashworthi
# Without acicular spines……………………………...………………..Hyboscolex
only H. equatorialis
6 Posterior parapodia with dorsal and ventral cirri………………………………...7
# Posterior parapodia without dorsal cirri. Acicular spines present….Sclerocheilus
only S. antarcticus
7 Without branchiae………………………………………………………………..8
# With branchiae in anterior chaetigers……………………………….Scalibregma
only S. inflatum
8 Without acicular spines………………………………………Pseudoscalibregma Rounded nuchal crest dorsally on prostomium. Broad dorsal and ventral cirri
………………………………………………………………………………...P. bransfieldium Dorsal and ventral cirri enlarged, foliose to wing-like in posterior segments (Fig. 9C)
…………………………………………………………………….P. papilia sp.n.
Dorsal and ventral cirri drop-shaped, with bacillary glands………………………..P. ursapium
# With acicular spines (App.-Fig. 3M)……………………………….Oligobregma
Chaetigers 1 and 2 with one row of acicular spines each……………………………….O. blakei Chaetigers 1 and 2 with two rows, chaetiger 3 with one row of acicular spines. Interramal sense
organs present………………………………………………………………………….O. collare Chaetigers 1 and 2 with inconspicious acicular spines. Dorsal surface papillated
…………………………………………………………………………………….O. hartmanae
With Y-shaped eyes. One row of acicular spines in chaetigers 1 – 3………………….O. notiale
Chaetigers 1 and 2 with two rows of acicular spines…………………………O. pseudocollare Chaetigers 1 and 2 with two rows, chaetigers 3 and 4 with one row of acicular spines
………………………………………………………………………………..O. quadrispinosa
Acicular spines as O. collare. No interramal sense organs present………………………O. sp. 1
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 53
Annotated species list
Asclerocheilus ashworthi BLAKE 1981
Station (No of specimens): 46-7 (1)
Distribution: subantarctic, 223 – 2889 m
Records: Blake, 1981
Axiokebuita millsi POCKLINGTON & FOURNIER 1987
Stations (No of specimens): 143-1 (62)
Distribution: cosmopolitan, 532 – 3687 m
Records: Pocklington & Fournier, 1987
Axiokebuita minuta (HARTMAN 1967)
Stations (No of specimens): 46-7 (1), 131-3 (6), 133-3 (1), 134-4 (3), 141-10 (11),
74-6 (8), 80-9 (7), 81-8 (1), 133-2 (15), 150-6 (2)
Distribution: cosmopolitan, 180 – 4419 m
Records: Blake, 1981; Hartman, 1967; 1978; Persson & Pleijel, 2005
Hyboscolex equatorialis BLAKE 1981
Stations (No of specimens): 141-10 (1)
Distribution: South Sandwich Trench, west coast of South America, 0 – 2313 m
Records: Blake, 1981
Kesun abyssorum MONRO 1930
Stations (No of specimens): 46-7 (86), 131-3 (1), 134-4 (3), 141-10 (8), 21-7 (1),
59-5 (14), 78-9 (24), 80-9 (8), 81-8 (1), 94-14 (1), 102-13 (3), 110-8 (4),
121-11 (14), 133-2 (2), 142-5 (4), 150-6 (26), 153-7 (54), 154-9 (11)
Distribution: South Atlantic and Atlantic sector of the Southern Ocean, 193 – 4817 m
Records: Augener, 1932; Hartman, 1966; 1978; Monro, 1930; 1939
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 54
Oligobregma blakei SCHÜLLER & HILBIG 2007
Stations (No of specimens): 46-7 (1), 121-11 (1), 133-2 (2)
Distribution: Weddell and Scotia Sea, 1582 – 2889 m
Records: Schüller & Hilbig, 2007
Oligobregma collare (LEVENSTEIN 1978)
Stations (No of specimens): 42-2 (4), 46-7 (3), 131-3 (8), 134-4 (2), 59-5 (1), 74-6 (1),
78-9 (7), 80-9 (1), 110-8 (1), 121-11 (11), 133-2 (3), 142-5 (1), 150-6 (1),
151-7 (1), 153-7 (10), 154-9 (1)
Distribution: subantarctic, 1178 – 4698 m
Records: Hartman, 1967; 1978; Blake, 1981
Oligobregma hartmanae BLAKE 1981
Stations (No of specimens): 59-5 (3), 133-2 (2); 150-6 (1)
Distribution: Antarctic sector of the Southern Ocean, 505 – 4651 m
Records: Blake, 1981
Oligobregma notiale BLAKE 1981
Stations (No of specimens): 42-2 (7)
Distribution: subantarctic, 18 – 3690 m
Records: Blake, 1981
Oligobregma pseudocollare SCHÜLLER & HILBIG 2007
Stations ( no specimens): 46-7 (20), 131-3 (1), 143-1 (8), 80-9 (2), 121-11 (1),
133-2 (8), 150-6 (1), 153-7 (2)
Distribution: Weddell and Scotia Sea, 1582 – 3138 m
Records: Schüller & Hilbig, 2007
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 55
Oligobregma quadrispinosa SCHÜLLER & HILBIG 2007
Stations (No of specimens): 42-2 (5), 131-3 (8), 134-4 (2), 141-10 (1), 21-7 (1),
59-5 (6), 88-8 (1), 94-14 (2), 110-8 (1), 121-11 (1), 133-2 (1), 151-7 (1),
153-7 (10)
Distribution: Weddell and Scotia Sea, 1178 – 4928 m
Records: Schüller & Hilbig, 2007
Oligobregma sp. 1
Stations (No of specimens): 151-7 (1)
Pseudoscalibregma bransfieldium (HARTMAN 1967)
Stations (No of specimens): 42-2 (8), 46-7 (5), 132-2 (6), 143-1 (2), 74-6 (2), 80-9 (1),
88-8 (1), 133-2 (2), 150-6 (1), 151-7 (1), 153-7 (1)
Distribution: subantarctic, 335 – 4928 m
Records: Blake, 1981; Hartman, 1967; 1978
Pseudoscalibregma papilia n. sp.
Stations (No of specimens): 42-2 (3), 46-7 (2), 141-10 (3), 121-11 (1), 142-5 (1),
150-6 (5), 153-7 (1)
Pseudoscalibregma ursapium BLAKE 1981
Stations (No of specimens): 42-2 (1), 46-7 (1), 131-3 (1), 80-9 (3), 133-2 (5), 151-7 (1)
Distribution: subantarctic, 1178 – 3690m
Records: Blake, 1981
Scalibregma inflatum RATHKE 1843
Stations (No of specimens): 42-2 (3), 46-7 (2), 133-2 (1), 150-6 (2)
Distribution: cosmopolitan, 7 – 3690 m
Records: Blake, 1981; Ebbe (pers. comment), Ehlers, 1900; 1901; Fauchald, 1972;
Fauvel, 1941; Hartman, 1967; 1978; Hartmann-Schröder, 1965; 1996;
Hartmann-Schröder & Rosenfeldt, 1989; 1991; Hilbig & Blake, 2006; Monro,
1930
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 56
Sclerocheilus cf. antarcticus ASHWORTH 1915
Stations (No of specimens): 80-9 (1)
Distribution: Antarcic Peninsula and Weddell Sea, 45 – 3138 m
Records: Blake, 1981
Travisia kerguelensis MCINTOSH 1885
Stations (No of specimens): 46-7 (1), 143-1 (6), 59-5 (1), 74-6 (6), 78-9 (4), 133-2 (2),
150-6 (3), 153-7 (1), 154-9 (9)
Distribution: subantarctic, 13 – 4651 m
Records: Augener, 1932; Ehlers, 1897; 1900; 1901; 1912; Fauvel, 1941; Hartman,
1953; 1966; 1967; 1978; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Knox,
1962; McIntosh, 1885; Monro, 1930; 1936; 1939; Willey, 1902
Travisia cf. lithophila KINBERG 1866
Stations (No of specimens): 154-9 (1)
Distribution: subantarctic and Southern Pacific, 22 – 3789 m
Records: Hartman, 1952; 1966; Kinberg, 1866; 1858-1910
3.2.10 Key to the Sphaerodoridae MALMGREN 1867
Three genera of Sphaerodoridae have been found, contributing 891 specimens from
eight species. Four of the species are known to science, the other four are new and
described in this study.
1 Macrotubercles seated on dorsum (Fig. 10)……………………………………...2
# Macrotubercles on a short stem (App.-Fig. 3N)……………………..Clavodorum
only C. antarcticum
2 Macrotubercles in two rows…………………………………………...Ephesiella
Body long. More than 50 segments. First chaetiger with recurved hooks……….E. antarctica
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 57
Body shorter, less than 50 segments. Recurved hooks not apparent (Fig. 10A)
……………………………………………………………………………...E. hartmanae sp.n.
# Macrotubercles in at least four rows……………………………Sphaerodoropsis Two pairs of lateral antennae. Tentacular cirri and antennae long. Macrotubercles large and
distinctly spherical (Fig. 10E)…………………………………………………S. distincta sp.n. Three pairs of lateral antennae. Macrotubercles cone-shaped. Surface speckled with purple to
black pigments (Fig. 10H)……………………………………………………S. maculata sp.n.
Three pairs of lateral antennae. Macrotubercles well developed………………………...S. parva
With 7 – 9 rows of macrotubercles…………………………………………….S. polypapillata Two pairs of lateral antennae. Macrotubercles somewhat rectangular, little pronounced
(Fig. 10K)……………………………………………………………………….S. simplex sp.n.
Annotated species list
Clavodorum antarcticum HARTMANN-SCHRÖDER & ROSENFELDT 1990
Stations (No of specimens): 46-7 (1), 142-5 (1), 150-6 (1)
Distribution: Antarctic Peninsula and Weddell Sea, 142 – 3404 m
Records: Hartmann-Schröder & Rosenfeldt, 1990; 1992
Ephesiella antarctica (MCINTOSH 1885)
Stations (No of specimens): 42-2 (1), 46-7 (5), 143-1 (8), 142-5 (1), 153-7 (2)
Distribution: subantarctic, 220 – 3690 m
Records: Ehlers, 1912; 1913; Hartman,1964; 1978; Hartmann-Schröder & Rosenfeldt,
1992; McIntosh, 1885; Monro, 1930; 1936; 1939
Ephesiella hartmanae sp.n.
Stations (No of specimens): 143-1 (3), 16-10 (1), 74-6 (2), 80-9 (1), 133-2 (3), 150-6 (1)
Sphaerodoropsis distincta sp.n.
Stations (No of specimens): 46-7 (6)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 58
Sphaerodoropsis maculata sp.n.
Stations (No of specimens): 74-6 (23)
Sphaerodoropsis parva (EHLERS 1913)
Stations (No of specimens): 41-3 (2), 42-2 (82), 46-7 (333), 141-10 (21), 143-1 (11),
16-10 (1), 21-7 (1), 59-5 (6), 74-6 (87), 78-9 (3), 80-9 (6), 81-8 (28),
133-2 (189), 150-6 (2), 151-7 (11), 152-6 (1), 153-7 (3), 154-9 (2)
Distribution: Southern hemisphere, 0 – 4758 m
Records: Ehlers, 1913; Hartman, 1953; 1964; 1967; 1978; Hartmann-Schröder, 1965;
Hartmann-Schröder & Rosenfeldt, 1988; 1990; 1992; Wesenberg-Lund, 1961
Sphaerodoropsis polypapillata HARTMANN-SCHRÖDER & ROSENFELDT 1988
Stations (No of specimens): 42-2 (1), 131-3 (2), 80-9 (4), 121-11 (5), 153-7 (2)
Distribution: South Atlantic and Atlantic sector of the Southern Ocean, 96 – 3690 m
Records: Hartmann-Schröder & Rosenfeldt, 1988; 1992
Sphaerodoropsis simplex sp.n.
Stations (No of specimens): 46-7 (22), 141-10 (1), 74-6 (3), 80-9 (1), 150-6 (1)
3.2.11 Key to the Syllidae GRUBE 1850
The Syllidae are represented by 1331 specimens. In total twenty-one species from eight
genera have been found. The identification of four species is not without doubt, one of
them is a stolon probably of Autolytus simplex EHLERS 1900 (station 46-7). Six species
are new to science.
1 Without ventral cirri……………………………………………………………...2
# With ventral cirri…………………………………………………………………3
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 59
2 Two pairs of tentacular cirri, three antennae. Each chaetiger with a ciliary band.
Simple chaetae delicate, thinner than composite ones………………….Autolytus
only A. gibber
# Two pairs of tentacular cirri, three antennae. Chaetiger without ciliary bands.
Simple chaetae as robust as composite ones…………………………...Proceraea
only P. mclearanus
3 Tentacular cirri and dorsal cirri more or less smooth……………………………4
# Tentacular and dorsal cirri distinctly articulated……………………….Typosyllis Delicate species. Chaetal appendages bifid. Dorsal cirri alternate long (12 –15 articles) and short
(8 – 10 articles)………………………………………………………………………...T. hyalina Chaetae composite, unidentate falcigers. Long dorsal cirri with about 50 articles
…………………………………………………………………………………T. variegata
4 One pair of tentacular cirri……………………………………………………….5
# Two pairs of tentacular cirri……………………………………………………...7
5 Body epidermis covered with numerous fine papillae. Dorsal cirri flask shaped
(App.-Fig. 4A)……………………………………………………...Sphaerosyllis Sparsely papillated. Chaetiger 2 without dorsal cirri. Eyes black. Proventriculus in 4 –5 segments
(18 – 20 cell rows)………………………………………………………S. antarctica Sparsely papillated. Chaetiger 2 with dorsal cirri. Eyes red. Proventriculus in 5.5 segments (25 –
28 cell rows)……………………………………………………………………...S. joinvillensis Strongly papillated. Chaetiger 2 without dorsal cirri. Proventriculus in about 3 segments (13 – 17
cell rows)……………………………………………………………..S. lateropapillata uteae
# Epidermis without numerous papillae. Dorsal cirri variable in size and shape
…………………………………………………………………………………...6
6 Dorsal cirri long, slender. Eversible pharynx unarmed…………………Braniella
only B. palpata
# Dorsal cirri short, papilliform. Eversible pharynx with a single tooth…..Exogone Proventriculus in 3.5 – 4.5 segments (13 – 16 cell rows). Modified chaetae with triangular
blades. Two pairs of eyes. Lateral and median antennae similar……………….E. heterosetosa
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 60
Proventriculus in 3 segments. Long and short falcigers present with serrated blades (composite,
plus some simple in median body region). Median antenna prolonged…………..E. minuscula
Proventriculus in 1.5 segments (12 cell rows). Chaetal blades serrated, bidentate….E. sp. 2BH Proventriculus in 2 – 3 segments (~ 14 cell rows). Two pairs of eyes close to each other. Three
antennae similar in shape, in a narrow row between the eyes……………………….E. sp. 5 Proventriculus in 3 – 3.5 segments (~ 17 cell rows). Median antenna spindle-shaped,
longer than lateral ones……………………………………………………………………E. sp. 6 No eyes. Lateral antennae distinctly lateral in position. Proventriculus in about six segments
…………………………………………………………………………………………….E. sp. 7
7 Small forms of few millimeters. Palps fused at least partially. Dorsal cirri long
……………………………………………………………………………..Brania
only B. sp.1BH
# Larger forms. Palps maximally fused at base……………………………………8
8 Pharynx unarmed…………………………………………………………Syllides
only S. articulosus
# Pharynx with a single tooth. Dorsal cirri cylindrical………………….Pionosyllis Prostomium posteriorly notched. Notch covered by first subsequent segment. Two pairs of eyes
(App.Fig. 4B)………………………………………………………………………….P. comosa Prostomium with deep longitudinal groove and posterior incision. These not covered. Two pairs
of eyes (App.-Fig. 4C)……………………………………………………………P. epipharynx
Prostomium neither notched nor incised. Two pairs of eyes…………………………P. maxima
Prostomium with deep longitudinal groove and posterior incision. Eyes absent………….P. sp.1
Annotated species list
Autolytus gibber EHLERS 1897
Stations (No of specimens): 74-6 (1)
Distribution: subantarctic and Southern Pacific, 20 – 1047 m
Records: Ehlers, 1897; 1901; Fauvel, 1936; 1951; Gravier, 1906; 1907; Hartman, 1954;
1964; Hartmann-Schröder & Rosenfeldt, 1992; Monro, 1930; 1936
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 61
Brania sp. 1BH
Stations (No of specimens): 131-3 (2), 132-2 (1)
Braniella palpata HARTMAN 1967
Stations (No of specimens): 141-10 (2), 21-7 (1), 59-5 (1), 74-6 (3), 80-9 (2), 81-8 (1),
121-11 (2), 133-2 (32)
Distribution: cosmopolitan, 210 – 4419 m
Records: Ebbe (pers. comment); Hartman, 1967; 1978
Exogone heterosetosa MCINTOSH 1885
Stations (No of specimens): 42-2 (1), 74-6 (1)
Distribution: cosmopolitan, 0 – 3690 m
Records: Augener, 1912; Benham, 1921; 1927; Blankensteyn & Lana, 1986; Ehlers,
1897; 1901; 1913; Fauvel, 1916; 1936; Gravier, 1906; 1907; 1911; Hartman,
1953; 1964; Hartmann-Schröder, 1965; Hartmann-Schröder & Rosenfeldt, 1988;
1990; 1992; McIntosh, 1885; Monro, 1939; Wesenberg-Lund, 1961
Exogone minuscula HARTMAN 1953
Stations (No of specimens): 74-6 (8), 133-2 (1), 151-7 (18), 153-7 (1)
Distribution: Antarctic Peninsula and Weddell Sea, 10 – 2014 m
Records: Blankenstey & Lana, 1986; Hartman, 1953; 1964; 1967; 1978
Exogone sp. 2BH
Stations (No of specimens): 46-7 (1)
Exogone sp. 5
Stations (No of specimens): 74-6 (5)
Exogone sp. 6
Stations (No of specimens): 133-2 (16)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 62
Exogone sp. 7
Stations (No of specimens): 133-2 (42)
Pionosyllis comosa GRAVIER 1906
Stations (No of specimens): 74-6 (7)
Distribution: subantarctic, 6 – 1047 m
Records: Benham, 1921; 1927; Ehlers, 1912; 1913; Gravier, 1906; 1907; 1911; Harman,
1954; 1964; 1967; Monro, 1930
Pionosyllis epipharynx HARTMAN 1953
Stations (No of specimens): 46-7 (1), 133-3 (1), 141-10 (1), 74-6 (14), 133-2 (24),
150-6 (1), 153-7 (1), 154-9 (2)
Distribution: Weddell Sea and Antarctic Peninsula, 73 – 3784 m
Records: Hartman, 1964; 1967
Pionosyllis cf. maxima MONRO 1930
Stations (No of specimens): 74-6 (1)
Distribution: Weddell Sea, 238 – 1047 m
Records: Hartman, 1964; Hartmann-Schröder & Rosenfeldt, 1988; Monro, 1930
Pionosyllis sp. 1
Stations (No of specimens): 141-10 (1)
Proceraea cf. mclearanus (MCINTOSH 1885)
Stations (No of specimens): 74-6 (1)
Distribution: subantarctic and Southern Pacific, 18 – 1047 m
Records: Benham, 1927; Ehlers, 1912; 1913; Hartman, 1964; Hartmann-Schröder &
Rosenfeldt, 1990; 1992; McIntosh, 1885
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 63
Sphaerosyllis antarcticus GRAVIER 1907
Stations (No of specimens): 74-6 (1)
Distribution: Weddell Sea, 50 – 1047 m
Records: Hartmann-Schröder & Rosenfeldt, 1988; 1990; 1992
Sphaerosyllis joinvillensis HARTMANN-SCHRÖDER & ROSENFELDT 1988
Stations (No of specimens): 141-10 (1), 74-6 (12), 133-2 (285)
Distribution: Weddell Sea, 30 – 2313 m
Records: Hartmann-Schröder & Rsoenfeldt, 1988; 1992; San Martín & Parapar, 1996
Sphaerosyllis lateropapillata uteae HARTMANN-SCHRÖDER & ROSENFELDT 1988
Stations (No of specimens): 141-10 (12), 143-1 (1), 74-6 (156), 133-2 (217), 150-6 (5),
151-7 (4), 153-7 (38)
Distribution: Weddell Sea, 68 – 2313 m
Records: Hartmann-Schröder & Rosenfeldt, 1988; 1992
Syllides articulosus EHLERS 1897
Stations (No of specimens): 74-6 (1), 133-2 (317)
Distribution: subantarctic, 0 – 1582 m
Records: Augener, 1932; Blankensteyn & Lana, 1986; Ehlers, 1879, 1901; 1912; 1913;
Fauvel, 1916; Hartman, 1953; 1964; 1967; 1978; Hartmann-Schröder &
Rosenfeldt, 1988; 1990; 1992; Monro, 1939
Typosyllis cf. hyalina (GRUBE 1863)
Stations (No of specimens): 143-1 (8), 74-6 (1), 133-2 (5)
Distribution: cosmopolitan, 0 – 1582 m
Records: Ehlers, 1879; 1901; Fauvel, 1936; Gravier, 1911; Hartman, 1964; Hartmann-
Schröder, 1996; Hartmann-Schröder & Rosenfeldt, 1992; Licher, 1999; Willey,
1902
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 64
Typosyllis variegata (GRUBE 1860)
Stations ( no specimens): 143-1 (4), 133-2 (60)
Distribution: cosmopolitan, 0 – 1582 m
Records: Ehlers, 1900; 1901; Hartman, 1953; 1964; 1967; Hartmann-Schröder, 1996;
Hartmann-Schröder & Rosendfeldt, 1992, Licher, 1999; Monro, 1930;
Wesenberg-Lund, 1961
3.2.12 Key to the Terebellidae MALMGREN 1867
For the Terebellidae 683 specimens belonging to 32 species from at least 13 genera
have been found. Due to a rather poor condition of most specimens the identity of seven
species remains unclear. Four new species have been found.
1 Thoracic uncini in one row throughout…………………………………………..2
# Thoracic uncini at least partially in two rows……………………………………6
2 Branchiae present………………………………………………………………...3
# Branchiae absent…………………………………………………………………4
3 Three pairs of branchiae from segments 2 – 4. Notochaetae from first branchial
segment (segment 2), uncini from 4th chaetiger……………………..Streblosoma
only S. variouncinatum
# Three pairs of branchiae from segmenst 2 – 4. Notochaetae from second
branchial segment (segment 3), uncini from 3rd chaetiger………………Thelepus
only T. cincinnatus
4 Chaetae present…………………………………………………………………..5
# Chaetae absent. Ten thoracic segments……………………………….Hauchiella
only H. tribullata
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 65
5 Notochaetae from third segment, neurochaetae (uncini) absent…………...Lysilla
only L. sp. 1BH
# Notochaetae from third segment, uncini from chaetigers 7 – 12………Polycirrus
11 thoracic chaetigers, 12 abdominal uncinigers…………………………………P. antarcticus 11 thoracic setigers, 14 abdominal uncinigers. Notochaetae of one kind, slightly limbate, blade
weakly denticulated…………………………………………………………………...P. insignis
Notochaetae of two kinds. Long slender ones smooth, shorter ones with serrated knob...P. sp. 1
Notochaetae of two kinds. Short ones limbate with broad wings………………………...P. sp. 2
6 Branchiae present………………………………………………………………...7
# Branchiae absent…………………………………………………………………9
7 Two pairs of branchiae…………………………………………………………..8
# Three pairs of branchiae. Lateral lappets present. 17 thoracic segments
…………………………………………………………………………Thelepides
Branchial filaments in sessile bundles.….....................................................................T. koehleri
Branchiae as single filaments………………………………………………………...T. venustus
8 17 thoracic chaetigers. Branchiae smooth. Lateral lappets on segments 2 and 3.
Some anterior uncini with prolonged shafts……………………………Eupistella Two pairs of branchiae on segments 2 and 3, both with single filaments (App.-Fig. 4D) …………………………………………………………………………...E. grubei Two pairs of branchiae on segments 2 and 3. Second pair with two filaments (App.-Fig. 4E)
…………………………………………………………………………………………….E. sp. 1
# Number of thoracic chaetigers varies from 15 – 24. Branchiae stalked. Anterior
thoracic uncini long-handed (App.-Fig. 4F)…………………………………Pista Without lateral lappets on segment 2. Narrow lateral lappets on segment 4. Two pairs of
branchiae on segments 2 and 3……………………………………………………..P. corrientis
With lateral lappets on segment 2. Two pairs of branchiae on segments 2 and 3…….P. cristata Without lateral lappets on segment 2. Lappets on segment 1 greatly enlarged. One pair of
branchiae on segment 2……………………………………………………………...P. spinifera
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 66
9 Lateral lappets present………………………………………………………….10
# Lateral lappets absent…………………………………………………………...11
10 Notochaetae distally smooth or dentate. Uncini from segment 3. Segments
dorsally without ridges……………………………………………………Proclea
only P. graffii
# Third segment dorsally with transverse ridge……………………………..Leaena Thorax with 10 – 11 chaetigers. Uncini in double rows through seventh abdominal segment
……………………………………………………………………….L. antarctica 16 thoracic chaetigers. Second segment dorsally short and laterally enlarged, projecting forward
to form a ventral hood………………………………………………………………L. arenilega 17 thoracic chaetigers. Segment 3 with low dorsal ridge. Uncini in double rows through last
thoracic segment………………………………………………………………………L. collaris 15 thoracic chaetigers. Segments 2 and 3 with large lateral lappets and short digitiform processes
that remind of branchiae. Uncini in double rows through first abdominal segment ………………………………………………………………………L. pseudobranchiata 15 thoracic chaetigers. First ventral segment with ventral collar. Segments 2 and 3 with lateral
lappets. Uncini in double rows from chaetigers 7 – 15………………………….L. wandelensis
17 chaetigers. Segment 4 with lateral lappets partially covering segment 3……………...L. sp. 4
11 17 thoracic segments. Notochaetae smooth, uncini from chaetiger 7….Laphania
only L. boecki
# 14 thoracic segments. Notochaetae distally denticulated, uncini from chaetiger 2
……………………………………………………………………………Phisidia
14 thoracic chaetigers…………………………………………………………...P. rubrolineata
16 thoracic chaetigers. Short chaetae with hirsute blades………………………………...P. sp. 1
Annotated species list
Eupistella grubei (MCINTOSH 1885)
Stations (No of specimens): 74-6 (9), 80-9 (3), 133-2 (4)
Distribution: subantarctic, 430 – 4770 m
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 67
Records: Hartman, 1966; 1978; Levenstein, 1964; McIntosh, 1885
Eupistella sp. 1
Stations (No of specimens): 16-10 (2)
Hauchiella tribullata (MCINTOSH 1869)
Stations (No of specimens): 42-2 (1), 74-6 (2)
Distribution: cosmopolitan, 17 – 3690 m
Records: Hartman, 1966; 1978; Hartmann-Schröder, 1996; Hartmann-Schröder &
Rosenfeldt, 1989; 1991; Hessle, 1917; Holthe, 1986; Monro, 1930; 1936
Laphania cf. boecki MALMGREN 1866
Stations (No of specimens): 46-7 (2)
Distribution: cosmopolitan, 12 – 2889 m
Records: Hessle, 1917
Leaena antarctica MCINTOSH 1885
Stations (No of specimens): 81-8 (1), 102-13 (1), 133-2 (2), 153-7 (7)
Distribution: subantarctic, 13 – 4419 m
Records: Benham, 1927; Ehlers, 1897; 1900; 1901; 1913; Hartman, 1966; Hartmann-
Schröder & Rosenfeldt, 1989; 1991; Hessle, 1917; Levenstein, 1964; McIntosh,
1885; Monro, 1930; 1936
Leaena arenilega EHLERS 1913
Stations (No of specimens): 74-6 (5), 153-7 (1)
Distribution: subantarctic, 27 – 2014 m
Records: Benham, 1921; Ehlers, 1913; Hartman, 1966; 1978; Hartmann-Schröder &
Rosenfeldt, 1991; Hessle, 1917
Leaena collaris HESSLE 1917
Stations (No of specimens): 74-6 (2), 133-2 (64)
Distribution: subantarctic and Southern Atlantic, 50 – 1582 m
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 68
Records: Hartman, 1966; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Hessle, 1917;
Monro, 1930; 1936
Leaena cf. collaris HESSLE 1917
Stations (No of specimens): 16-10 (1), 74-6 (3)
Leaena pseudobranchiata LEVENSTEIN 1964
Stations (No of specimens): 74-6 (3)
Distribution: subantarctic, 206 – 1047 m
Records: Hartman, 1966; Levenstein, 1964
Leaena wandelensis GRAVIER 1907
Stations (No of specimens): 21-7 (1)
Distribution: subantarctic, 32 – 4574 m
Records: Benham, 1927; Gravier, 1907; 1911; Hartman, 1952; 1966; Levenstein, 1964
Leaena sp. 4
Stations (No of specimens): 74-6 (1), 154-9 (3)
Lysilla sp. 1BH
Stations (No of specimens): 46-7 (1)
Phisidia rubrolineata HARTMANN-SCHRÖDER & ROSENFELDT 1989
Stations (No of specimens): 133-2 (266)
Distribution: Weddell Sea, 120 – 1582 m
Records: Hartmann-Schröder & Rosenfeldt, 1989; 1991
Phisidia sp. 1BH
Stations (No of specimens): 150-6 (1)
Pista corrientis MCINTOSH 1885
Stations (No of specimens): 74-6 (1)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 69
Distribution: subantarctic and Southern Atlantic, 15 – 1047 m
Records: Benham, 1927; Ehlers, 1913; Fauvel, 1951; Hartman, 1952; 1966; 1967;
Hartmann-Schröder & Rosenfeldt, 1989; Hessle, 1917; Levenstein, 1964;
McIntosh, 1885; Monro, 1930; 1939
Pista cristata (MÜLLER 1776)
Stations (No of specimens): 74-6 (2)
Distribution: cosmopolitan, 0 – 1784 m
Records: Augener, 1932; Ehlers, 1900; 1901; Gravier, 1907; 1911; Hartman, 1966;
1967; Hartmann-Schröder, 1996; Hessle, 1917; Levenstein, 1964
Pista spinifera (EHLERS 1908)
Stations (No of specimens): 74-6 (5)
Distribution: subantarctic, 73 – 4758 m
Records: Augener, 1932; Ehlers, 1908; 1913; Gravier, 1911; Hartman, 1966; 1967
Polycirrus cf. antarcticus (WILLEY 1902)
Stations (No of specimens): 74-6 (1)
Distribution: subantarctic, 1047 m
Records: Hartman, 1966; Willey, 1902
Polycirrus insignis GRAVIER 1907
Stations (No of specimens): 46-7 (1), 143-1 (1) 74-6 (13), 78-9 (3), 133-2 (241),
142-5 (2), 150-6 (1), 154-9 (1)
Distribution: subantarctic, 40 – 3784 m
Records: Fauvel, 1951; Gravier, 1907; Hartmann, 1966; Hartmann-Schröder &
Rosenfeldt, 1989; Hessle, 1917
Polycirrus sp. 1
Stations (No of specimens): 59-5 (1)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 70
Polycirrus sp. 2
Stations (No of specimens): 74-6 (1)
Streblosoma variouncinatum HARTMANN-SCHRÖDER & ROSENFELDT 1991
Stations (No of specimens): 141-10 (2)
Distribution: Weddell Sea and Drake Passage, 134 – 2313 m
Records: Hartmann-Schröder, 1991
Thelepides koehleri GRAVIER 1911
Stations (No of specimens): 74-6 (2)
Distribution: Weddell Sea, 36 – 3020 m
Records: Gravier, 1911; Hartman, 1966; 1967; 1978; Hartmann-Schröder & Rosenfeldt,
1989; 1991
Thelepides venustus LEVENSTEIN 1964
Stations (No of specimens): 74-6 (3)
Distribution: subantarctic, 197 – 1047 m
Records: Hartman, 1966; Levenstein, 1964
cf. Thelepides venustus LEVENSTEIN 1964
Stations (No of specimens): 74-6 (1)
cf. Thelepus cincinnatus (FABRICIUS 1880)
Stations (No of specimens): 74-6 (1)
Distribution: cosmopolitan, 15 – 3239 m
Records: Augener, 1932; Benham, 1927; Fauvel, 136; 1951; Hartman, 1952; 1966;
1967; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Hessle, 1917; Levenstein,
1964; Monro, 1930; 1939; Willey, 1902
Thelepodinae sp. 1
Stations (No of specimens): 46-7 (7), 74-6 (1)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 71
Terebellidae sp. 1
Stations (No of specimens): 131-3 (1)
Terebellidae sp. 2
Stations (No of specimens): 46-7 (1)
Terebellidae sp. 3
Stations (No of specimens): 21-7 (1)
Terebellidae sp. 4
Stations (No of specimens): 46-7 (1)
3.2.13 Key to the Trichobranchidae MALMGREN 1866
Five species of Trichobranchidae have been found. Two genera could be identified
without doubt, the taxonomy of two species remains unresolved. A total of 211
specimens were collected.
1 Four pairs of lanceolate branchiae. 16 thoracic chaetigers, uncini from chaetiger
4……………………………………………………………………Octobranchus
only O. antarcticus
# One single branchia of four lamellate branchial lobes (App.-Fig. 4G). 18 thoracic
chaetigers, uncini from chaetiger 6…………………………………..Terebellides
Branchial lobes basally fused………………………………………………………….T. stroemi
Branchial lobes basally free…………………………………………………………..T. sp. 1BH
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 72
Annotated species list
Octobranchus antarcticus MONRO 1936
Stations (No of specimens): 46-7 (1), 141-10 (1), 74-6 (19), 80-9 (4), 133-2 (119),
153-7 (1)
Distribution: Weddell Sea and Antarctic Peninsula, 267 – 3138 m
Records: Hartman, 1966; 1967; Monro, 1936
Terebellides stroemi SARS 1835
Stations (No of specimens): 42-2 (1), 46-7 (2), 131-3 (1), 16-10 (1), 59-5 (1), 74-6 (6),
78-9 (12), 80-9 (5), 81-8 (2), 94-14 (2), 102-13 (1), 110-8 (3), 121-11 (15),
133-2 (1), 142-5 (6), 150-6 (4)
Distribution: cosmopolitan, 18 – 4817 m
Records: Augener, 1932; Ehlers, 1897; 1900; 1908; Fauchald, 1972; Hartman, 1966;
Hartmann-Schröder, 1996; Hessle, 1917; Holthe, 1986
cf. Terebellides sp. 1BH
Stations (No of specimens): 80-9 (1)
Trichobranchidae sp. 1
Stations (No of specimens): 46-7 (1)
Trichobranchidae sp. 2
Stations (No of specimens): 141-10 (1)
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 73
3.3 Descriptions of new species
3.3.1 Ampharetidae MALMGREN 1866
3.3.1.1 Anobothrus pseudoampharete sp.n.
Genus Anobothrus LEVINSEN 1884
Anobothrus pseudoampharete sp.n.
Holotype. ANDEEP III, eastern Weddell Sea, st. 74-6, 20 February 2005, 71°18.42’ S,
13°58.21’ W, 1047 m, EBS, ZMH P-24741
Paratypes. ANDEEP III, eastern Weddell Sea, st. 74-6, 20 February 2005, 71°18.42’ S,
13°58.21’ W, 1047 m, EBS; 50 specimens, ZMH P-24742
Etymology. The name refers to the strong reminiscence of this species to Ampharete
kerguelensis at first sight
Diagnosis. The species can be recognized by the palae which are wide at the base and
then abruptly tapering to a long, delicate tip.
Description
Holotype complete except for lack of branchiae, 5 mm long and 0.5 mm wide for 30
chaetigers.
A species of median size, between 3 – 13 mm long. Body long, gradually tapering to the
posterior end (Fig. 4A). Color in alcohol light tan to white.
Prostomium slightly scoop-shaped, fused to peristomium and anterior segments. 15
thoracic chaetigers present, first bearig pairwise whirls of 15 – 20 palae. Palae stout and
broad at base suddenly tapering to a delicate tip (Fig. 4B). First two subsequent
segments only with notopodia. Twelve thoracic uncinigers and up to 15 abdominal
uncinigers. Fifth-to-last thoracic unciniger with elevated parapodia connected by a
dorsal ridge, with modified chaetae and uncinigers (Fig. 4E, G).
Thoracic chaetae limbate, of two sizes (Fig. 4D). Limbate chaetae of fifth to last
chaetiger of three sizes plus some simple capillaries (Fig. 4E). Thoracic uncini with
three rows of small teeth, laterally kidney shaped (Fig. 4F). Those of fifth-to last
chaetiger also with three rows of teeth, more or less round in lateral view (Fig. 4G).
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 74
Abdominal uncinigers lacking notopodia, neuropodia with uncini with four rows of
small teeth, one row of three teeth and one median main tooth (Fig. 4H).
Pygidium with uneven margin, cirri lacking, anus terminal (Fig. 4A).
Eight pairs of branchiae present, arranged in a row. Branchiae rather robust, digitiform
(Fig. 4C), reaching back to about third chaetiger.
Remarks. Only one further species of the genus Anobothrus LEVINSEN 1884 is known
for the Southern Ocean to date. This species, A. gracilis (MALMGREN 1866), bears very
long palae that gradually taper in width from base to tip. The palae of A.
pseudoampharete n.sp. in contrast are shorter and very wide in their complete basal
half. They suddenly taper to a fine, long tip in their distal half.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 75
Fig. 4: Anobothrus pseudoampharete sp.n. A- lateral view, B- palae, C- branchiae, D- thoracic
notochaetae, E- notochaetae 5th-to-last thoracic segment, F- thoracic uncinus, G- thoracic uncinua 5th-to-
last segment, lateral and frontal view, H- abdominal uncini, lateral and frontal view
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 76
3.3.2 Hesionidae GRUBE 1850
3.3.2.1 Amphiduros serratus sp.n.
Genus Amphiduros HARTMAN 1959
Amphiduros serratus sp.n.
Holotype. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 132-2, 06 March
2002, 65°17.74'S, 53°22.82'W, 2084-2086 m, EBS, ZMH P-24743
Paratypes. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 132-2, 06 March
2002, 65°17.74'S, 53°22.82'W, 2084-2086 m, EBS, 1 specimens, ZMH P-24744
Additional material. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 134-4, 09
March 2002, 5°19.20'S, 48°3.81'W, 4066-4069 m, EBS, 1 specimens
Etymology. The name refers to the serration of the notopodial chaetae
Diagnosis. Distinguishing characters of this species are the types of the notopodial
(chambered and serrated) and neuropodial (composite, with chambered shafts) chaetae.
Description
Holotype incomplete, 3.0 mm long and 1.0 mm wide for 10 chaetigers. Antennae lost.
Color in alcohol white.Prostomium about as long as wide. Palps biarticulated with
terminal article about twice as long as basal. Antennae short and delicate throughout,
median antenna attached medially on prostomium. Peristomium dorsally reduced,
laterally visible. Eyes absent.
Tentactular cirri eight pairs, arranged as 2-2-2-2, all articulated. Length about that of
body width (Fig. 5A).
Parapodia biramous (Fig. 5B). Notopodium a short massive lobe, neuropodium longer,
with a somewhat pointed tip. Dorsal cirri articulated, long, similar to tentacular cirri.
Ventral cirri smooth, shorter, clearly extending tip of neuropodia.
Notochaetae simple, chambered and serrated at distal margin (Fig. 5C). Neurochaetae
composite falcigers with chambered shafts and long appendages with very delicate tips
(Fig. 5D).
Remarks. Amphiduros serratus sp.n. is the second species of the genus. It can clearly be
distinguished by the absence of eyes and the form of the antennae. Amphiduros
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 77
fuscescens (MARENZELLER 1875) in contrast has large eyes and strongly tapering
antennae with delicate tips (Pleijel, 2001).
3.3.2.2 Micropodarke cylindripalpata sp.n.
Genus Micropodarke OKUDA 1938
Micropodarke cylindripalpata sp.n.
Holotype. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 46-7, 30
January 2002, 50°38.35’S, 53°57.36’W, 2889 m, EBS, ZMH P-24745
Paratypes. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 46-7, 30
January 2002, 50°38.35’S, 53°57.36’W, 2889 m, EBS, 2 specimens, ZMH P-24746
Additional material. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 42-2,
27 January 2002, 59°40.29’S, 57°35.43’W, 3683-3690 m, EBS, 9 specimens
Etymology. The name refers to the presence of pin-shaped biarticulated palps
Diagnosis. The species can be recognized by the pin-shaped form of the palps in
combination with the button-shaped tip of the parapodial lobes.
Description
Holotype incomplete, 3.0 mm long and 1.0 mm wide for 14 chaetigers. Color in alcohol
white.
Prostomium oval in shape, slightly wider than long. Palps biarticulated and pin-shaped.
Two antennae, attached frontodorsally between palps. Peristomium indistinct, dorsally
reduced. Everted pharynx with eleven distal papillae. Eyes absent (Fig. 6A).
Body segmentation sutures indistinct, segmentation apparent due to parapodia
arrangement. Six pairs of tentactular cirri on three segments, arranged in 2-2-2. Cirri
smooth, length about half the width of body.
Parapodia uniramous (Fig. 6B). Parapodial lobe robust with somewhat button-shaped
tip. Dorsal cirri resembling tentacular cirri in size and shape, ventral cirri similar,
slightly shorter.
Chaetae all composite falcigers with long appendages with very delicate tips (Fig. 6C).
Chaetae arranged in dense fans.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 78
Remarks. The species is the only species of the genus Micropodarke OKUDA 1938
described for the Southern Oceans so far and is thus easily recognized. The other only
known species of this genus, M. dubia (HESSLE 1925), is only reported from temperate
and tropic waters of the Pacific and Indic. Distinguishing characters of the species are
the shape of the palpal articles and chaetal appendages.
3.3.2.3 Ophiodromus calligocervix sp.n.
Genus Ophiodromus SARS 1862
Ophiodromus calligocervix sp.n.
Holotype. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 132-2, 06 March
2002, 65°17.74'S, 53°22.82'W, 2084-2086 m, EBS, ZMH P-24747
Paratypes. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 132-2, 06 March
2002, 65°17.74'S, 53°22.82'W, 2084-2086 m, EBS, 1 specimens, ZMH P-24748
Additional material. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 134-4, 09
March 2002, 5°19.20'S, 48°3.81'W, 4066-4069 m, EBS, 6 specimens
Etymology. The name refers to the distinct dark coloring of the posterior end of the
prostomium
Diagnosis. The species can be recognized by the distinct coloring of the posterior
margin of the prostomium that is mostly present in ethanol preservation. The round
shape of the prostomium is also characteristic, even when antennae and tentacular cirri
are lost.
Description
Holotype incomplete, 1.7 mm long and 0.5 mm wide for eight chaetigers. Color in
alcohol white.
Promstomium about as long as wide, almost round in shape. Three antennae, median
one attached mediofrontally on prostomium. Palps biarticulated with basal article large
and stout, terminal article of about equal length but more narrow. Peristomium well
developed, dorsally reduced, laterally visible. Eyes absent (Fig. 5E). Boarder part
between prostomium and peristomium distinctly colored, light brown in alcohol.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 79
Six pairs of tentacular cirri, length about as body width, robust, often lost, on one to two
visible segments.
First chaetae in segment 2; parapodia biramous. Notopodia increasing in size from
anterior to posterior. Neuropodia with a somewhat pointed tip (Fig. 5F).
Dorsal and ventral cirri smooth, tapering to apex. Dorsal cirri about 1.5 times the length
of neuropodia; ventral cirri shorter than neuropodia.
Notochaetae simple capillaries, interiorly chambered (Fig. 5G). Neurochaetae
composite falcigers; shaft of chaetae chambered, appendage long, about half as long as
shaft, with very delicate tip (Fig. 5H).
Remarks. The species seems very common and is easily recognized due to a dark
transverse pigmentation on the dorsoposterior margin of the prostomium. In contrast to
O. comatus (EHLERS 1912) which is found in the Southern Ocean deep sea as well, the
species lacks eyes. Also the dorsal and ventral cirri are not as distinctly articulated in O.
calligocervix n.sp. as in O. comatus.
3.3.2.4 Parasyllidea delicata sp.n.
Genus Parasyllidea PETTIBONE 1961
Parasyllidea delicata sp.n.
Holotype. ANDEEP III, Weddell Sea, Antarctic Peninsula, Sta. 142-5, 18 March 2005,
62°11.36'S, 49°27.62'W, 3403 m, EBS, ZMH P-24749
Paratypes. ANDEEP III, Weddell Sea, Antarctic Peninsula, Sta. 142-5, 18 March 2005,
62°11.36'S, 49°27.62'W, 3403 m, EBS, 2 specimens, ZMH P-24750
Additional material. ANDEEP III, Weddell Sea, South Orkney Islands, Sta. 150-6, 20
March 2005, 61°49.13'S, 47°27.51'W, 1970 m, EBS, 5 specimens
Etymology. The name refers to the delicate structure of the tentacular and dorsal cirri
Diagnosis. The species can be recognized by its very delicate and long tentacular and
dorsal cirri and the lack of eyes.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 80
Description
Holotype incomplete. Complete paratype of 29 chaetigers, 8 mm long and 1 mm wide.
Color in alcohol white.
Prostomium trapezoid, about as long as wide. Two antennae present, inserted frontally
between the palps, very thin, about as long as first palpal article. Palps biarticulated with
both articles long and slender. Pharynx smooth with fimbriated margin. Peristomium a
slender ring, dorsally reduced. Eyes absent (Fig. 6D).
Six pairs of tentacular cirri arranged in 2-2-2. First five pairs slender and delicate, about
as long as body width. Third dorsal pair more robust, even longer and indistinctly
articulated.
Parapodia uniramous, increasing in size towards middle of body, decreasing again
towards pygidium. One acicula present ending in a stout tip. Dorsal and ventral cirri
similar in shape, slender, dorsal cirri about 1.5 times as long as ventral cirri (Fig. 6F).
Chaetae all composite falcigers in a dense fan. Appendages shortest at margins of
parapodial lobe, increasing in size towards median tip of lobe (Fig. 6G).
Pygidium a smooth ring, pointing terminally (Fig. 6E).
Remarks. Parasyllidea delicata sp.n. is the only species of the genus Parasyllidea
PETTIBONE 1961 described for the Southern Ocean so far. It is found in deep waters
below 1000 m depth in contrast to P. australiensis HARTMANN-SCHRÖDER 1980, which
is described for the Australian coast’s sublitoral. The only other known species and type
species of this genus, P. humesi PETTIBONE 1961 is a commensale species associated
with the bivalve Tellina nymphalis (LAMARCK 1818) also known for shallow waters
(Pettibone, 1961).
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 81
Fig. 5: A-D Amphiduros serratus sp.n. A-anterior segments, B- parapodium, C- notochaeta, D-
neurochaeta. E-H Ophiodromus calligocervix sp.n. E-anterior segments, F- parapodium, G- notochaeta,
H- neurochaeta
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 82
Fig. 6: A-C Micropodarke cylindripalpata sp.n. A-anterior segments, B- parapodium, C- neurochaeta,
D-G Parasyllidea delicata sp.n. D-anterior segments, E- posterior segments, F- parapodium, G-
neurochaeta
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 83
3.3.3 Opheliidae MALMGREN 1867
3.3.3.1 Ammotrypanella MCINTOSH 1879
Genus Ammotrypanella MCINTOSH 1879
Typespecies: A. arctica MCINTOSH 1879
Generic rediagnosis
Body long and thin. Ventral groove along whole length of body. Prostomium bluntly
rounded to conical with small palpode, peristomium indistinct. Eyes absent.
Anterior parapodia button shaped with long, bent chaetae arranged in tufts. Parapodia
embedded into lateral groove in median region, becoming more distinct in posterior
region. Parapodia with branchiae in third quarter of body. Posterior chaetae longer than
median ones, stiff and straight. All chaetae simple.
Branchiae flat, wide at base, tapering to top.
Posterior end of body laterally flattened with chaetigers reduced in length. Pygidium
with or without anal tube, this with or without ventral cirrus.
3.3.3.1.1 Ammotrypanella arctica MCINTOSH 1879
Ammotrypanella arctica MCINTOSH 1879
Holotype. Davis Strait, Greenland (1785 fms), Globigerina – ooze, H.M.S. Valerous,
Sta. 16, 55°10’N, 25°58’W, 1875 (NHM, 1921.5.1.2392)
Additional material. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 42-2,
27 January 2002, 59°40.29’S, 57°35.43’W, 3683-3690 m, EBS, 2 specimens, Weddell
Sea, Antarctic Peninsula, Sta. 131-3, 05 March 2002, 65°19.83’S, 51°31.62’W, 3049-
3050 m, EBS, 7 specimens, Sta. 134-4, 09 March 2002, 65°19.20’S, 48°3.81’W, 4066-
4069 m, EBS, 1 specimen
Additional records.
Hartman & Fauchald (1971) – A66, 38°46.7’N, 70°08.8’W, 2802 m; A70, 36°23’N,
67°58’W, 4680 m; A121, 35°50’N, 65°11’W, 4800m; A125, 37°24’-37°26’N, 65°54’-
65°50’W, 4825 m; A122, 35°50’-35°52’N, 64°57.5’-64°58’W, 4833 m; A124, 37°26’-
37°25’N, 63°59.5-63°58’W, 4862 m; A120, 34°43’-34°40.5’N, 66°32.8-66°35’W,
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 84
5018-5023 m; Ch84, 36°24.4’N, 67°56’W, 4749 m; Ch100, 33°56.8’N, 65°47’W, 4743-
4892 m
Diagnosis. The species can be recognized by the characters of the anal tube which are
the lack of a ventral cirrus and the presence of numerous short cirri on the posterior
margin.
Redescription
Holotype incomplete, body long and thin, 15 mm long and 1 mm wide for 42
chaetigers.
A moderately large species of 9 – 33 mm length and 0.5 – 2 mm; number of chaetigers
37 – 43. With ventral groove along whole length of body. Posterior body region
laterally flattened. Color in alcohol white to a light tan.
Prostomium conical with a short palpode. Peristomium a thin ring around the lips.
Nuchal organs apparent as narrow lateral grooves, eyes absent (Fig. 7C-D).
First following segment without chaetae. Anterior 8 – 10 parapodia button shaped with
long, stiff chaetae in bushy fascicles . Following parapodia embedded in lateral groove.
Posterior parapodia more distinct with long, straight chaetae. All chaetae simple.
Branchiae can be present in chaetigers 22 – 32, all branchiae of similar length, flat with
wide base, tapering to top.
Posterior abranchiate chaetigers reduced in length, close to each other. Pygidium with
anal tube; length of anal tube about that of last 5 – 8 chaetigers; posterior margin with
numerous short cirri, without ventral cirrus (Fig. 7E).
Remarks. A. arctica is the type species of the genus. The holotype is of poor condition,
the diagnostic characters, however, are still present. Therefore it is without doubt that
the newly found specimens from the Southern Ocean belong to this species.
3.3.3.1.2 Ammotrypanella cirrosa sp.n.
Ammotrypanella cirrosa sp.n.
Holotype. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 131-3, 05 March
2002, 65°19.83’S, 51°31.62’W, 3049-3050 m, EBS, ZMH P-24751
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 85
Paratypes. ANDEEP I – II; Weddell Sea, Antarctic Peninsula, Sta. 131-3, 05 March
2002, 65°19.83’S, 51°31.62’W, 3049-3050 m, EBS, 63 specimens, ZMH P-24752
Additional material. Sta. 134-4, 09 March 2002, 65°19.20’S, 48°3.81’W, 4066-4069 m,
EBS, 2 specimens, Scotia Sea northeast off Elephant Island, Sta. 42-2, 27 January 2002,
59°40.29’S, 57°35.43’W, 3683-3690 m, EBS, 4 specimens, South Sandwich Islands,
east off Monatgu Island, Sta. 141-10, 23 March 2002, 58°25.08’S, 25°0.77’W, 2258-
2313 m, EBS, 1 specimen
Etymology. The name refers to the presence of numerous small cirri and a large ventral
cirrus on the posterior margin of the anal tube.
Diagnosis. The species can be recognized by the presence of a thick ventral cirrus and
numerous small cirri on the posterior margin of the anal tube.
Description
Holotype complete, body long and thin, 16 mm long and 1.5 mm wide for 40 chaetigers.
A median species of 7 – 25 mm length and 0.5 – 2 mm; number of chaetigers 37 – 42.
Ventral groove present along whole length of body. Posterior body region bilaterally
flattened. Color in alcohol white to a light tan.
Prostomium conical ending in a short palpode. Peristomium indistinct, a thin ring
around the lips. First subsequent segment without chaetae. Nuchal organs apparent as
narrow grooves laterally, eyes absent (Fig. 7A).
Anterior 7 – 8 parapodia button shaped, chaetae long, stiff, in bushy fascicles. Median
parapodia embedded in lateral groove, becoming more distinct in posterior part of body;
posterior chaetae long and straight. All chaetae simple.
Branchiae flat with wide base, tapering to top; all of similar length; present in region
from chaetigers 23 – 31, exact number variable.
Posterior chaetigers shorter, close to each other. Pygidium with anal tube; length of anal
tube about that of last 5 – 8 chaetigers; posterior margin with few short cirri; ventral
cirrus robust, about half the length of anal tube (Fig. 7B).
Remarks. The species is most similar to A. arctica. The most apparent diagnostic
difference is the presence of a robust ventral cirrus on the anterior margin of the anal
tube. Other than that the anal tube is very similar to that of A. arctica and recognition of
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 86
this species becomes difficult when the ventral cirrus is lost. In that case the structure of
the branchiae can be considered, as they are less flattened and more robust than in A.
arctica.
3.3.3.1.3 Ammotrypanella mcintoshi sp.n.
Ammotrypanella mcintoshi sp.n.
Holotype. ANDEEP III, st. 16-10, Southern Atlantic, off South Africa, 26 January 2005,
41°07.55’ S, 9°55.94’E, 4720 m, EBS, ZMH P-24753
Paratypes. ANDEEP III, st. 21-7, Southern Atlantic, 29 January 2005, 47°39.87'S,
04°15.79'E, 4574 m, EBS, 1 specimens, ANDEEP III, st. 59-5, Atlantic Sector Southern
Ocean, 14 February 2005, 67°30.75'S, 00°00.23'W, 4651 m, EBS, 1 specimens, ZMH
P-24754
Etymology. The name is chosen in honour of W.C. McIntosh who defined the genus
Ammotrypanella in 1879
Diagnosis. The diagnostic character of this species is the lack of an anal tube.
Description
Holotype complete, body long and thin, 6 mm long and 0.5 mm wide for 35 chaetigers.
A small species of 5 – 16 mm length and 0.5 – 1 mm; number of chaetigers about 30 –
35 or more, very variable. With ventral and lateral groove along whole length of body.
Posterior body region bilaterally flattened. Color in alcohol white to a light tan.
Prostomium conical, bearing a short palpode. Peristomium a thin ring around the lips.
First subsequent segment apparent as a narrow ring from above, without chaetae.
Nuchal organs indistinct narrow lateral grooves, eyes absent (Fig. 8A).
Anterior 8 – 10 parapodia button shaped with long, stiff chaetae in bushy fascicles.
Following parapodia indistinct, embedded in lateral groove. Posterior parapodia more
distinct with long, straight chaetae. All chaetae simple capillaries.
Branchiae present in third quarter of body, all of similar length, flat with wide base,
tapering to top.
Posterior abranchiate chaetigers reduced in length, close to each other. Pygidium
without anal tube (Fig. 8B).
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 87
Remarks. The species is most similar to the type species of the genus Ammotrypanella
MCINTOSH 1879, A. arctica. In contrast to all other species of the genus the species
lacks an anal tube and thus can easily be recognized at first sight.
3.3.3.1.4 Ammotrypanella princessa sp.n.
Ammotrypanella princessa sp.n.
Holotype. ANDEEP I – II, Weddell Sea, Antarctic Peninsula, Sta. 134-4, 09 March
2002, 65°19.20’S, 48°3.81’W, 4066-4069 m, EBS,ZMH P-24755
Paratypes. Scotia Sea northeast off Elephant Island, Sta. 42-2, 27 January 2002,
59°40.29’S, 57°35.43’W, 3683-3690 m, EBS, 1 specimen, Weddell Sea, Antarctic
Peninsula, Sta. 135-4, 09 March 2002, 65°0.06’S, 43°1.19’W, 4678 m, EBS, 1
specimen, ZMH P-24756
Etymology. The name refers to the crown-shaped prostomium and the long, slightly
bent chaetae along the anterior and median body that give the species a noble, feminine
look.
Diagnosis. Distinguishing characters of the species are the shape of the prostomium, the
presence of relatively long chaetae in the median body region, and the smooth margin of
the anal tube in combination with the presence of a ventral cirrus.
Description
Holotype complete, body long and thin, 11 mm long and 1 mm wide for 35 chaetigers.
Length of species 5 – 11 mm, width about 02 – 1 mm; number of chaetigers 33 – 35.
With ventral groove along whole length of body. Posterior body region laterally
flattened. Color in alcohol white to a light tan.
Prostomium crown-shaped to conical, with a short palpode. Peristomium limited to the
lips. First following segment achaetous. Nuchal organs apparent as lateral grooves, eyes
absent (Fig. 7G).
Anterior eight parapodia button-shaped with prolonged, stiff chaetae in bushy fascicles.
Median parapodia less distinct, embedded in lateral groove, with few long, slightly bent
chaetae. Posterior parapodia prominent, chaetae long and straight. All chaetae simple.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 88
Branchiae present in various numbers in third quarter of body (about chaetigers 23 –30),
all of similar length, flat with wide base, tapering to top.
Posterior abranchiate chaetigers reduced in length, close to each other. Pygidium with
anal tube; about as long as last six chaetigers, opening slightly ventrally; posterior
margin smooth, with short and stout ventral cirrus (Fig. 7F).
Remarks. The species stands out by its median parapodia. These are more distinct than
in A. arctica and A. cirrosa sp.n., with long, slightly bent chaetae. Therefore, the
difference in chaetation between the anterior and median body region is less apparent.
The prostomium is not completely conical, its form rather reminds of a crown. A.
princessa sp.n. and A. cirrosa sp.n.have the ventral cirrus on the posterior margin of the
anal tube in common. The margin of the anal tube, however, is smooth in A. princessa
sp.n..
3.3.3.2 Ophelina Örsted, 1843
3.3.3.2.1 Ophelina ammotrypanella sp.n.
Ophelina ammotrypanella sp.n.
Holotype. ANDEEP I – II; Weddell Sea, Antarctic Peninsula, Sta. 131-3, 05 March
2002, 65°19.83’S, 51°31.62’W, 3049-3050 m, EBS, ZMH P-24757
Paratypes. ANDEEP III, Weddell Sea, South Orkney Islands, Sta. 150-6, 20 March
2005, 61°49.13'S, 47°27.51'W, 1970 m, EBS, 5 specimens, ZMH P-24758
Additional material. ANDEEP III, Weddell Sea, South Orkney Islands, Sta. 151-7, 21
March 2005, 61°45.67'S, 47°07.19'W, 1178 m, EBS, 2 specimens
Etymology. The name refers to the close resemblance of this species to Ammotrypanella
arctica McIntosh, 1879 which only differs from it in the lack of branchiae in anterior
and median segments.
Diagnosis. The species can be recognized by the limtiation of enlarged branchiae to the
third body quarter and an anal tube without a ventral cirrus.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 89
Description
Holotype complete, 30.5 mm long and 2 mm wide for 45 chaetigers.
A large species of about 14 – 31 mm length and 1 – 2 mm width for 32 – 45 chaetigers.
Color in alcohol a light tan.
Prostomium conical with a distinct palpode. Peristomium forming a ring around the
lips. Nuchal organs indistinct, but present. Eyes absent (Fig. 8E).
First subsequent segment achaetous. Following segments with small, reduced
parapodia. Anterior chaetigers with apparent annulation, this and segmental sutures
fading towards median body region. Segmental sutures more apparent in posterior body
region again.
Chaetae simple capillaries throughout, anterior ones long and curved, median ones
short, and posterior ones long and straight.
Branchiae present in anterior and median chaetigers, very small and indistinct in first
body half. Very large, somewhat limbate branchiae in third quarter of body, up to 15 of
them present, lacking in posterior chaetigers.
Posterior chaetigers laterally flattened and strongly decreasing in size. Pygidium with
robust anal tube. Margin of this with numerous fine cirri, ventral cirrus lacking (Fig.
8F).
Remarks. The species is most similar to O. setigera (HARTMAN 1978). Both species
share the prolonged chaetae in anterior segments and the lack of an ventral cirrus on the
anal tube. The anterior branchiae of O. ammotrypanella sp.n. are however smaller and
less distinct than that of O. setigera, at first sight the species is therefore reminiscent of
Ammotrypanella arctica.
3.3.3.2.2 Ophelina robusta sp.n.
Ophelina robusta sp.n.
Holotype. ANDEEP I – II; Weddell Sea, Antarctic Peninsula, Sta. 131-3, 05 March
2002, 65°19.83’S, 51°31.62’W, 3049-3050 m, EBS, ZMH P-24759
Paratype. ANDEEP III, Weddell Sea, Sta. 121-11, 14 March 2005, 63°38.27'S,
50°37.16'W, 2668 m, EBS, 5 specimens, ZMH P-24760
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 90
Etymology. The name refers to the presence of a very robust ventral cirrus on the anal
tube.
Diagnosis. A diagnostic character of this species is the presence of median sized
branchiae in the first 7 – 8 parapodia. Enlarged branchiae are present in the posterior
region of the body. The anal tube bears a robust ventral cirrus.
Description
Holotype complete, 18 mm long and 1.5 mm wide for 30 chaetigers.
A species of median size. Length about 18 – 23 mm, width 1 – 2 mm, number of
chaetigers between 30 – 35. Color in alcohol a light tan.
Prostomium conical, carrying a distinct palpode. Peristomium limited to the lips.
Nuchal organs present, but indistinct. Eyes absent (Fig. 8C).
First follwing segment achaetous. Subsequent segments with reduced button-shaped
parapodia. Anterior chaetigers with slight annulation. Segmental sutures indistinct in
median body region becoming more apparent in posterior body region again.
Chaetae simple capillaries throughout, anterior ones long and curved, median ones
short, and posterior ones longer but few.
Anterior 7 – 8 branchiae distinct, of median length. Very small and indistinct branchiae
in median body region becoming long and flat in posterior part of body.
Posterior chaetigers laterally flattened, terminally with an anal tube. Anal tube
pseudoarticulated, bearing a robust ventral cirrus (Fig. 8D).
Remarks. The species is similar to O. setigera and O. ammotrypanella sharing the
presence of prolonged chaetae in anterior segments. It differs from both species by
bearing a robust ventral cirrus on the anal tube. The anterior branchiae are larger than
that of O. ammotrypanella, but still shorter than the posterior branchiae of both species.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 91
Fig. 7: A-B Ammotrypanella cirrosa sp.n. A- entire animal, B- posterior segments, C-E Ammotrypanella
arctica MCINTOSH 1879 C- anterior segments dorsal view, D- anterior segments lateral view, E- posterior
segments, F-G Ammotrypanella princessa sp.n. F- posterior segments, G- anterior segments
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 92
Fig. 8: A-B Ammotrypanella mcintoshi sp.n. A- anterior segments, B- posterior segments, C-D Ophelina
robusta sp.n, C- anterior segments, D- posterior segments E-F Ophelina ammotrypanella sp.n. E- anterior
segments, F- posterior segments
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 93
3.3.4 Scalibregmatidae MALMGREN 1867
3.3.4.1 Pseudoscalibregma papilia sp.n.
Genus Pseudoscalibregma ASHWORTH 1901
Pseudoscalibregma papilia sp.n.
Holotype. ANDEEP I – II, South Sandwich Islands, Sta. 141-10, 23 March 2002,
58°24.08’S, 25°0.77’W, 2258-2313 m, EBS, ZMH P-24761
Paratypes. ANDEEP I – II, South Sandwich Islands, Sta. 141-10, 23 March 2002,
58°24.08’S, 25°0.77’W, 2258-2313 m, EBS, 2 specimens, ZMH P-24762
Additional material. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 42-2,
27 January 2002, 59°40.29’S, 57°35.43’W, 3683-3690 m, EBS, 3 specimens, Scotia Sea
northeast off Elephant Island, Sta. 46-7, 30 January 2002, 50°38.35’S, 53°57.36’W,
2889 m, EBS, 2 specimens
Etymology. The species is named after the shape of the posterior parapodia which
remind of butterfly wings.
Diagnosis. The species can be recognized by prominent, almost foliose dorsal and
ventral cirri in its posterior parapodia, distinctly rounded prostomial lobes and a rather
smooth to irregularly wrinkled body surface.
Description
Holotype. complete, 6 mm long and 1 mm wide for 33 chaetigers.
A moderately large species of 5 – 12 mm length and 0.5 – 1 mm width. Number of
chaetigers 26 – 33 (Fig. 9A). Color in alcohol white to a light tan. Body sometimes
expanded in anterior region to about chaetiger 12.
Prostomium with two spherical lobes anterolaterally; no eyes, nuchal organs not
apparent. Peristomium a single achaetous ring, well developed (Fig. 9B).
Body surface almost smooth, sometimes irregularly wrinkled, a scheme in annulation
not apparent. Anterior parapodia with reduced parapodial lobes, dorsal and ventral cirri,
these rapidly increasing in size in median region; posterior dorsal and ventral cirri of
large size, almost foliose, ventral cirri larger than dorsal ones; interramal sense organs
missing (Fig. 9C).
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 94
All parapodia with simple chaetae; furcate chaetae with unequal tynes covered by fine
hairs, present from chaetiger 2 (Fig. 9D).
Pygidium terminal, formed by a ring of large tubercles carrying cirri of different lengths
(Fig. 9E).
Remarks. The species is most similar to P. bransfieldium (HARTMAN 1967) which is
also very common in the Southern Ocean (Blake, 1981). They have in common the
moderately large size and the lack of a schematic annulation (unlike P. ursapium e.g.
which is covered by prominent tubercles). It can easily be distinguished from this
species by the lack of a prominent nuchal organ dorsally on the prostomium, the
distinctly spherical form of the anterolateral prostomial lobes, and the exceptionally
large size of the posterior dorsal and ventral cirri.
3.3.5 Sphaerodoridae MALMGREN 1867
3.3.5.1 Ephesiella hartmanae sp.n.
Genus Ephesiella Chamberlin, 1919 sensu Hartman & Fauchald, 1971
Ephesiella hartmanae sp.n.
Holotype. ANDEEP I – II, South Sandwich Islands, east off Monatgu Island, Sta. 143-1,
25 March 2002, 58°44.69’ S, 25°10.27’ W, 774 m, EBS, ZMH P-24763
Paratypes. ANDEEP I – II, South Sandwich Islands, east off Monatgu Island, Sta. 143-
1, 25 March 2002, 58°44.69’ S, 25°10.27’ W, 774 m, EBS, 2 specimens, ZMH P-24764
Etymology. The name refers to Olga Hartman, who contributed strongly to our
knowledge about the Southern Ocean polychaetes through detailed and thorough
taxonomic studies.
Diagnosis. The species can be recognized by the limited number of segments of less
than 50 and the lack of recurved hooks in the first parapodia.
Description
Holotype complete with 39 chaetigers. Length of body 4 mm, width 0.5 mm.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 95
A small and thin species of 3 – 4 mm length and 0.5 – 1 mm width for up to 40
chaetigers. Color in alcohol white to a light tan. Whole body, including prostomium and
pygidium covered with numerous small tubercles.
Prostomium with one median and one pair of lateral short antennae. One pair of
tentacular cirri present. Eyes absent (Fig. 10A).
Chaetigers with well developed parapodia. Dorsally with two lateral rows of
macrotuberclese each bearing a small papilla on top. A distinct row of microtubercles
dorsally to the macrotubercles.
Parapodia uniramous with a well developed ventral cirrus reaching the tip of the
parapodial lobe. A distinct prechaetal lobe dorsally on parapodium and a small
postchaetal lobe present (Fig. 10F). Chaetae composite falcigers with short appendanges
(Fig. 10C). Recurved hooks absent.
Pygidium with two large bulbous like appendages similar to macrotubercles and two
short cirri similar to microtubercles (Fig. 10B).
Remarks. The species is the third species of this genus described for the Southern Ocean
(Fauchald, 1974). In contrast to both other species from that region, E. antarctica
(MCINTOSH 1885) and E. pallida FAUCHALD 1974, it lacks recurved hooks in the first
parapodia. The chaetal appendages of E. hartmanae sp.n. are similar to those of E.
antarctica, the number of segments is similar to E. pallida.
3.3.5.2 Sphaerodoropsis HARTMAN & FAUCHALD 1971
3.3.5.2.1 Sphaerodoropsis distincta sp.n.
Sphaerodoropsis distincta sp.n.
Holotype. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 46-7, 30
January, 60°38.35’ S, 53°57.36’ W, 2889 m, EBS, ZMH P-24765
Paratypes. ANDEEP I – II, Scotia Sea northeast off Elephant Island, Sta. 46-7, 30
January, 60°38.35’ S, 53°57.36’ W, 2889 m, EBS, 5 specimens, ZMH P-24766
Etymology. The name refers to the well-defined and comparably large size of the
macrotubercles.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 96
Diagnosis. Diagnostic characters of this species are a distinct spherical and pronounced
shape of the macrotubercles and the presence of two pairs of lateral antennae.
Description
Holotype complete, 2 mm long and 0.5 mm wide for 13 chaetigers.
A delicate species of 1 – 4 mm length and 0.5 – 1 mm width. Chaetigers number 10 –
18. Body grub shaped, completely covered with fine tubercles (Fig. 10E). Color in
alcohol white.
Prostomium with one long median and two pairs of long lateral antennae. One pair of
tentacular cirri of similar length present. Eyes absent.
First chaetigers with reduced parapodia, subsequent parapodia well developed. Dorsum
covered with four rows of dorsolateral macrotubercles. These strongly pronounced,
spherical, sometimes distinctly white in color.
Parapodia uniramous, dorsally smooth, ventrally with one small tubercle and a short
ventral cirrus (Fig. 10D). One postchaetal lobe present. Parapodial lobe supported by
one acicula. Chaetae composite with long falcigerous appendages (Fig 10G).
Pygidium with two large bulbus like appendages, these spherical to triangular in shape
(Fig. 10E).
Remarks. At first sight the species seems to resemble S. parva (EHLERS 1913). The
macrotubercles take the same position in S. distincta sp.n. as in S. parva, their outline is
however more spherical and the size is larger in relation to total body size. In contrast to
S. parva the species only bears two lateral antennae, and all antennae are comparably
long.
3.3.5.2.2 Sphaerodoropsis maculata sp.n.
Sphaerodoropsis maculata sp.n.
Holotype. ANDEEP III, eastern Weddell Sea, st. 74-6, 20 January 2005, 71°18.42’ S,
13°58.21’ W, 1047 m, EBS, ZMH P-24767
Paratypes. ANDEEP III, eastern Weddell Sea, st. 74-6, 20 January 2005, 71°18.42’ S,
13°58.21’ W, 1047 m, EBS, 19 specimens, ZMH P-24768
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 97
Etymology. The name refers to the pigmentation of the body which is speckled with
black to purple dots.
Diagnosis. The macrotubercles are cone-shaped. The body is speckled with dark
pigments. Three pairs of lateral antennae are present.
Description
Holotype complete, 3 mm long and 0.5 mm wide for 15 chaetigers.
A small species up to 3 mm long and 0.5 mm wide for 14 – 15 chaetigers. Body grub
shaped, sparsely covered with distinct microtubercles. Distinct coloration due two
numerous purple spots throughout the body that resist fixation in ethanol, rest of body
white (Fig. 10H).
Prostomium with long antennae, one median and three lateral pairs present. One
additional pair of tentacular cirri present. Eyes indistinct brown dots dorsally on the
prostomium, visible under high magnification.
Dorsum with four pairs of lemon shaped macrotubercles and four rows of enlarged
mircotubercles in addition to the smaller microtubercles. Parapodia of all chaetigers
well developed with a distinct postchaetal lobe and a short ventral cirrus. No tubercles
on parapodial lobes present (Fig. 10I). Chaetae composite with long falcigerous
appendages (Fig. 10J).
Pygidium with two large bulbous appendages of triangular shape, and a median, long
cirrus (Fig. 10H).
Remarks. The species stands out from all other species found in the deep Southern
Ocean by a distinct pigmentation. The entire surface is spreckled with black to purple
dots of various sizes. The prostomium bears a pair of light brown eyes that seem to
vanish in fixation. The original position of the eyes can then only be determinde under a
microscope. The macrotubercles are somewhat lemon to cone-shaped and not distinctly
spherical as known for S. parva and S. distincta sp.n. Three pairs of lateral antennae are
present
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 98
3.3.5.2.3 Sphaerodoropsis simplex sp.n.
Sphaerodoropsis simplex sp.n.
Holotype. ANDEEP I –II, Scotia Sea northeast off Elephant Island, Sta. 46-7, 30
January, 60°38.35’ S, 53°57.36’ W, 2889 m, EBS, ZMH P-24769
Paratypes. ANDEEP I –II, Scotia Sea northeast off Elephant Island, Sta. 46-7, 30
January, 60°38.35’ S, 53°57.36’ W, 2889 m, EBS, 21 specimens, ZMH P-24770
Etymology. The name refers to the weakly developed macrotubercles.
Diagnosis. The species can be recognized by a stout body form and weakly pronounced
macrotubercles in lateral position.
Description
Holotype complete, 1.5 mm long and 0.5 mm wide for 16 chaetigers.
A small species with 1 – 2.5 mm length and 0.5 – 1 mm width for 10 – 21 chaetigers.
Body grub shaped, robust, completely covered with fine tubercles (Fig. 10K). Color in
alcohol white.
Prostomium with one median and two pairs of lateral antennae. One pair of tentacular
cirri present. Eyes absent.
Chaetigers with distinct parapodia, those of first chaetiger reduced. Dorsum covered
with four rows of dorsolateral macrotubercles. These somewhat rectangular, only little
pronounced. An additional transverse ridge formed by small mircotubercles across the
dorsum (Fig. 10K).
Parapodia uniramous, dorsally with two small tubercles. Ventral cirrus and postchaetal
lobe short. Parapodial lobe supported by one acicula (Fig. 10M). Chaetae composite
with short falcigerous appendages (Fig. 10L).
Pygidium with two bulbous appendages and short anal cirri (Fig. 10K).
Remarks. The shape of the macrotubercles is an outstanding characteristic of this
species. It differs from that of other Southern Ocean species in being only little
pronounced and almost square in outline. The body is more compact, almost a third as
wide as long. The species’ general appearance is most similar to S. parva, the lack of a
third pair of antennae however puts it closer to S. distincta sp.n.
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 99
Fig. 9: Pseudoscalibregma papilia sp.n. A- entire animal, B- anterior segments, C- posterior parapodium,
D- furcate chaetae, E- pygidium
RESULTS – COMMUNITY COMPOSITION & SPECIES IDENTIFICATION 100
Fig. 10: A-C, F Ephesiella hartmanae sp.n. A- anterior segments, B- posterior segments, C- composite
chaeta, F- parapodium, D-E, G Sphaerodoropsis distincta sp.n. D- parapodium, E- entire animal, G-
composite chaeta, H-I Sphaerodoropsis maculata sp.n. H- entire animal, I- parapodium, J-composite
chaeta, K-M Sphaerodoropsis simplex sp.n. K- entire animal, L- composite chaeta, M- parapodium
RESULTS- COMMUNITY ANALYSES
101
3.4 Univariate and multivariate community analyses
3.4.1 Analysis of the epi- and supranet
The EBS bears two nets for sampling, the epi- and the supranet. Samples from both nets
were fixed, sorted and counted separately. Additionally, individuals found in the gear
outside the nets were treated likewise and labeled with ‘Ü’ (abbreviation for
“Überstand” supernatant). For the expedition ANDEEP III all stations are represented
by an epi- and a supranet. For ANDEEP I-II only the samples of station 134-4 are
complete. The other stations are represented by each one net only.
In order to test the null hypothesis that there is no significant difference between the
similarities of the nets of one station and that of different stations, a one-way ANOSIM
was performed on the species assemblage matrix of ANDEEP III (standardized to 1000
m trawling distance). The result with R = 0.628 shows (Fig. 11) that the null hypothesis
can be rejected with a level of significance of 0.1 %.
Fig. 11: Comparison of epi- and supranet similarities of EBS samples of ANDEEP III by means of one-
way ANOSIM with 999 permutations
RESULTS- COMMUNITY ANALYSES
102
A standard operation of a student’s t-test on the unstandardized family data of ANDEEP
III (App.-Tab. 2) results in a rejection of the null hypothesis of 0.012 % (degrees of
freedom: 90, t = -2.58, standard deviation = 209).
It is therefore statistically proven that the nets of one station are significantly more
similar to each other than nets of different stations. For further analyses the nets of each
station are since summed up and treated as one sample.
3.4.2 Species richness and biodiversity
During the expeditions ANDEEP I, II and III a total of 14,176 polychaete specimens
were sampled. The individuals were sorted into 47 different families (App.-Tab. 3). The
most abundant families were the Spionidae, Hesionidae, Syllidae, and Polynoidae with
more than 1000 individuals each (App.-Fig. 2A-B).
Of the 47 families the Ampharetidae, Glyceridae, Goniadidae, Hesionidae, Nephtyidae,
Nereididae, Opheliidae, Sabellariidae, Scalibregmatidae, Sphaerodoridae, Syllidae,
Terebellidae, and Trichobranchidae have been determined to species level, 155 different
species were distinguished.
Based on the resulting species assemblage matrix for ANDEEP I-III a species
accumulation plot has been drawn. The curve shows no saturation (Fig. 12) and
consequently suggests an undersampling of the sampled areas.
Fig. 12: Species accumulation plot based on species assemblage matrix of ANDEEP I-III
RESULTS- COMMUNITY ANALYSES
103
3.4.2.1 Species richness (Margalef’s Index)
The great differences in sampling distance and sample volume between the stations of
the expeditions ANDEEP I-III do not allow an analysis of the overall data volume by
means of quantitative biodiversity measures as e.g. the Shannon Index. Therefore, the
Margalef’s Index for species richness [d] is chosen to compare all stations. This index
relates the number of species (respectively families) with the number of individuals at
each station. It does not consider the abundance of species (respectively family) – the
dependence on the sample volume is therefore distinctly weaker than for quantitative
indices.
Fig. 13 presents the Margalef’s Index for all stations of ANDEEP I-III at family level.
In addition, the trawling distances of station are shown. A correlation between the size
of the sampled area and the family richness [dF] is seen for stations within the Drake
Passage (st. 41-3, 42-2, 46-7), the South Sandwich Trench (st. 141-10, 143-1), near
King George Island (st. 153-7, 154-9), and within the central Weddell Sea (st. 131-3,
132-2, 133-3, 134-4). This is evidence that the differences in family richness of stations
within these regions might be biased by the trawling distances. No correlation is
apparent for stations connecting the eastern Weddell Sea with the central Weddell Sea
(st. 88-8, 94-14, 102-13, and 110-8). While the trawling distances decrease from east to
west, family richness increases. The dF’s of these stations are lower than that of stations
near the eastern Weddell Sea coast (st. 74-6, 78-9, 80-9, 81-8) and that of st. 16-10, 21-
7 and 59-5 connecting the South Atlantic with the eastern Weddell Sea. The highest
family richness is reported for the Drake Passage, the Southern Atlantic and Eastern
Weddell Sea, and the area around the King George Islands with dF’s > 3.8. Those of the
central Weddell Sea lie between 1.6 and 3.7. The smallest family richness is found at st.
152-6 (dF = 0.9) positioned in the Drake Passage near Elephant Island.
A similar picture is presented when the taxonomic level is elevated to the observation of
species (Fig. 14). Correlation of species richness to trawling distances is found for the
same areas as at family level. Also, st. 132-2, 133-3, 134-4, and 135-4 (central Weddell
Sea), and st. 88-8, 94-14, 102-13, and 110-8 on the transect between the central Weddell
Sea and the eastern Weddell Sea show the lowest species richnesses of dS’s < 4. High
species richnesses are found at st. 150-6 (South Orkney Islands), 153-7 (King-George
RESULTS- COMMUNITY ANALYSES
104
Island), 42-2 and 46-7 (Drake Passage), and st. 80-9 (eastern Weddell Sea). The highest
value is found in the eastern Weddell Sea at station 74-6 [dF = 9.2]. It is based on the
smallest area sampled. The smallest dS’s are found at stations 135-4 (central Weddell
Sea) and 152-6 (near Elephant Island).
Margalef's Index
0
1
2
3
4
5
6
41-3
42-2
46-7
131-
313
2-2
133-
313
4-4
135-
414
1-10
143-
116
-10
21-7
59-5
74-6
78-9
80-9
81-8
88-8
94-1
410
2-13
110-
812
1-11
133-
214
2-5
150-
615
1-7
152-
615
3-7
154-
9
station
fam
ily ri
chne
ss
05001000150020002500300035004000
traw
ling
dist
ance
(m)
dF
traw ling distance (m)
Fig. 13: Family richness for ANDEEP I-III-stations (Margalef’s Index)
Margalef's Index
0123456789
10
41-3
42-2
46-7
131-
313
2-2
133-
313
4-4
135-
414
1-10
143-
116
-10
21-7
59-5
74-6
78-9
80-9
81-8
88-8
94-1
410
2-13
110-
812
1-11
133-
214
2-5
150-
615
1-7
152-
615
3-7
154-
9
station
spec
ies
richn
es
0
500
1000
1500
2000
2500
3000
3500
4000
traw
ling
dist
ance
(m)
dStrawling distance (m)
Fig. 14: Species richness for ANDEEP I-III-stations (Margalef’s Index)
RESULTS- COMMUNITY ANALYSES
105
3.4.2.2 Biodiversity and Evenness
Quantitative biodiversity measures are applied to the species data of the expedition
ANDEEP III. In contrast to ANDEEP I/II the samples are complete, no vials were lost
during transfer from the vessel to the home laboratories. Therefore, comparability of
abundances can be achieved by standardization to 1000 m2 sampling area (App.-Tab. 7).
As biodiversity measures the Shannon Index and the Pielou’s Evenness Index were
chosen. Both indices have proven to be suitable for most benthic diversity data and are
commonly used. Comparable data, especially of different taxa, is therefore available.
During ANDEEP III a total of 10,635 specimens belonging to 46 different families were
sampled. The most abundant families were the Spionidae, Hesionidae, Syllidae, and
Polynoidae. The sampling volume ranged between 58 specimens (st. 102-13, sampling
area 3283 m2, central Weddell Sea) and 4226 specimens (st. 133-2, sampling area 1164
m2, central Weddell Sea). Standardized to 1000 m2 sampling area a theoretical total of
8471 specimens was sampled, the most abundant families being the same as in the
unstandardized data.
At family level the highest diversity (H’ > 2.5) is found in the Southern Atlantic (st. 16-
10, 21-7, 59-5), the eastern Weddell Sea (st. 78-9, 80-9, 81-8), and around King George
Island (st. 153-7). The central Weddell Sea is clearly the least diverse area sampled
(H’= 1.6 – 2.4). Within the central Weddell Sea the easternmost stations (st. 88-8, 94-
14) are less diverse than that close to the Antarctic Peninsula (st. 133-2, 142-5),
resulting in an east to west gradient in family diversity. The least diverse station is st.
152-6 near Elephant Island with H’ = 0.3. The highest evenness is found at station 142-
5 in the western part of the central Weddell Sea with J = 0.89. The lowest is found for
station 152-6 (Elephant Island, J = 0.2). Relatively similar values are found for stations
from the Southern Atlantic to the eastern Weddell Sea with J = 0.7 – 0.85. Evenness in
the central Weddell Sea (st. 88-8 to 133-2) and around King George Island (st. 153-7
and 154-9) lies below J = 0.8. (Fig. 15).
On a geographical scale species diversity is similar to family diversity (Fig. 16). The
central Weddell Sea (st. 88-8 to 142-5) is less diverse than the easternmost and
westernmost sites. Also, st. 152-6 is least diverse (H’ = 1.04). In comparison to family
level, the differences between the diversity indices of the stations are more distinct. The
RESULTS- COMMUNITY ANALYSES
106
highest diversity is found at st. 150-6 (South Orkney Islands, H’ = 3.04), 16-10 (South
Africa, H’ = 3.02), 74-6 (eastern Weddell Sea, H’ = 3.0), and 154-9 (King George
Island, H’ = 2.87). The most diverse stations in the central Weddell Sea are st. 121-11
and 133-2 (J = 2.59). Evenness is highest for st. 152-6 with J = 0.95. With exception of
st. 74-6 (J = 0.72), the evenness for stations from the Southern Atlantic and the eastern
Weddell Sea ranges from J= 0.8 (st. 80-9, 81-8) and J= 0.92 (st. 16-10). Evenness of the
central Weddell Sea (st. 88-8 to 142-5) ranges from J = 0.61 – 0.9, that of the King
George Island stations are J = 0.76 (st. 153-7) and J = 0.89 (st. 154-9).
Biodiversity and Evenness
0
0,5
1
1,5
2
2,5
3
16-1
021
-759
-574
-678
-980
-981
-888
-894
-14
102-
1311
0-8
121-
1113
3-2
142-
515
0-6
151-
715
2-6
153-
715
4-9
station
Shan
non
Inde
x/ P
ielo
u's
Even
ness
0
510
15
20
2530
35
40
tota
l fam
ilies
J'H'(loge)S
Fig. 15: Family diversity and evenness of ANDEEP III-stations (Shannon Index, Pielou’s Evenness)
Biodiversity and Evenness
0
0,5
1
1,5
2
2,5
3
3,5
16-1
021
-759
-574
-678
-980
-981
-888
-894
-14
102-
1311
0-8
121-
1113
3-2
142-
515
0-6
151-
715
2-6
153-
715
4-9
station
Shan
non
Inde
x/ P
ielo
u's
Even
ness
0
10
20
30
40
50
60
70
spec
ies
tota
l
J'
H'(loge)
S
Fig. 16: Species diversity and evenness of ANDEEP III-stations (Shannon Index, Pielou’s Evenness)
RESULTS- COMMUNITY ANALYSES
107
3.4.3 Clustering and Multi Dimensional Scaling (MDS)
With help of the Bray-Curtis Index similarities between different stations can be
defined. These statistical numbers are presented in similarity resemblance matrices
(App.-Tab. 8-10) based on which dendrograms of the relationships between stations
(clustering) and two- to multidimensional plots (MDS) can be calculated to graphically
visualize the similarities.
In the following chapter cluster and MDS plots for all ANDEEP stations are presented.
3.4.3.1 ANDEEP I-III
To minimize bias resulting from different trawling distances and sample volumes the
species matrix was transformed to a presence/ absence matrix. After calculation of
Bray-Curtis similarities, cluster and MDS analyses for the stations of ANDEEP I-III at
species and family level were performed.
Clustering of species results in eleven statistically supported groups with 1 – 11 stations
(Fig. 17). The similarities between stations lie below 50 %. Geographical relations are
not apparent in the analysis. The largest cluster with eleven stations includes stations of
the Drake Passage (st. 42-2), the central Weddell Sea (st. 110-8, 121-11, 131-1, and
134-4), and the eastern Weddell Sea (st. 78-9, 102-13). Also, st. 16-10 and 59-5
between South Africa and the eastern Weddell Sea, and st. 153-7 and 154-9 off King
George Island are found in this main cluster.
The MDS plot (Fig. 18) suggests the presence of one main clusters and six solitaire
stations (st. 41-3, 152-6 from the Drake Passage, and st. 132-2, 133-3, 135-4, and 142-5
from the central Weddell Sea). The remaining Weddell Sea stations (WS) and all
eastern Weddell Sea stations (EWS) are concentrated in the upper part of the main
clusters. Also within the WS and EWS sites stations 16-10, 21-7 (SA, Southern
Atlantic), and 59-5 (EA, eastern Atlantic sector of the Southern Ocean), as well as the
stations from King Georg Island (KGI), st. 153-7 and 154-9, are found, indicating high
similarities between the polychaete assemblages in the Southern Atlantic, the Weddell
Sea and west off the Antarctic Peninsula. Stations 141-10 and 143-1 from the South
RESULTS- COMMUNITY ANALYSES
108
Sandwich Trench (SST) and the stations of the Drake Passage and South Orkney Islands
(DP, st. 42-2 and 46-7, SOI, st. 150-6, 151-7) form the bottom part of the main cluster.
They are less close to the WS and EWS stations than the SA, EA, and KGI stations. A
clear formation of a separate clusters is, however, not apparent.
Fig. 17: Cluster analysis of ANDEEP I-III-stations (pres/abs – species matrix, Bray-Curtis similarities)
Fig. 18: MDS plot of ANDEEP I-III-stations (pres/abs – species matrix, Bray-Curtis similarities)
RESULTS- COMMUNITY ANALYSES
109
Clustering and MDS ordination of the family data result in a similarly weak resolution
as that of the species data. Therefore only the MDS plot is presented (Fig. 19). A main
cluster including the stations 153-7 and 154-9 (King George Island), stations 16-10, 21-
7 and 59-5 (South Africa to Eastern Antarctica), stations 141-10 and 143-1 (South
Sandwitch Trench), stations 150-6 and 151-7 (South Orkney Island), and some stations
from the East and Central Weddell Sea (74-6, 78-9, 80-8, 81-8, 121-11, 131-3, 133-2)
can be observed. The stations from the central and eastern Weddell Sea are positioned
in the lower part of the ordination while the stations positioned closer to the Drake
Passage lie in the upper part. Below the main cluster, stations 88-8, 94-14, 102-13, and
110-8, positioned on a transect connecting the eastern with the central Weddell Sea,
form a small group, together with station 135-4 (WS). These stations seem to be similar
in their faunal composition. Also stations 132-2, 133-3 and 134-4 from the central
Weddell Sea are positioned below the main cluster. Although these stations are not part
of the main cluster, a close relation to the other stations from the Weddell Sea is
evident. St. 41-3 (DP), in contrast, lies above the main cluster closer to the stations from
the Drake Passage and the South Sandwich Trench. St. 152-6 (EI) is the station least
similar to all other stations.
A
B
Fig. 19. MDS plot of ANDEEP I-III-stations,
A- entire, B- detail of main cluster (pres/abs – family matrix, Bray-Curtis similarities)
RESULTS- COMMUNITY ANALYSES
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3.4.3.2 ANDEEP I/II
As for the analysis of the ANDEEP I-III data, the species assemblage matrix of
ANDEEP I/II was transformed to a presence/ absence matrix. After calculating the
similarities by Bray-Curtis Index, cluster and MDS analyses at species level were
performed.
Cluster analysis (Fig. 20) of the species data results in two solitaire stations (st. 132-2
and 133-3 from the Weddell Sea) and two statistically supported clusters. Both clusters
include stations from the Weddell Sea, the Drake Passage, and the South Sandwich
Trench.
The similarities between stations lie below 40 %. This results in a poorly resolved MDS
plot (Fig. 21). The ordination of stations seems rather random without clear groupings.
It becomes apparent that the stations of the Weddell Sea (st. 131-3 to 134-5) are
concentrated in the lower part of the plot, that of the Drake Passage (st. 41-3, 42-2, and
46-7) and the South Sandwich Trench (st. 141-10 and 143-1) in the central and upper
part.
Fig. 20: Cluster analysis of ANDEEP I/II-stations (pres/abs – species matrix, Bray-Curtis similarities)
A comparison of this result with station depths and the species matrices (Tab. 1, App.-
Tab. 4) shows that all stations below 2000 m and with more than ten species cluster
together (stations 131-3, 134-4, 141-10, 42-2, and 46-7). For better resolution of station
similarities, MDS analysis is applied to an excerpt of these stations (Fig. 22). The
RESULTS- COMMUNITY ANALYSES
111
resulting plot indicates a relation between faunal composition and geographical
position. Stations of the same basins (stations 131-3 and 134-4 from the Weddell Sea,
stations 42-2 and 46-7 from the Drake Passage, and station 141-10 from the South
Sandwich Trench) are ordinated closest to each other. Additionally, similarities between
the Drake Passage and the Weddell Sea are suggested. The similarities of the South
Sandwich Trench (st. 141-10) to the Weddell Sea (st. 131-3 and 134-4) are lower than
to the Drake Passage (st. 42-2 and 46-7).
Fig. 21: MDS plot of ANDEEP I/II-stations (pres/abs – species matrix, Bray-Curtis similarities)
Fig. 22: MDS plot of ANDEEP I/II-stations deeper 2000 m and with at least ten different species
(pres/abs – species matrix, Bray-Curtis similarities)
RESULTS- COMMUNITY ANALYSES
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3.4.3.3 ANDEEP III
Before calculation of the resemblance matrix for ANDEEP III (App.-Tab. 10), a fourth
root overall transformation was used to strengthen the position of rare species and
weaken that of very abundant species. The Bray-Curtis Index is then applied, cluster
and MDS analyses are carried out.
Cluster analysis results in four solitaire stations of which st. 152-6 (Drake Passage, near
Elephant Island) is the one least similar to all others (Fig. 23). In addition, four clusters
can be recognized. The first combines stations 74-6 from the eastern Weddell Sea and
133-2 from the central Weddell Sea. In a second cluster station 21-7 from the Southern
Atlantic, and st. 88-8 and 94-14 between the eastern and central Weddell Sea are
included. Both remaining clusters include stations from the Southern Atlantic, the
eastern Weddell Sea, the central Weddell Sea, and off King George Island (western
Antarctic Peninsula).
Fig. 23: Cluster analysis of ANDEEP III-stations (standardized species matrix, Bray-Curtis similarities)
The MDS plot (Fig. 24) supports the solitaire positions of stations 81-8 (EWS), 142-5
(WS), 151-7 (SOI), and 152-6 (EI), and the high similarity between stations 74-6
(EWS) and 133-2 (WS). All remaining stations are combined in one main cluster. In its
lower part the South Atlantic stations (st. 16-10, 21-7, and 59-5) group within stations
RESULTS- COMMUNITY ANALYSES
113
from the eastern Weddell Sea (st. 81-8 to 110-8). In the upper part the stations off King
George Island (st. 153-7 and 154-9) and the South Orkney Islands (st. 150-6) cluster
with stations of the eastern and central Weddell Sea (st. 78-9, 80-9 and 121-11).
Fig. 24: MDS plot of ANDEEP III-stations (standardized species matrix, Bray-Curtis similarities)
3.4.4 Correlation of polychaete communities to environmental data (BIO-ENV)
During the expeditions ANDEEP I/II data about sediment type at different sites were
collected. Information are taken from Howe et al. (2004) and Diaz (2004) and presented
in App.-Tab. 11. Taken into account are minimum and maximum grain size, water
depth, and O2 depth in the sediment layer. The grain sizes of stations 41-3, 42-2, 46-7,
and 143-7 were only expressed in words, not measures. Measures were therefore
complemented through comparison of the overall description of these stations to other
stations. In addition, O2 depths for stations 41-3 and 143-1 are unknown.
A correlation of the ecological data with the species data of ANDEEP I/II is achieved
with the Primer v 6.1.6 function BIO-ENV (compare material and methods). The results
give an idea which combination of environmental factors might play an important role
in determining species composition of stations. Two different station combinations are
analyzed. First all stations of ANDEEP I/II are considered. The station list is then
RESULTS- COMMUNITY ANALYSES
114
reduced to stations deeper than 2000 m and with at least ten species. The resulting five
best matches and their significance level are presented in Tab. 2.
Tab. 2: Most likely factor combinations explaining the species distribution of ANDEEP I-II, B 1- best
match, to B 5- fifth best match, factors: minimum grain size (1), maximum grain size (2), water
depth (3), O2 depth (4)
Station matrix B 1 B 2 B 3 B 4 B 5 Significance (%)
All stations 4 3 1,3 3,4 1,3,4 29
Reduced stations 2,4 1,2,4 2 1,2 1 12
Considering all stations of the expedition, minimum grain size and water depth play an
important role in the species distribution. Also O2 depth has to be taken into account.
After reducing the number of stations to those below 2000 m depth and with at least ten
different species, grain size seems to be the most important factor.
In analogy to the species assemblage matrices the family composition of stations is
correlated to ecological factors. This is of interest because the family assemblage
matrices consider all polychaete specimens found during the expedition, and therefore
represent a more complete excerpt of the real community. During the analysis two
different station lists are taken into account. First, all stations of ANDEEP I/II are
included; second, all stations with less than ten species and 2000 m water depth are
excluded.
Tab. 3: Most likely factor combinations explaining the family distribution of ANDEEP I-II, B 1- best
match, to B 5- fifth best match, factors: minimum grain size (1), maximum grain size (2), water
depth (3), O2 depth (4)
Station matrix B 1 B 2 B 3 B 4 B 5 Significance (%)
All stations 2 1,2 2,4 1,2,4 1 60
> 2000 m depth, min. 10 species 4 1,4 1 2,4 1,2,4 6
The significance level comparing all stations is very high (60 %), the results show no
evidence for a relation between depth and family composition (Tab. 3). The significance
RESULTS- COMMUNITY ANALYSES
115
level of the reduced station list is distinctly lower. The excerpt of all stations below
2000 m and with at least ten species favors a correlation between sediment characters,
especially minimum grain size and O2 depths, to family composition.
3.4.5 Correlation of species to stations similarities (BV Step)
In addition to the correlation to ecological factors, species composition can be
correlated to a set of species that has the greatest influence on the similarities between
stations by BV Step-analysis. Due to varying sampling volumes this analysis is only
applied to the standardized data of ANDEEP III.
The result (with a significance level of 1 %) suggests a set of 15 species to most likely
determine the station similarities. These species are: Aglaophamus paramalmgreni,
Ammotrypanella arctica, Ampharete kerguelensis, Amphicteis sp. 3, Kefersteinia
fauveli, Kesun abyssorum, Micropodarke cylindricaudata sp.n., Ophelina
ammotrypanella sp. n., Ophelina robusta sp.n., Ophiodromus calligocervix sp.n.,
Ophiodromus comatus, Polycirrus insignis, Pseudoscalibregma papilia sp.n.,
Sosanopsis kerguelensis and Travisia kerguelensis.
3.4.6 Average Taxonomic Distinctness (AvTD) and Variation in Taxonomic
Distinctness (VarTD)
The AvTD and the VarTD for all stations of ANDEEP I-III were calculated and are
separately compared with an expected range (95 % significance range) based on a
master list (all species found during ANDEEP I-III). The results are presented in two
funnel plots.
The AvTD funnel plot shows (Fig. 25) that stations 81-8 (eastern Weddell Sea), 121-11
and 131-3 (central Weddell Sea) have a significantly smaller average taxonomic
distinctness, indicating that the species found at these stations are more closely related
to each other than expected. The AvTDs of stations 134-4 (Weddell Sea), 16-10, 78-9,
and 74-6 (South Atlantic and eastern Weddell Sea) lie at the lower border of the
RESULTS- COMMUNITY ANALYSES
116
expected values for the according number of species. Stations 152-6, 133-3 and 135-4
(off Elephant Island and central Weddell Sea) show the highest AvTDs. Also stations
141-10 (South Sandwich Trench) and 46-7 (Drake Passage) have AvTDs close to being
significantly higher than expected.
The VarTD shows almost the inverse result (Fig. 26) of the AvTD, both values being
strongly negatively correlated in most cases. Stations 121-11 and 131-3 show a
significantly higher VarTD than the expected values, with Λ+ around 400 (expected
value for Λ+ between 100 – 300). Also the VarTDs of stations 134-4, 110-8, 16-10, and
78-9 (South Atlantic, eastern and central Weddell Sea) lie above the 95 % significance
range. Stations 81-8, 133-2 and 74-6 (eastern and central Weddell Sea) have increased
VarTDs, the increase being not significant. The lowest VarTDs are found for stations
152-6, 133-3 and 135-4 (off Elephant Island and central Weddell Sea).
Due to the negative correlation of the results the joint analysis of AvTD and VarTD
allows a better comparison of stations with the expected values (Fig. 27). Stations 152-
6, 133-3 and 134-5, that were at the border of being significantly different from the
expected in the analyses above, now lie within their expected range. Also does station
134-4. Stations 131-3, 121-11, 110-8, 81-8 (eastern and central Weddell Sea), and 16-10
(South Atlantic) show significant difference, with values expected for stations with
lower species abundances. Stations 42-2 (Drake Passage), 78-9 and 74-6 (eastern
Weddell Sea), show expected values. All remaining stations have values significantly
higher than the expected, indicating that the diversity of these stations is based on
species that are less taxonomically related than expected.
RESULTS- COMMUNITY ANALYSES
117
Fig. 25: AvTD of ANDEEP I-III-stations (95 % significance range)
Fig. 26: VarTD of ANDEEP I-III-stations (95 % significance range)
RESULTS- COMMUNITY ANALYSES
118
A
B
Fig. 27: AvTD/ VarTD- joint analysis for ANDEEP I-III-stations (95 % significance range), A- entire
result, B- detail
RESULTS- VERTICAL AND GLOBAL DISTRIBUTION PATTERNS
119
3.5 Zoogeography
3.5.1 Vertical distribution patterns in the Southern Ocean
The following results present the vertical distribution of the Southern Ocean deep-sea
polychaetes collected during the expeditions ANDEEP I-III.
App.-Fig. 5 illustrates the depth ranges in which species were found. The least number
of species (15) is found at stations shallower than 1000 m depth. Of these, fourteen
species are also found in deeper waters, only Axiokebuita millsi is limited to the shallow
depth. Thirty species are only found in depths shallower than 3000 m, these include
Gyptis incompta, Sphaerosyllis antarctica, Syllides articulosus, and all species of the
genera Phisidia, Pista, and Typosyllis.
A total of thirty-six species was collected from depths of 0 m to 5000 m, respectively
1000 m to 5000 m, and consequently can be considered eurybath species. The most
abundant of these are Aglaophamus paramalmgreni, Aglaophamus trissophyllus,
Anobothrus pseudoampharete sp. n., Glycera kergulensis, Kefersteinia fauveli,
Ophelina breviata, Ophelina gymnopyge, Sosanopsis kerguelensis, Sphaerodoropsis
parva, and Terebellides stroemi. Fifty-six species are exclusively found below 2000 m,
fourteen of them being collected from below 3000 m and seven from below 4000 m.
The depth range richest in species is that between 1000-3000 m.
3.5.2 Global distribution patterns
Global distribution patterns were compiled for all named species found during
ANDEEP I-III including data from former records (compare chapter 2.4.2). New
species are not included. App.-Tab. 12 gives the complete list of 84 species and a global
distribution category.
As shown in App.-Fig. 6, the distribution of the thirteen families analyzed is not limited
to the Southern Ocean but covers all ocean basins world-wide. Most species records
outside the Southern Ocean originate from Atlantic sites, especially from the Northern
Atlantic. Records from the Pacific are mostly from the west coast of South and North
RESULTS- VERTICAL AND GLOBAL DISTRIBUTION PATTERNS
120
America, or from sites close to Australia. Records from the Indic are rare, maily due to
an undersampling of this ocean.
A separate view on the single families shows that distribution patterns differ distinctly
at this taxonomic level. In the following an overview of the family distribution is given
in alphabetical order.
Characterized by a high number of species (thirteen named species are observed, all
from different genera), the Ampharetidae are very wide spread with a clear tendency to
being bipolar (App.-Fig. 8). Records from temperate waters are rare. The centre of
distribution lies in subantarctic waters. Four cosmopolitan species are found (App.-Fig.
7A). Amphicteis gunneri, Melinna cristata, and Muggoides cinctus are recorded from
northern-hemisphere waters with a connection to the Atlantic ocean, Anobothrus
gracilis is additionally found in the North Pacific. Five species seem to be endemic to
the Southern Ocean. Ampharete kerguelensis, Grubianella arctica, and Sosanopsis
kerguelensis are additionally found at ANDEEP station 16-10 in the eastern South
Atlantic.
The Glyeridae and Goniadidae are each represented by one species only in this analysis.
While Glycera kerguelensis is restricted to the subantarctic region, Goniada maculata
has been reported from Atlantic and Pacific sites in the northern hemisphere and from
the Red Sea (App.-Fig. 9-10).
The three hesionid species are mainly found in Antarctic and subantarctic waters, Gyptis
incompta reaching into the South Pacific (App.-Fig. 11). A similar distribution is found
for Nereis eugeniae, the only Nereididae included (App.-Fig. 12). A locally restricted
distributional pattern is observed for the Nephtyidae that are only represented by the
genus Aglaophamus (App.-Fig. 13).
The Opheliidae are represented by two genera that show distinct differences in their
distributional patterns. The species of the genus Ophelina are only recorded from the
southernmost Atlantic and the Atlantic district of the Southern Ocean (App.-Fig. 14).
Ophelina gymnopyge and O. scaphigera were to date only known for the Weddell Sea
while O. breviata and O. nematoides have been reported for the subantarctic region.
During the ANDEEP cruises these four species were additionally found at station 16-10
north of the Antarctic Convergence. Ophelina setigera is reported from the Weddell Sea
quadrant in the Southern Ocean, in addition records from the South Atlantic were found
RESULTS- VERTICAL AND GLOBAL DISTRIBUTION PATTERNS
121
in the Angola Basin (SE Atlantic, Ebbe, pers. comment).The second genus found is
represented by Ammotrypanella arctica which has a clearly cosmopolitan distribution
(App.-Fig. 7B).
Only one sabellariid species is found (Phalacrostemma elegans) which is also known
for southern and northern Atlantic waters (App.-Fig. 15).
The most common distributional pattern within the Scalibregmatidae is a subantarctic
one (App.-Fig. 16). Also two species restricted to the Weddel Sea are found, namley
Oligobregma hartmanae and Sclerocheilus antarcticus (App.-Tab. 12). Travisia
kerguelensis and T. lithophila are found throughout the Southern Ocean, T. lithophila is
also recorded from the east Australian coasts. Hyboscolex equatorialis is reported for
some sites at the pacific coasts of South America. Altogether three potentially
cosmopolitan species are found (App.-Fig. 7C). Of these Scalibregma inflatum shows
the distribution furthest spread, while Axiokebuita millsi and Axiokebuita minuta show a
tendency to a bipolar distribution.
Of the four sphaerodorid species, Sphaerodoropsis parva shows the widest distribution,
being reported for the whole Southern hemisphere (App.-Tab. 12). Sphaerodoropsis
polypapillata has so far only been recorded for the Weddell Sea, during this study new
records from station 16-10 are found. This is evidence for a South Atlantic distribution
of this species (App.-Fig. 17). Ephesiella antarctica shows a circumpolar distribution
while Clavorodum antarcticum seems to be endemic for the Weddell Sea (App.-Fig.
7D).
The distibutional patterns within the Syllidae are very heterogenous. Of the 14 species
found, six show a restricted distribution (App.-Fig. 7E), including both Sphaerosyllis
species (Sphaerosyllis joinvillensis and Sphaerosyllis lateropapillata uteae) as well as
Pionosyllis maxima and Pionosyllis epipharynx (App.-Fig. 18). Autolytus gibber and
Proceraea mclearanus show a subantarctic and South Pacific distribution. In spite of
the high number of locally restricted species, four cosmopolitan species are found with
Braniella palpata, Exogone heterosetosa, Typosyllis hyalina, and Typosyllis variegata.
The widest distribution of all families analyzed is discovered in the Terebellidae.
Hereby a clear difference between the genera can be found. Most species are locally
restricted (3 species) or subantarctic (11 species) (App.-Fig. 7F). Among these are the
species of the genera Leaena, Thelepides, and Polycirrus (App.-Fig. 19). Only Leaena
RESULTS- VERTICAL AND GLOBAL DISTRIBUTION PATTERNS
122
collaris is found at station 16-10 and thus north of the Antarctic Convergence. The
genus Pista shows a far spread distribution, Pista cristata being a cosmopolite that is
reported for the North Pacific, North Atlantic, as well as the Red Sea and the
Mediterranean. Four further cosmopolitan species are found (Laphania boeckii,
Hauchiella tribullata, Thelepus cincinnatus, and Proclea graffii), Laphania boeckii and
Hauchiella tribullata are concentrated in polar waters of the Arctic and south of South
Africa.
A similar difference between the genera is observed within the Trichobranchidae. While
Octobranchus antarcticus is restricted to the Weddell Sea, Terebellides stroemi is
supposed to be a cosmopolitan species (App.-Fig. 20).
Differences in species richness and diversity (Fig. 14, 16) give evidence that sites in the
Drake Passage, around the Antarctic Peninsula, and the eastern Weddell Sea are
influenced by adjacent ocean basins. These influences might be missing in the central
Weddell Sea resulting in lower diversities. Additionally, clustering and MDS analyses
presented under chapters 3.4.3.1 – 3.4.3.3 indicate that stations from South Atlantic,
eastern Weddell Sea and central Weddell Sea sites are closely related. The same applies
to stations from the Drake Passage, off King George Island and the South Sandwich
Trench. The results suggest the presence of an east to west gradient in polychaete
composition in the Southern Ocean originating from different faunal influences from
Atlantic and Pacific waters. If such a longitudinal gradient exists it should be reflected
in an east to west shift in the distribution patterns of species found at eastern and
western sites. Species known for Pacific waters are expected to be present in higher
numbers at stations in the Drake Passage and around the Antarctic Peninsula. Similarly,
species reported from Indic and eastern Southern Atlantic waters are mainly expected at
eastern Weddell Sea sites. App.-Fig. 21-29 present the distribution charts for the regions
as defined in the community analysis. No clear difference is found between
distributional patterns of eastern and western sites. Species recorded for the South
Pacific are found at all sites including those close to South Africa and in the eastern
Weddell Sea. In return, species from the northern Atlantic have been collected from
west of the Antarctic Peninsula. In contrast, it becomes obvious that with rising number
RESULTS- VERTICAL AND GLOBAL DISTRIBUTION PATTERNS
123
of stations per region and therefore of species per region the number of world-wide
records changes but not the extent of the patterns.
The same effect can be seen when comparing the global patterns for the different depth
ranges as defined above (App.-Fig. 30-34). Depths between 1000-5000 meter contain
species from ocean basins world-wide. The four charts resulting are almost identical
(App.-Fig. 31-34). Only the chart regarding the species found in depths shallower than
1000 m is different (App.-Fig. 30). Species records are exclusively found from southern
hemisphere sites. The chart, however, is only based on one station. The species number
included is smaller than that from the other depth ranges possibly causing the different
pattern.
DISCUSSION
124
4 Discussion 4.1 Efficiency of the gear and methods used for sampling
The epibenthic sledge is constructed to function at any depth, and on soft sediment as
well as primary hard substrate (Brenke, 2005). The width of the netboxes (1 m) allows
an easy determination of the sampled area with given trawling distance. The mesh size
of the net cones is 500 µm, plus an additional filter of 300 µm in the net buckets. This
combination ensures the sampling of macrofaunal elements of all sizes while at the
same time the bycatch of hugh sediment volumes is minimized. Valves and
mechanically closing flaps secure the catch inside the net buckets and cone nets during
heaving of the EBS (Brenke, 2005). Brenke (2005) gives instructions about the handling
of the EBS and the calculation of the trawling distance. During the expedition ANDEEP
I–III these instructions were followed as closely as possible. Individual adjustments to
weather and current conditions were carried out when required. The calculation of the
trawling distance and the sampling area allows standardization of abundances to 1000
m2 (Basford et al., 1989; Svavarsson et al, 1990; Brattegard & Fossa, 1991; Brandt,
1993; Brandt & Piepenburg, 1994). A quantitative comparability of qualitative data is
achieved this way. Nevertheless, values have to be handled with great care since the
EBS does not function without a systematic error. Brenke (2005) predicted a minimum
systematic error of 25 % at all depths. He states, however, that this prediction is only
preliminary. A higher number of deployments is needed to give a more adequate value.
During his analysis of the deep-sea benthos of the Angola Basin Brenke (2005) also
found out that there is no significant difference in the sampling result of the epi- and
supranet. This is supported by the findings in this study. Neither at species level nor at
family level a difference in the composition of net samples could be statistically proven.
However, the sample volume of the epinet (7733 individuals) is distinctly larger than
that of the supranet (2580 individuals).
In comparison to the great box corer that is preferably used for gathering quantitative
data, the EBS covers greater areas and thus samples a larger number of specimens (Tab.
4). The box corer is limited to a small sampling area of usually 0.25 m2 per deployment.
In addition, it creates a blast wave before touching the ground that represses large water
masses and with them many epibenthic organisms. The chance to catch a nearly
DISCUSSION
125
representative extract of the species present seems more likely with an EBS. Due to its
movement, the EBS catches part of the up-churned and repressed sediment and water
masses, and also some organisms fleeing to the frontward direction of the gear.
Tab. 4: Comparison of EBS and box corer deployments during ANDEEP I/II (Hilbig, 2004)
station location depth [EBS] N [EBS] N [GKG]area
analyzed (m2) [EBS]
area analyzed (m2)
[GKG] 42 Drake Passage 3690 m 687 95 2318 0.2 46 Drake Passage 2889 m 1418 166 2232 0.2
131 Weddell Sea 3050 m 217 8 1898 0.1 132 Weddell Sea 2086 m 35 45 1328 0.2 133 Weddell Sea 1123 m 6 55 908 0.1 134 Weddell Sea 4069 m 51 4 2378 0.1 141 South Sandwich Islands 2313 m 478 17 1508 0.1
However, the polychaete community sampled by the EBS seems in its structure very
similar to that of box-corer samples known from former studies in the Southern Ocean.
At ANDEEP I/II site 133 in the Weddell Sea Ampharetidae, Polynoidae, and Syllidae
were among the most abundant families in EBS and in box corer samples (Hilbig,
2004). The polychaete communities also agree well with studies from different world-
wide sites (App.-Tab. 13). Spionidae and Cirratulidae are known to be among the most
abundant polychaete families in deep-sea communities. The same applies to scale
worms as Polynoidae, and to some extent to Ampharetidae, Syllidae, and Opheliidae.
Distinct differences from the expected are found for Pholoididae and Paraonidae. The
Pholoididae are present in remarkably high numbers in some samples. Most findings
originate from stations located around the Antarctic Peninsula, the Drake Passage, and
the South Sandwich Trench. In the Weddell Sea only few Pholoididae are found. The
family Pholoididae is only little studied to date. Therefore an explanation for this
distribution cannot be given. The contribution of Paraonidae to the assemblages is rather
low in the EBS samples. Paraonidae are known to be one of the most abundant
polychaete families in the northern hemisphere (Cosso-Saradin et al., 1998; Glover et
al., 2001; Thistle et al., 1985). The lack of their dominance in the Weddell Sea was
already discovered in box-corer samples from ANDEEP I-II (Hilbig, 2004). Both
studies indicate that Paraonidae are less important in the Southern Ocean deep sea than
DISCUSSION
126
in more temperate regions. A potential relation of these findings to the sampling method
can be neglected.
4.2 Polychaete abundance and family composition
Experiences from deep-sea basins from lower latitudes indicate that polychaetes make
up around 40-60 % of macrobenthic communtities in the deep sea (App.-Tab. 14). This
study predicts a comparably smaller percentage of polychaete contributions to the
samples of the ANDEEP expeditions. In comparison to the Polychaeta (10,635
individuals) peracaridan crustaceans were represented by 22,974 individuals in the EBS
samples of ANDEEP III (Brökeland et al., submitted). An overview of the percentual
invertebrate composition of the ANDEEP expeditions is not yet available. In the AGT
samples the polychaetes contributed 19 % to all invertebrate individuals found (Linse et
al., submitted). It has to be considered that the AGT is not constructed for the sampling
of macrofaunal elements. An undersampling of the polychaetes is thus underlying this
value. However, a reduced contribution of polychaetes in the Southern Ocean deep-sea
benthos is not unexpected. Brandt & Schnack (1999) found an increased dominance of
crustaceans in Arctic Seas near Greenland. On the basis of the recent database (App.-
Tab. 14) a shift in dominance from Polychaeta to Crustacea with increasing latitude
might be observed.
Still, Polychaeta are one of the most flexible and successful invertebrate taxa in the deep
seas. Their presence in large numbers and the finding of many cosmopolitan species in
the ANDEEP samples indicate that the Southern Ocean deep sea is not as isolated in its
faunal composition as the Antarctic shelf (Hilbig, 2004). A faunal exchange with lower
latitude deep-sea basins is imaginable. The Antarctic Circumpolar Current (ACC), that
is known to be one major distributional barrier between the Southern Ocean shelf and
temperate waters, does not reach down into great depths. In contrast, the Antarctic deep
water flows below into lower latitudes and can be identified as far north as the North
Atlantic (e.g., Knox & Lowry, 1977).
The family composition of polychaetes found in the Atlantic sector of the deep Southern
Ocean shows some resemblance to that of other ocean basins (App.-Tab. 13). The
DISCUSSION
127
Spionidae are among the most abundant as are the Cirratulidae. Remarkable is the high
abundance of Syllidae and Hesionidae in this study. Both are common, but usually not
dominant families. The high abundance mainly originates from st. 133-2 (central
Weddell Sea) and 74-6 (Eastern Weddell Sea). Both stations are characterized by high
individual densities that are not found at adjacent sites. Cluster analysis indicates strong
similarities of these stations (Fig. 17). A correlation of the polychaete assemblages of
ANDEEP I-II with environmental factors (after Howe et al. (2004) and Diaz (2004))
suggests that family composition is strongly correlated to sediment grain size at sites.
Comparable sediment parameters might be one explanation for the high similarities of
st. 74-6 (eastern Weddell Sea) and 133-2 (central Weddell Sea). Additionally, Syllidae
are known to often be associated with sponges. Hilbig et al. (2006) found high syllid
abundances on the southeastern Weddell Sea shelf and explained it by strong influences
of epibenthic sponge communities. However, the AGT samples of ANDEEP III did not
show increased numbers of sponges in the eastern Weddell Sea compared to the
remaining stations (Linse et al, submitted). Further sampling is needed to investigate the
unusual high abundance of syllids and hesionids.
Remarkable family compositions are also found for stations 131-3, 121-11 (central
Weddell Sea), and 154-9 (King George Island) where Opheliidae are most abundant. At
station 152-6 near Elephant Island almost exclusively Cirratulidae are present. The
Cirratulidae all belong to one species. Specimens were surrounded by a dense fibrous
material, presumably of organic origin. This is possible evidence for a special food fall
or disturbance that favored the dominance of Cirratulidae at this station. Cirratulidae are
typical pioneer polychaetes. They are able to fast settle in disturbed environment (e.g.,
Borowski & Thiel, 1998; Glover et al., 2001) and become very abundant.
Rather typical deep-sea communities as known e.g. for the Atlantic deep sea are found
at stations 59-5, 81-8, 142-5 (Eastern and central Weddell Sea below 3000 m) and 151-
7 (South Orkney Islands, below 1000 m). Most abundant here are Spionidae,
Cirratulidae (except for st. 151-7), Ampharetidae, as well as Paraonidae. The great
depth range of these stations supports the result of the BIO-ENV-anlysis that depth is a
subordinated factor determining polychaete family composition. Grain size and oxygene
depth (after Howe et al. (2004) and Diaz (2004)) have been identified as more
important. The great variety in family composition in the ANDEEP samples is probably
DISCUSSION
128
due to further ecological factors. Some of the different communities might present
different successive stages of communities recovering from disturbance (e.g., eddies,
iceberg rafting, sudden food input, lack of seasonal food input due to prolonged ice
coverage).
DISCUSSION
129
4.3 Species identification, composition and descriptions of new species
Species identification was mainly carried out using monographs on Southern Ocean
polychaetes (compare chapter 2.2). Original species descriptions were only consulted
when identification with secondary literature were ambiguous. The risk of possible
discontinuities in the naming of species remains when original descriptions are not
regarded. However, the monographs are standard works and a high percentage of
published studies rely on them. Their usage results in species lists of good
comparability.
In this study 155 species from 13 different families were distinguished. The families
belong to four orders/ suborders (Glyceriformia, Nereidiformia, Opheliida, and
Terebellida, taxonomy based on Fauchald, 1977) that differ in their life strategies.
While Glyceriformia and Nereidiformia are vagile hunters, Opheliida are mainly
endobenthic deposit feeders. The Terebellida are tube-dwelling suspension and deposit
feeders. The 13 families make up 47 % of all specimens found in the analyzed samples.
With a percentage of ~ 30 % the number of new species in the samples is comparable to
that known from former studies in the Southern Ocean (Hartman, 1964; 1966;1967;
1978; Hartmann-Schröder & Rosenfeldt, 1988; 1989; 1990; 1991; 1992). Although the
Southern Ocean has been subject to intensified sampling during the last century, this
high amount of new species indicates an undersampling of the area. A percentage of ~
46 % species found were formerly named. All of these have been recorded for the
Southern Ocean before, although not necessarily in the same depths. It remains to verify
that morphologically equal speciemens of different sites are members of one species.
They might also be genetically separated cryptic species. Molecular studies on this topic
are desirable. First attempts for crustacean and mulluscs have been made (e.g., Held,
2003; Held & Leese, 2007; Held & Wägele, 2005; Linse et al., 2007; Raupach &
Wägele, 2006). In addition, Mincks & Glover (pers. comment) are working on Southern
Ocean polychaete samples with special focus on Spionidae.
The 155 species found during this study are included in new identification keys. The
most recent, published monographs including keys for polychaetes of the Southern
Ocean were presented by Hartman (1964; 1966; 1967, 1978). Since then a great number
of new species has been described, but keys including these new species are not
DISCUSSION
130
published to date. The presented keys therefore represent an useful addition to recent
taxonomic literature.
4.3.1 Species composition of families
As mentioned, the Ampharetidae, together with the Terebellidae, are the most speciose
family in the samples. Species are mostly present in low abundances. High abundances
and a large distribution is limited to few species such as Sosanopsis kerguelensis,
Anobothrus pseudoampharete sp. n., Ampharete kerguelensis, and species of the genus
Amphicteis. Consequently, the Ampharetidae do not contribute dominant species to any
station, although they are among the most abundant families at some stations (e.g., 16-
10, 21-7 (South Atlantic), 121-11, 134-4 (Weddell Sea)). The percentage of unknown
and unidentified species within this family lies above 50 %. The weak knowledge of the
ampharetid fauna of the deep Southern Ocean results from an undersampling of the area
and often poor conditions of specimens after sampling.
The drawn picture partially also applies to the Terebellidae. The family is not as
abundant at single stations as Ampharetidae. Its high presence in the sum of all samples
results from high abundances of Polycirrus insignis and Phisidia rubrolineata at station
133-2. The particularities of this station have been discussed above.
The Opheliidae and Scalibregmatidae are both present with median abundance and
median diversity. The Opheliidae show a less even distribution than the
Scalibregmatidae. They are very dominant at some stations (e.g., 131-3, 121-11, 154-9),
although their high abundance is not due to dominance of single species as seen in the
cases discussed above.
The Trichobranchidae, Nephtyidae, Sphaerodoridae, and Glyceridae are present at most
stations. However, only one to a few species are found. The Glyceridae are known to be
a species rare family. In the samples, they are only represented by Glycera kerguelensis.
For the Sphaerodoridae only Sphaerodoropsis parva is found in high abundance. Two
species build the main community of the Nephtyidae (Aglaophamus paramalmgreni and
A. trissophyllus) and the Trichobranchidae (Octobranchus antarcticus and Terebellides
stroemi). While the determination of the nephtyid species is undoubted, it may be
DISCUSSION
131
possible that the two trichobranchid species are clusters of cryptic species.
Trichobranchidae are only distinguished by few characters that are hard to determine in
preserved species. For Terebellides stroemi a subspecies has already been described,
differing in the shape of the uncini (T. stroemi kerguelensis).
The Nephtyidae, as well as the Sphaerodoridae are considered basal polychaetes (see
App.-Fig. 1). The species found are mainly endemic for the Southern Ocean. Therefore,
these species are presumably not invaded species, high abundance of single species
might be a sign of slow evolution rates. Also, a fast reaction to local food input is a
possible explanation.
The families Goniadidae, Nereididae and Sabellariidae are very rare, with low numbers
of species. They seem to play a subordinated role in the Southen Ocean deep sea.
4.3.2 Descriptions of new species
Descriptions of 15 new species were given in this study. In addition, the genus
Ammotrypanella MCINTOSH 1879 was revised. Except for the hesionid genera
Micropodarke and Parasyllidea, all genera were already known for the Southern Ocean.
The formerly classified species of these two genera were only reported from northern
hemisphere waters and southern hemisphere waters close to Australia.
4.3.2.1 Ampharetidae MALMGREN 1866
The only species of the genus Anobothrus LEVINSEN 1884 recorded for the Southern
Ocean were A. gracilis (MALMGREN 1866) and A. patagonicus (KINBERG 1867)
(Hartman, 1966; Parapar & San Martín, 1997). With A. pseudoampharete sp.n. a third
Antarctic species is introduced. The former species A. antarcticus MONRO 1939 has
been excluded from the genus by now, being the type species of a new genus,
Anobothrella antarctica (MONRO 1939). Almost half of all species known for the genus
are reported from the Southern Ocean. Anobothrus pseudoampharete sp.n. clearly
differs from A. gracilis and A. patagonicus by the shape of its palae. These are similar
DISCUSSION
132
to that of Ampharete kerguelensis. Anobothrus gracilis bears very long, thin palae, that
of A. patagonicus are small and similar to the notosetae (Hartman, 1966). The palae of
A. pseudoampharete sp.n. are of median length with a broad base and a suddenly
tapering, long delicate tip.
4.3.2.2 Hesionidae GRUBE 1850
Within the Hesionidae four species are newly described, all belonging to different
genera. The genus Ophiodromus SARS 1861 has formerly been reported for the
Southern Ocean with one species, Ophiodromus comatus (EHLERS 1913). The most
apparent difference between this species and Ophiodromus calligocervix sp.n. is the
presence of eyes in O. comatus. The tentacular cirri, as well as the doral cirri of O.
calligocervix sp.n. are stouter. A distinct articulation of the dorsal cirri as known for O.
comatus could not be discovered.
The genus Amphiduros HARTMAN 1959 was originally believed to consist of five
species. The holotypes of all formerly known species originated from northern
hemisphere waters, specimens from Australia and Papua New Guinea were also
recorded (Pleijel, 2001). Pleijel (2001) revised the genus. He excluded the species
Amphiduros alensis BLAKE & HILBIG 1990 and formed a new genus Amphiduropsis.
The remaining species were synonymized as A. fuscescens (MARENZELLER 1875).
Amphiduros serratus sp.n. is consequently the second species of the genus at present
time. It differs from the type species in the lack of eyes, also the tips of the antennae are
less fine. While A. fuscescens is only reported from warm to temperate waters on both
hemispheres, A. serratus sp.n. seems to be a cold water species of greater depths.
Micropodarke cylindripalpata sp.n. is the second species described for the genus. As
formerly Amphiduros, the genus Micropodarke OKUDA 1938 was revised by Pleijel &
Rouse, 2005. They synonymized the formerly distinguished species M. dubia (HESSLE
1925), M. amemiyai OKUDA 1938, and M. trilobata HARTMANN-SCHRÖDER 1983,
leaving M. dubia the only species. This species has a more or less cosmopolitan
distribution. Records are present in temperate waters from the Pacific and Indic ocean,
records from the Atlantic and polar regions are lacking. Micropodarke cylindripalpata
DISCUSSION
133
sp.n. is clearly different in its distribution pattern, being found in much colder waters
and greater depths. Morphologically the two species differ in the shape of their palps.
The second article of M. dubia is equal to the first while it is clearly thicker and pin-
shaped in M. cylindripalpata sp.n. The chaetal appendages of M. dubia are serrated
while that of M. cylidripalpata sp.n. are smooth.
Parasyllidea delicata sp.n. can be easily recognized by the lack of eyes . Both other
species of this genus, P. humesi PETTIBONE 1961 and P. australiensis HARTMANN-
SCHRÖDER 1980, bear four eyes. The chaetal appendages of P. delicata sp.n. end in a
simple tip while they are bifid in P. australiensis. While P. delicata sp.n. is obviously
adapted to cold waters, P. australiensis is reported for temperate waters of the
Australian coast (Hartmann-Schröder & Hartmann, 1980). P. humesi was described to
live in a commensale relationship associated with Tellina nymphalis (LAMARCK 1818),
a bivalve known for shallow waters (Pettibone, 1961).
4.3.2.3 Opheliidae MALMGREN 1867
The genus Ammotrypanella MCINTOSH 1879 was to date defined by the description of
its only species A. arctica (McIntosh, 1879). According to McIntosh (1879), the main
characters distinguishing it from Ophelina aulogaster (RATHKE 1843) are a bluntly
rounded prostomium, bent silky bristles in the anterior seven to eight segments, median
bristles sunk in lateral grooves, and short caudal segments with strong bristles. Fauchald
(1977) summarized the generic characters of Ammotrypanella. In his monograph he
defines the presence of two ventral cirri on the anal tube as a key feature of the genus.
This character is, however, not present in the type species A. arctica which totally lacks
a ventral cirrus. In contrast, McIntosh (1879) does not mention these ventral cirri. The
finding of three new species of the genus enabled to rewrite the diagnosis and to define
characters more clearly.
The genus shows some plesiomorphic characters known for other opheliid genera such
as Ophelina ÖRSTEDT 1843 and Antiobactrum CHAMBERLIN 1919. These characters are
a long and thin habitus with a ventral groove, a conical prostomium with a palpode, and
the possible presence of an anal tube with or without ventral cirrus.
DISCUSSION
134
Considered autapomorphies for Ammotrypanella are prolonged, bent setae in tufts in
anterior parapodia in combination with a limitation of branchiae to the posterior body
region.
The type species A. arctica has been described from polar and temperate waters.
Whether all records belong to this species has yet to be proven since the genus has been
considered monotypic until now. It is, however, without doubt that the species has a
bipolar distribution. The holotype originates from Arctic waters, the new records from
Antarctic waters (App.-Fig. 14). Records of Hartman and Fauchald (1971) originate
from a transect along the Hudson Canyon to Bermuda. Records of Fauvel (1914) and
Levenstein (1978) do not represent specimens of A. arctica. Although both authors
point out many similarities to the former description of McIntosh (1879), the specimens
described present branchiae in the anterior and median body region. This character
clearly excludes them from the genus Ammotrypanella. It is rather proposed that the
specimens belong to the genus Ophelina.
Comparing all four species of Ammotrypanella, it becomes apparent that the shape of
the anal tube is the most distinguishing character within the genus. The presence of a
ventral cirrus and the shape of the posterior margin of the anal tube occur in various
combinations that clearly differentiate the species (Tab. 5).
Tab. 5: Anal-tube characters of the species of Ammotrypanella MCINTOSH 1879
species posterior margin ventral cirrus
A. arctica numerous fine anal cirri absent
A. cirrosa numerous fine anal cirri present
A. mcintoshi no anal tube no anal tube
A. princessa smooth present
Five of the ten species of the genus Ophelina found in this study are new to science.
Two of them have been classified and described. Except for O. cylindricaudata
(HANSEN 1878), all species formerly known for the Southern Ocean have been found in
the samples proposing that the species of Ophelina have a common distribution, none of
them can be considered rare species. In comparison to the most abundant opheliid
species (O. breviata (EHLERS 1913), O. gymnopyge EHLERS 1908), and O. nematoides
(EHLERS 1913)) the two new species are of rather large size, O. ammotrypanella sp.n. is
DISCUSSION
135
larger than 30 mm in some cases. Both new species are most similar to O. setigera
HARTMAN 1978, sharing a robust body and prolonged anterior chaetae. The anterior
chaetae of both species are distinctly shorter than described for O. setigera. Ophelina
robusta sp.n. in addition bears a robust ventral cirrus on its anal tube.
4.3.2.4 Scalibregmatidae MALMGREN 1867
The new scalibregmatid species, Pseudoscalibregma papilia sp.n. is the 15th
scalibregmatid species known for Subantarctic and Antarctic waters. It can be
distinguished from its most related species P. bransfieldium HARTMAN 1967 by an
extraordinary well development of the dorsal and ventral cirri in the median and
posterior parapodia, and the lack of a clearly apparent nuchal crest dorsally on the
prostomium.
Pseudosaclibregma papilia sp.n. is sampled from depths between 2889 and 3690 m, and
therefore seems to represent a true deep-sea species. All together 12 of the 15 recorded
scalibregmatid species from the Southern Ocean are found in the deep sea (Tab. 6),
proposing that the Southern Ocean inhabits a highly divers scalibregmatid deep-sea
fauna.
4.3.2.5 Sphaerodoridae MALMGREN 1867
The Sphaerodoridae were represented with 50 % of species undescribed in the samples.
Especially the genus Sphaerodoropsis has grown after this study to a total number of 48
named species world wide. Known for the Southern Ocean are eight species including
the formerly described S. arctowskyensis HARTMANN-SCHRÖDER & ROSENFELDT 1988,
S. oculata FAUCHALD 1974, S. parva (EHLERS 1913), S. polypapillata HARTMANN-
SCHRÖDER & ROSENFELDT 1988, and S. pyenos FAUCHALD 1974 (Hartman & Fauchald,
1971; Fauchald, 1974, Hartmann-Schröder & Rosenfeldt, 1988; 1990; 1992). Tab. 7
gives a comparison of the eight species concentrating on the most distinguishing
DISCUSSION
136
Tab. 6: Distribution of scalibregmatid species known for the Southern Ocean (after Blake, 1981; Schüller
& Hilbig, 2007)
Species distribution depth (m)
Ascleirocheilus ashworthi BLAKE 1981 off Elephant Island
west of Antipodes Island
223-2892
384-397
Hyboscolex equatorialis BLAKE 1981 Ecuador, Peru
South Sandwich Trench
intertidal
2258-2313
Axiokebuita millsi POCKLINGTON & FOURNIER 1987 South Sandwich Trench 773-752
Axiokebuita minuta HARTMAN 1967
South Shetland Islands
South Orkney Islands
Antarctic Peninsula
Ross Sea
Weddell Sea
off Elephant Island
South Sandwich Trench
300
593-598
412
589-608
1122-4069
2889-2892
2258-2313
Oligobregma blakei SCHÜLLER & HILBIG 2007 off Elephant Island 2889-2892
Oligobregma collare (LEVENSTEIN 1975)
Drake Passage
Ross Sea
Weddell Sea
Bellinghausen Sea
2889-3806
6070 (?)
1622-4575
2562
Oligobregma hartmanae BLAKE 1981 Weddell Sea 585
Oligobregma notiale BLAKE 1981
Antarctic Peninsula
Ona Basin
Weddell Sea
Budd and Knox Coasts
Ross Sea
18-97
3683-3690
28
909-923
Oligobregma pseudocollare SCHÜLLER & HILBIG 2007
South Sandwic Trench
off Elephant Island
Weddell Sea
773-752
2889-2892
3049-3050
Oligobregma quadrispinosa SCHÜLLER & HILBIG 2007
Ona Basin
Weddell Sea
South Sandwich Trench
3683-3690
3049-4069
2258-2313
Pseudosaclibregma bransfieldium (HARTMAN 1967)
Antarctic Peninsula
Bransfield Strait
Weddell Sea
off Elephant Island
Ona Basin
South Sandwich Trench
Ross Sea
335
769
412-2086
2889-2892
3683-3690
773-752
916
Pseudoscalibregma papilia n.sp.
Ona Basin
off Elephant Island
South Sandwich Trench
3683-3690
2889-2892
2258-2313
Pseudoscalibregma ursapium BLAKE 1981
Ross Sea
Ona Basin
off Elephamt Island
Weddell Sea
2143-2154
3683-3690
2889-2892
3049-3050
Scalibregma inflatum RATHKE 1843 cosmopolitan intertidal-abyssal
Sclerocheilus antarcticus ASHWORTH 1915 Antarctic Peninsula
Bransfield Strait
45-90
210-426
DISCUSSION
137
characters. These are the number of lateral antennae, the number of macrotubercle rows,
and the parapodial structure.
Tab. 7: Differentiating characters of species of the genus Sphaerodoropsis known for the Southern Ocean
(after Fauchald, 1974; Hartmann-Schröder & Rosenfeldt, 1988; 1990; 1992)
species
Longitudinal
macrotubercle
rows (max)
Lateral antennae Parapodial lobes Parapodial
papillae
S. arctowskyensis 7 3, short, oval post- & prechaetal 1 anterobasal
S. distincta 4 2, long postchaetal 1 ventral
S. maculata 4 3, long postchaetal none
S. oculata 4 2, basally swollen prechaetal 1 median
S. parva 4 3 prechaetal up to 8
S. polypapillata 9 3 prechaetal 1 anterior,
1 posterior
S. pyenos 11 2 prechaetal, foliose unknown
S. simplex 4 2 postchaetal 2 dorsal
It becomes apparent that one group of species has four rows of macrotubercles on the
dorsum, the number of rows in the other species is clearly higher. All newly described
species are characterized by the presence of four rows. The number of lateral antennae
is either two or three pairs in this genus. Within the group of species with four tubercle
rows only S. parva and S. maculata sp.n. share the presence of three pairs of lateral
antennae. They can clearly be distinguished by the shape of the parapodia, that of S.
maculata sp.n. lacking papillae, and the presence of numerous colored spots on the
entire surface of S. maculata sp.n. Of all species with two pairs of lateral antennae, S.
distincta sp.n. stands out by the unusual pronouncation of its macrotubercles. These are
greatly enlarged. In contrast, S. simplex sp.n. has small, somewhat flattened
macrotubercles that are unusual for this genus.
With its high number of species, the genus Sphaerodoropsis is the largest among the
Sphaerodoridae. The variability in macrotubercle number and the presence of a large
group of species with four rows suggests that the genus is a generic cluster. A careful
revision of this group might lead to the formation of different genera for the known
DISCUSSION
138
species. Also the number of lateral antennae might play an important role in the
definition of genera.
The new species of the genus Ephesiella, E. hartmanae sp.n., can easily be recognized
by the absence of recurved hooks in the first parapodium. The only species known with
that trait is E. gallardi FAUCHALD 1974. It differs from E. hartmanae sp.n. in having
two kinds of chaetae. E. gallardi is recorded from South Vietnam (Fauchald, 1974),
further species known from the Southern Ocean are E. antarctica, E. mühlenhardtae
HARTMANN-SCHRÖDER & ROSENFELDT 1988, and E. pallida FAUCHALD 1974. Apart
from the difference in lacking recurved hooks in the first parapodia, the parapodial and
chaetal structure of E. hartmanae sp.n. is different from that of these three species. E.
hartmanae sp.n. bears a postchaetal and a prechaetal lobe (unlike E. antarctica), a
papillation of the parapodial lobe is not apparent (unlike E. mühlenhardtae and E.
pallida). Chaetae are composite with more or less smooth appendages (that of E.
mühlenhardtae bear 4 – 6 teeth (Hartmann-Schröder & Rosenfeldt, 1988)) of median
size (short in E. pallida).
DISCUSSION
139
4.4 Polychaete diversity in the Southern Ocean
As mentioned before, the samples of ANDEEP I/II are incomplete. This results in a
reduction of the actual sample size and polychaete abundance for these stations. Due to
the resemblance of the community structure in the epi- and supranet, a significant
reduction in species richness is not expected.
Analysis of the species assemblage matrices indicates that the deep Southern Ocean is
still undersampled. The rising species accumulation plot proposes that the found
number of species is lower than the expected. Continuing the graph to a maximum, a
total number of about 180 species for the 13 analyzed families is expected (Fig. 28).
Due to the great structuring of the Southern Ocean deep sea and the small area sampled
this number is presumably still underpredicted.
Fig. 28: Hypthetical course of the species accumulation plot resulting in a saturation (ANDEEP I-III)
Biodiversity of all stations of the ANDEEP expeditions was calculated with the
Margalef’s Index. Additionally, the stations of ANDEEP III were standardized to
achieve quantitative samples of 1000 m2 sampling area. To this quantitative data the
Shannon Index and Pielou’s Evenness Index were applied. These indices are commonly
used in biodiversity studies of different areas and invertebrate taxa, and comparability
of the data is achieved.
The Margalef’s Index was applied to the species assemblage matrix, as well as to the
family assemblage matrix of ANDEEP I-III. Families are taxonomic artefacts and not
DISCUSSION
140
comparable natural units. Biodiversity and systematic studies on families are therefore
not favorable. However, for the present data analysis of family level is desirable for two
reasons: At family level all polychaetes found during ANDEEP I-III are included while
the species analyzed are limited to 13 families. In addition, polychaete families combine
species of similar life forms and serve as indicators for ecological diversity. The
environmental structure of areas as well as the occurrence of disturbances might
therefore be easier detected by family diversity than species diversity.
At family level diversity is highest on the transect between South Africa and the eastern
Weddell Sea (st. 16-10 to 81-8) and off King George Island on the western side of the
Antarctic Peninsula (st. 153-7 and 154-9). The lowest diversity was found at stations
connecting the eastern Weddell Sea with the sites close to the Antarctic Peninsula (st.
88-8 to 110-8). Also the central Weddell Sea shows comparably low diversities
(ANDEEP III st. 121-11 and 133-2, ANDEEP I/II st. 131-3 – 135-4). A correlation of
low diversities with trawling distance and depth can be neglected. Trawling distances
were greatest on the transect between the eastern and central Weddell Sea, decreased
diversity is thus not due to reduced sample volumes. Stations in the central Weddell Sea
were located between 1123 m (st. 133-3) and 4928 m (st. 88-8) depth, depths of the
Southern Atlantic and eastern Weddell Sea stations ranged from 1047 m (st. 74-6) to
4720 m (st. 16-10). Nevertheless, the result is not unexpected. The analysis of isopod
samples from ANDEEP I/II also resulted in the lowest diversity in the central Weddell
Sea with the exception of st. 131-3 (Brandt et al., 2004). In addition, Hilbig et al (2006)
reported lower polychaete diversity in the deep Weddell Sea than on the south eastern
Weddell Sea shelf and the Antarctic Peninsula region. Brandt et al. (2005) explained
increased species richness for Isopoda in the Antarctic Peninsula region and off the
South Shetland Islands by possible faunal exchanges with South American benthic
communities. This could also apply to the polychaete fauna and lead to increased family
diversity around the Antarctic Peninsula. Influences from the Southern Atlantic might
serve as an explanation for high diversities at stations in the eastern Weddell Sea.
The lowest diversity was found at st. 152-6. More than 94 % of all specimens found,
belong to one cirratulid species. A possible explanation was already given in chapter
4.2.
DISCUSSION
141
At species level the differences between stations are more distinct than at family level.
While the geographical differences found for the family diversity are also recognized at
species level it becomes apparent that some stations are more divers than others. The
most divers stations are st. 42-2 and st. 46-7 in the Drake Passage, st. 74-6 in the eastern
Weddell Sea, st. 150-6 off South Orkney Islands, and st. 153-7 near King George
Island.
The high diversity at stations in the Drake Passage might be due to the joining of
different water masses in this area. The Drake Passage is the southernmost connection
between the Pacific and the Atlantic Ocean. Faunal elements of both oceans are found
in this region. Also, the Antarctic bottom water formed in the Weddell Sea passes the
Drake Passage, before entering the South Atlantic. Some species are found in the Drake
Passage and in the Atlantic or Pacific ocean, but not in the Weddell Sea (e.g.,
Phalacrostemma elegans, Hauchiella tribullata, or Proclea graffii). Other species are
present in the Drake Passage and in the Weddell Sea, and further records from the
Atlantic or Pacific are known. Among those are Ampharete kerguelensis, Amphicteis
gunneri, Ammytropanella arctica, Axiokebuita minuta, Sphaerodoropsis parva, and
Terebellides stroemi. The Drake Passage consequently combines polychaete species
recorded from three ocean basins, explaining the high diversity found.
The remaining stations pointed out above are all located on the continental slope that
begins in depths of approximately 600 m in the Southern Ocean. The continental slope
serves as a direct faunal connection between shelf and abyssal plain. Unlike temperate
oceans with steep temperature gradients, the continental slope of the Southern Ocean
offers a potential habitat for species of great depth ranges from the abyssal plain as well
as from the shelf. The vertical distribution patterns of the species analyzed support the
impression that eurybathy is a common trait among polychaetes in the Southern Ocean
(App.-Fig. 5, see also Hilbig, 2004; Hilbig & Blake, 2006, Hilbig et al. 2006). High
diversity of the slope stations is therefore evidence for the presence of faunal elements
from shelf and deep-sea communities. In addition, sedimentation rates are higher on the
slope than in the abyss. An increase in food input might also account for increased
diversity on the continental slope.
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The Shannon Index applied on the standardized family and species assemblage matrices
of ANDEEP III, supports the impression that the central Weddell Sea is the least divers
region sampled. Within the central Weddell Sea an increase in diversity from eastern
(st. 88-8) to western stations (st. 142-5) can be observed. Hilbig et al. (2006) assumed
that differences found in polychaete communities originate from differences in food
income. The Antarctic Peninsula region is known to be very productive. Primary
production in the Southern Ocean is highest during the austral summer when the ice
cover breaks apart to form the pack ice and a sufficient amount of light is available. A
decrease in diversity from the western to the eastern Weddell Sea sites might be
correlated to a decrease in primary production with increasing distance to the Antarctic
Peninsula.
The highest species diversity is found at st. 150-6 off South Orkney Islands and st. 16-
10 in the South Atlantic. These stations do not have remarkably high species
abundances. The highest species abundances are found at st. 74-6 (eastern Weddell Sea)
and 133-2 (central Weddell Sea). However, the evenness of these stations is lower than
that of st. 16-10 and 150-6 explaining lower diversities. Station 133-2 is characterized
by the dominance of single species (e.g., Phisidia rubrolineata, Polycirrus insignis,
Syllides articulosus, Gyptis incompta); at st. 74-6 Anobothrus pseudoampharete sp.n.,
Sphaerodoropsis parva, and Sphaerosyllis lateropapillata uteae are the most abundant
species found. In general, evenness is not apparently correlated to geographical regions.
Differences in evenness are presumably explained by local differences of sites.
On a general scale, the diversity of the Southern Ocean deep-sea polychaetes in this
study is comparable to that of former studies from the Southern Ocean and other ocean
basins. Hilbig et al. (2006) found diversity values of 1.906 and higher and an evenness
between 0.628 – 0.980 for stations in the eastern Weddell Sea and around the Antarctic
Peninsula. Hilbig and Blake (2006) report polychaete diversities between 2.5 – 3.5
(Shannon Index) and an evenness of 0.5 and upper for shelf to deep-sea stations off the
Gulf of Farallones, north-eastern Pacific. Compared to different invertebrate taxa the
polychaete diversity is equally high or even higher, underlining the importance of
polychaetes for benthic communities. The diversity of isopods from ANDEEP I/II EBS
samples ranged between 1.71 and 3.62 with an evenness of above 0.5 (Brandt et al.,
2004). This supports the common idea that polychaetes are among the most robust
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invertebrate taxa with a very high tolerance towards great depth ranges, extreme
environments, and disturbance (Hilbig & Blake, 2006).
In conclusion, the results give evidence that depth is not a limiting factor for polychaete
diversity in the deep Southern Ocean. A reduced input of food due to a strong
seasonality in primary production and low sedimentation rates, are more likely to
influence the diversity patterns within the central Weddell Sea. In addition, faunal
exchanges with adjacent ocean basins might lead to increased diversities in the eastern
Weddell Sea, the Drake Passage, and the western side of the Antarctic Peninsula.
4.5 Similarities between sampling areas
All cluster analyses clearly differentiate st. 152-6 from all other stations. The
peculiarities of this station have been discussed formerly. It is therefore excluded from
further discussions.
In general, similarities are comparably low, lying below 60 % in all analyses. This
might be due to an undersampling of the area. Further data will presumably result in
more distinct similarity patterns.
Clustering of all samples of ANDEEP I-III at family level and at species level show that
the stations of the Southern Atlantic (st. 16-10, 21-7 and 59-9) and off King George
Island (st. 153-7, 154-9) cluster within stations from the eastern and central Weddell
Sea. Most stations are located in one main cluster. Within this cluster geographical
ranges can be identified. At family level the stations of the eastern Weddell Sea and the
Southern Atlantic are concentrated on one side of the cluster. Stations off King George
Island are closer to the Drake Passage and off the South Orkney Islands on the other
side of the cluster. The central Weddell Sea stations are found on both sides. A similar
ordination is found at species level. A difference is found for the stations off King
George Island that are more similar to the eastern Weddell Sea sites than to the Drake
Passage in this analysis.
Clustering of the standardized species data of ANDEEP III also indicates a strong
resemblance of sites off King George Island and the Southern Atlantic to the eastern and
central Weddell Sea. A similar clustering of stations in the Weddell Sea and Drake
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144
Passage is found in a former study by Hilbig et al. (2006). The material in that study
originated from box corer samples taken during the EASIZ II expedition (1998) to the
eastern Weddell Sea shelf, deep-sea sites in the eastern Weddell Sea and stations around
the Antarctic Peninsula and the Drake Passage. The authors distinguished three different
groups, the Peninsula shelf group (including three sites of the eastern Weddell shelf),
the deep station group and the south-eastern Weddell Sea shelf group.
It has formerly been noticed that stations 74-6 (eastern Weddell Sea) and 133-2 (central
Weddell Sea) share the highest family and species abundances. Clustering and MDS
plots indicate high similarities of these stations. Both stations are located on the
continental slope. However, st. 74-6 lies at the east coast of the Weddell Sea while st.
133-2 is a north-western Weddell Sea site. A faunal overlap of shelf stations from the
eastern Weddell Sea and the Antarctic Peninsula has formerly been reported (Hilbig et
al., 2006). This supports the idea that the sampled area hosts a more or less uniform
faunal assemblage that underlies local variations due to different depths and abiotic
factors. The presence of the similar faunal compositions near the east coast and the west
coast of the Weddell Sea can be explained by two models (Fig. 29):
Fig. 29: Possible ways of polychaete dispersal within the Weddell Sea (Southern Ocean)
DISCUSSION
145
Model one proposes a connection through the Weddell shelf with a distribution along
the coastlines. Dispersal is most likely from east to west due to clockwise currents in the
Weddell Sea. Polychaetes are not very vagile, an active distribution over long distances
against current systems is not probable.
The second possibility is a connection through the deep Weddell Sea. As mentioned
earlier, eurybathy is a common trait among polychaetes in the Southern Ocean.
Submergence of shelf and slope species to deeper waters at the east and the west coast
of the Weddell Sea by down-slope currents is very likely, followed by a subsequent
transport across the Weddell abyssal plain by the Antarctic bottom water. Few
individuals could have been able to emerge to lower depths in both parts of the Weddell
Sea again, in spite of a lack of currents in that direction. Emergence against current
direction has already been reported for isopods in the Southern Ocean (Brandt et al.,
2004). The higher abundance of species in the coastal zones might result from the fact
that the deep sea is not the optimal habitat for submerging shallow water species. They
are able to survive and reproduce there but not as successfully as on the continental
slope and shelf.
Both models are supported by the data of this study. A great number of species are
present at stations 74-6 and 133-2 and are lacking on the Weddell abyssal plain. Among
these are Gyptis incompta, Kefersteinia fauveli, Leaena collaris, Polycirrus insignis,
Sphaerosyllis joinvillensis, Sphaerosyllis lateropapillata uteae, Syllides articulosus, and
Travisia kerguelensis. All of these species occur in comparably high abundance and can
not be considered rare species. A lack of records in the Weddell Sea abyss therefore
indicates absence in that area. Species found in high numbers at stations 74-6 and 133-2
and in lower numbers in the deep Weddell Sea are e.g., Amage sculpta, Amphicteis
gracilis, Axiokebuita minuta, Neosabellides elongatus, Ophiodromus comatus, and
Terebellides stroemi. Regarding this, a connection of the two sites through the abyssal
plain seems possible. The dataset is evidence for emergence but does not prove it. The
presence of species on the shelf of both coastal sides of the Weddell Sea and in the
abyssal plain might also result from distribution along the shelf and submergence of
species into the deep sea.
From the analysis of ANDEEP I/II stations it becomes apparent that the Weddell Sea
stations (st. 131-3, 134-4) group together as do the stations from the Drake Passage (st.
DISCUSSION
146
42-2, 46-7). St. 141-10 is more similar to the Drake Passage than to the Weddell Sea. A
possible explanation of that pattern is the flow of the Antarctic bottom water. The
Antarctic bottom water originates from the Weddell abyssal plain. It enters the Drake
Passage and flows past the South Sandwich Trench into the Southern Atlantic. With it,
faunal elements from the deep Weddell Sea might be transported into the South
Atlantic, decreasing in number with distance. Consequently, the Weddell Sea stations
are more similar to Drake Passage stations than to stations of the South Sandwich
Trench. Furthermore, the South Sandwich Trench and Drake Passage stations might
share some Atlantic and even Pacific elements that are rare in the Weddell Sea.
The similarity analyses show that the Atlantic sector of the Southern Ocean seems to
host a potentially uniform poylchaete fauna. Geographical gradients between eastern
and western sites, as known from former studies (Hilbig et al., 2006) are present.
However, these are only indistinct and based on low similarities. The differences in
polychaete composition found for different station are presumably rather due to local
differences of environmental factors and events of disturbance. Also, it becomes evident
that the Weddell deep sea is not isolated from the Southern Atlantic. A faunal exchange
between the basins occurs, which is not hindered by the Antarctic Circumpolar Current.
The ACC does not reach into these depths and therefore cannot be considered a
distributional barrier for the Antarctic deep-sea fauna (Brandt et al., 2006).
4.6 Influence of environmental factors and species on similarities
The Bray-Curtis Index is based on the presence and abundance of species to determine
the similarities between stations. Some species thus have a greater impact on the
similarities than others. These are not necessarily the most abundant species, but species
that show distinct differences in their distribution within the samples. The species
determined to have the greatest impact in the similarities of the ANDEEP III stations
were Aglaophamus paramalmgreni, Ammotrypanella arctica, Ampharete kerguelensis,
Amphicteis sp. 3, Kefersteinia fauveli, Kesun abyssorum, Micropodarke cylindripalpata
sp.n., Ophelina ammotrypanella sp.n., Ophelina robusta sp.n., Ophiodromus
DISCUSSION
147
calligocervix sp.n., Ophiodromus comatus, Polycirrus insignis, Pseudoscalibregma
papilia sp.n., Sosanopsis kerguelensis, and Travisia kerguelensis.
Among these species five are vagile predators; four of them belong to the Hesionidae.
Syllidae, the most speciose vagile family in the samples, is not among them. The stated
filter feeders are all but one Ampharetidae. The suspension feeders are to equal parts
Opheliidae and Scalibregmatidae. Those families might hence be of high ecological
importance, reacting towards environmental changes not by being present or absent but
by changes in their species composition.
Of all species stated, only Ophiodromus calligocervix sp.n., Polycirrus insignis, and
Travisia kerguelensis are present in depths shallower than 1000 m. Amphicteis sp. 3,
Ophelina robusta sp.n., Polycirrus insignis,and Pseudoscalibregma papilia sp.n. are
only present above 4000 m. The depth ranges of the species are very wide, proposing
that eurybath species have the strongest influence on similarities of stations. This results
in a weakening of the importance of depth as a differentiation factor for stations below
1000 m. This finding is also supported by the BIO-ENV analysis of the ANDEEP I/II
stations. The results show that depths is the most important factor when station 143-1
(774m depth) is included. After the exclusion of this slope station, the grain size and
oxygen availability in the sediment are more likely explanations for station similarities.
Sedimentary parameters regarded in this study are taken from Howe et al. (2004) and
Day (2004). The data originate from the expeditions ANDEEP I/II and were taken
directly at the sample sites. The coarsest sediment was found in the South Sandwich
Trench (st. 141-10 and 143-1) with silty sand (35-40 µm) to pebble of up to 235 µm
grain size. The environment is energetic with stronger currents and higher sedimentation
rates compared to the Drake Passage and Weddell Sea sites. Up to 28 polychaete
families have been found at the sites (st. 141-10), indicating a strongly structured
environment. The polychaete community at these stations is dominated by Spionidae, a
pioneer taxon typical for disturbed environments. Also, vagile hunters and deposit
feeders are present. Because of their higher vagility or burrowing abilities they have an
advantage to suspension feeders on coarse and mobile sediment. Grain sizes in the
Drake Passage are slightly larger than in the Weddell Sea. In the Drake Passage the
highest number of different families has been found (16-33 families per station). As for
the South Sandwich Trench, vagile hunters are present in high abundance. Also
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Spionidae and Cirratulidae, both indicators for disturbances, are among the most
abundant families. Further burrowing deposit feeders and suspension feeders play a
subordinated role. In the finer sediments in the Weddell Sea with slower sedimentation
rates, suspension and deposit feeders dominate the polychaete communities. Pioneer
taxa as Spionidae and Cirratulidae are less abundant. At st. 131-3 the highest number of
families is reported for this region with 18 families. As suggested by the BIO-ENV
analysis, a high correlation between the grain size and the polychaete composition is
observed. Coarse sediment results in highly structured communities dominated by
vagile families. Vagile hunters need hard surfaces to crawl on and approach their prey.
The abundance of pioneer taxa indicates disturbance events. Fine sediments, in contrast,
favor the presence of suspension feeders; these are less vagile and more sensitive
towards fast sedimentation. Also deposit feeders are abundant in fine sediments; they
are mostly burrowing forms that easily move through fine sediments. The general
importance of grain size in contrast to depth was already reported by Bilyard and Carey
(1979) for polychaete compositions in the western Beaufort Sea. It thus seems to be a
global characteristic rather than one specific for the Southern Ocean.
4.7 Taxonomic diversity
Most stations show a taxonomic diversity that lies among or above the expected. The
expected value is calculated based on a masterlist including all species found in this
study. The species accumulation plot has shown that the area is strongly undersampled.
The masterlist is consequently shorter than expected. This might explain why strong
differences to expected AvTDs and VarTDs have not been found for the samples. St.
152-6 has been formerly pointed out to be peculiar, its taxonomic diversity lies far
below the expected. The same applies to st. 135-4. The station is characterized by low
abundances. However, the samples of this station are incomplete; the result is therefore
strongly biased. Surprisingly, st. 131-3 and 121-11 in the central Weddell Sea show
values below the expected although they are among the most speciose stations. This can
be explained by the species composition which is dominated by Hesionidae (only st.
131-3), and species of Oligobregma and Ophelina. At st. 121-11 species of the genera
DISCUSSION
149
Ammotrypanella, Oligobregma, and Ophelina dominate the community. The AvTD is
therefore distinctly lower than expected for stations with comparable species numbers.
Both stations seem to present an environment that favors the presence of Opheliidae and
Scalibregmatidae. Both families probably underwent radiation events in some genera in
the Southern Ocean, among which Ophelina and Oligobregma are very successful. At
the mentioned stations species of both genera co-exist, proposing that conditions are
optimal and interspecific competition is strongly reduced.
DISCUSSION
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4.8 Zoogeography and vertical distribution
4.8.1 Origin of the Southern Ocean deep-sea fauna
During the early Triassic (245 Ma ago) the continent of Antarctica was imbedded in
Gondwana. It had direct connections to the recent South America, South Africa, India
and Australia. During the Jurrasic and the early Cretaceous, Gondwana broke apart and
the continent of Antarctica slowly moved southwards. While it isolated from South
America, Africa and India, the Australian continent was still closely attached to
Antarctica. A first gap between Antarctica and Australia occured during the late
Cretaceous. In the following 50 M years, the gaps between the Antarctic continent and
the other continental plates enlarged and Antarctica reached the approximate southern
position it has today (after Smith et al., 1994). About 30 – 28 M years ago the Drake
Passage opened into deep depths, allowing the ACC to develop (Lawver & Gahagan,
2003; Lawver et al., 1992). Barker (2001) postulated an even younger date of the deep
opening and the origin of the ACC. The glaciation of Antarctica and the cooling of the
deep ocean first started in the Eocene- Oligocene boundary about 34 M years ago
(Barker & Thomas, 2004). It was not abrupt but interrupted by several interglacial
phases (Lear et al, 2000). The cooling of the Weddell Sea and the Antarctic Peninsula
was supported by the Weddell Gyre which transported cold water masses and ice to the
Antarctic Peninsula. The ACC resulted in an isolation of the cold water masses around
the Antarctic continent and supported the glaciation process (Barker & Thomas, 2004).
Although the wind-driven ACC needs a constant deep-sea connection to exist, it does
not reach down to the sea floor. The Weddell Sea Bottom Water slowly flows
northwards beneath the ACC, similar deep-water flows exist in the Pacific and Indic
ocean (Barker & Thomas, 2004).
The development of the Antarctic continent and the Southern Ocean had several
consequences for the fauna and flora on land and in the sea. The extreme cooling
resulted in a great extinction of species. Some species were able to adapt to this extreme
environment and in course of the isolation, underwent numerous radiation events. The
adaptation to cold temperatures on the shelf facilitated a submergence of species into
the deep sea. Submergence helped the species escape from the glaciation of the shelf
DISCUSSION
151
and resulting disturbances from down-slope transport of glaciogenic depris (Thatje et
al., 2005). During the process of submergence species, however, had to adjust to
increased pressure and reduced food input.
The glaciation of the Antarctic continent and its surrounding waters did not have such
dramatic consequences for the deep-sea fauna. The deep sea of the Southern Ocean has
constant temperatures around 2°C and is hence as cold as the deep sea world wide. It is
thus proposed that part of the Southern Ocean deep-sea fauna originates from
submerging shelf species while also relict species occur. Additionally, the ACC does
not isolate the Southern Ocean deep sea from adjacent ocean basins. It is imaginable
that part of the deep-sea fauna secondarily invaded the Southern Ocean from temperate
deep-sea basins.
4.8.2 Vertical distribution
The vertical distribution patterns show that most polychaete species in the Southern
Ocean have wide depth ranges. Species limited to stations above 1000 m are rare. Most
species are found between 1000 and 3000 m. In depths between 2000 and 3000 m, a
faunal break occurs. Below 3000 m species are found that do not occur in shallower
samples. At the same time some species found above 2000 m do not occur below. The
result indicates that eurybathy is a common trait among Southern Ocean deep-sea
polychaetes. At the same time classical shelf and deep-sea faunas exist. Three categories
of polychaetes are found: shelf species, deep-sea species that only occur below 3000 m,
and eurybath species.
The Southern Ocean is characterized by a deep shelf (down to 600-800 m) and lack of
temperature isoclines. It is therefore a common idea that submergence of shelf species
and emergence of deep-sea species is facilitated within most invertebrate groups.
Evidence for the Isopoda is given by Brandt, 1991, Brandt et al., 2004, and Brandt et al.,
2006. The Antarctic shelf fauna is reported to be isolated. It is characterized by high
species richness and a high percentage of endemism that originates from radiation
events after the cooling of the Antarctic continent (Arntz et al, 1997; Barnes & de
Grave, 2000; 2001; Brandt et al., 2004; Crame, 2000; de Broyer et al., 2002; Gray,
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152
2001) ~ 34 Mio years ago (Barker & Thomas, 2004). This model seems to fit to many
invertebrate taxa such as Isopoda.
For Isopoda, large depth ranges as found for Polychaeta are reported (Brandt et al.,
2006). However, recent studies have presented new evidence that eurybathy is not a
special characteristic of polychaetes in the Southern Ocean but is rather common among
polychaetes world wide (Brandt et al.; 2006; Hilbig & Blake, 2006; Hilbig et al. 2006).
This supports the idea that the Southern Ocean deep-sea poylchaete community does
not originate only from relict and submerging shelf species. The far vertical distribution
is evidence for a secondary invasion of the Antarctic deep sea from adjacent ocean
basins such as the Pacific or Atlantic.
4.8.3 Global distribution patterns
A faunal exchange between the Southern Ocean deep sea and adjacent deep-sea basins
is strongly supported by the global distribution patterns found for some polychaete
species in this study. No difference could be determined between different depth zones.
Species communities from the continental slope (1000 to 3000 m) show as far a
distribution as species from the deep sea. Many species found are spread far beyond the
subantarctic region while others are more or less regionally restricted.
Distinct differences between the families analyzed can be seen. The species of the
Goniadidae, Glyceridae, Nereididae, and Nephtyidae show distribution patterns centred
in the Southern Ocean. However, only few species were found for these families. The
genera, in contrast to the species are reported for all ocean basins world wide (e.g.,
several species of Goniada, Glycera, and Nereis are known for the Northern Atlantic
(e.g., Hartman, 1950; Hartmann-Schröder, 1996)). These records might support the
theory of a faunal exchange between temperate deep-sea basins and the Southern
Ocean. After invasion of species of these genera into the Southern Ocean, speciation
could have resulted in the evolution of locally restricted species. A second possibility is
that the found species originate from relict species. It is not determinable if species of
the same genera already lived on the Antarctic plates and speciation took place during
glaciation, or if they invaded the Southern Ocean after the cooling.
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153
The hesionid species found are also only reported from the Southern Ocean. They are
present in higher abundance and with more species than the formerly mentioned
families. It might be a general characteristic of Hesionidae that most species do not have
wide distribution ranges. However, the genera of the Hesionidae found are also known
for other ocean basins – Amphiduros fuscescens (MARENZELLER 1875), Parasyllidea
humesi PETTIBONE 1961, and P. australiensis HARTMANN-SCHRÖDER 1980 are reported
from temperate waters of both hemispheres (Hartmann-Schröder, 1980; Pettibone,
1961; Pleijel, 2001). As for the formerly stated families, the sampled hesionid species
therefore either evolved from relict species, or from species that invaded the Southern
Ocean after glaciation.
A remarkable pattern is observed for the Trichobranchidae. Only two named species
were found in the samples. While Terebellides stroemi is a cosmopolitan species,
Octobranchus antarcticus is endemic for the Southern Ocean. The Trichobranchidae
resemble the Ampharetidae and Terebellidae in their morphology and feeding strategy.
All three families are tube dwelling, selective suspension feeders that use long tentacles
to catch particles from the water column and the sediment surface. The tubes are usually
partially imbedded in the sediment, and the animals are known to leave them
sometimes. The three families also have similar distribution patterns. Locally restricted
and cosmopolitan species are found in the Southern Ocean. An explanation might be
found in the great variances in reproductive strategies. Wilson (1991) reports brooding
and free swimming larvae (lecithotrophy or direct development) for the Ampharetidae
and Terebellidae. Also encapsulation of embryos in gelatinous masses is reported for
some Terebellidae and for the cosmopolitan trichobranchid T. stroemi. Free swimming
larvae with direct development presumably settle close to their place of hatching. The
larvae need to feed shortly after hatching. However, dispersal over large distances
aggravates feeding. In addition, larvae are put at risk of mechanical disturbance and
declined conditions. Lecithotrophy of larvae and encapsulation of embryos in contrast
permit widespread dispersal of species. The cosmopolitan species Melinna cristata e.g.,
has free-living lecitotrophic larvae (Wilson, 1991). Especially the encapsulation in
gelatinous masses might be an advantage when the brood is drifted away. It might
protect embryos from mechanical disturbance and changing abiotic factors during
development, and reduce embryo mortality.
DISCUSSION
154
Similar distributional patterns are found for Scalibregmatidae and Opheliidae. For both
families species with wide-spread distributions and locally restricted species are found.
The two families are very similar in their life strategies and presumably closely related
(Rouse & Fauchald, 1997). Their high abundance in samples world wide (e.g., Glover et
al., 2001) suggests a high adaptive ability to different environments. The Opheliidae
have free-living larval stages (Wilson, 1991). Distribution of larvae by bottom-near
currents and settlement of species in different localities are likely. Wide distribution
areas for the Opheliidae are not surprising. Larval development of the Scalibregmatidae
is not known. However, the cosmopolitan distribution of some species suggests
distribution by larval dispersal. Both families are burrowing forms. A second way of
species distribution might be dispersal of adult specimens. Benthic storms (eddies) and
turbidity currents (Scheltemar, 1994) are common phenomena in the Southern Ocean
deep sea. Adult specimens might be carried away with the sediment. Substrate
heterogeneity is supposedly less distinct on the abyssal plains than in shallow waters.
Thus specimens surviving the dispersal might be able to settle at different localities
afterwards.
Rouse & Fauchald (1997) on basis of cladistic analyses proposed that Opheliidae and
Scalibregmatidae are basal forms. The high age of these families would be an additional
explanatory factor for the wide distribution of species. Because of the high number of
endemic species in the Southern Ocean, these probably originate from local speciation
events. A subsequent invasion into the Pacific Ocean via the Humboldt Current and into
the Atlantic via the Antarctic bottom water can be postulated for some species (e.g.,
Hyboscolex equatorialis, Ammotrypanella arctic, and the genus Axiokebuita).
The similarities in life strategies might have an influence on the similarities of
distribution patterns – a phenomenom already discussed for Ampharetidae,
Terebellidae, and Trichobranchidae. The difference of Opheliidae and Scalibregmatidae
to the formerly discussed is the fact that the classification of species within these
families is not as reliable as in the other families. The genera Travisia and Kesun were
formerly believed to be opheliid genera (e.g., Hartman, 1966; Fauchald, 1977). Persson
& Pleijel (2005) however stated that Travisia is a scalibregmatid genus. The close
resemblance of Kesun to Travisia leads to the assumption that Kesun is also a
scalibregmatid genus. In addition, the identification of Axiokebutia millsi, A. minuta and
DISCUSSION
155
Scalibregma inflatum might not be correct for all records. This leads to a general
problem of the analysis of global distribution patterns. Potential misidentifications and
discontinuities in naming in the historical data can bias the analyses. Since most species
in this study are well studied and only records from taxonomic publications are
considered, the risk of bias is reduced.
The species of the Sphaerodoridae are primarily restricted to the Subantarctic region.
Only Sphaerodoropsis parva has been reported for the Southern Pacific and the South
Atlantic. This species is the most abundant sphaerodorid species in the samples. It
supposedly originates from the Southern Ocean and invades lower latitudes via the
Humboldt Current and the Antarctic bottom water. The Sphaerodoridae are vagile,
omnivorous forms that contribute around 2 – 5 % to all polychaete species in different
areas of continental slopes and abyssal plains, with low abundances (< 1 % of
individuals). Within the family free-living larvae are found, the existance of
lecithotrophic larvae is predicted (Fauchald, 1977). The combination of free larval
stages (potentially high dispersal rates), presence of only few speciose genera in the
Southern Ocean, and narrow distribution ranges at species level indicates that the
Sphaerodoridae are highly adapted to certain environmental niches. Within the Southern
Ocean several speciation events might have occurred resulting in the high number of
endemic species.
The Syllidae are also vagile, omnivorous forms. Most species are brooders (Wilson,
1991). Brooding is reported for almost all genera found in the samples, thus dispersal of
species through larval stages is very unlikely. Most species show restricted
distributions; especially the species of the genus Sphaerosyllis seem to be endemic for
the Weddell Sea and originate from local radiation events. The reproduction strategy of
the genus Typosyllis is unknown. This genus contributes two of the four cosmopolitan
syllid species found. An explanation for a cosmopolitan distribution of syllid species
cannot be given; a secondary invasion of the Southern Ocean by these species is likely.
The results are congruent with the common observation that the distribution potential of
species is related to the reproduction and environmental robustness of species. Free
larval stages contribute to the dispersal of species (e.g., Grahame & Branch, 1985).
Lecithotrophic larvae with prolonged planctonic stages might favor great dispersal even
in waters of low nutrient content. They are often found in the deep sea (e.g., Herring,
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2002). In addition, adults of species with high environmental robustness might be able
to settle in new localities after being carried away with sediment transport. Brooding
seems to be more common in species of narrow distribution as found for the Syllidae.
The cosmopolitan distribution of some species is an indication for secondary invasion
of these species into the Southern Ocean after glaciation. Typosyllis variegata,
Terebellides stroemi, Melinna cristata, and Scalibregma inflatum for example are
reported for depths from the shelf down into the deep sea in oceans world wide. Their
distribution patterns are patchy. Distinct distributional directions from the Southern
Ocean into adjacent basins as found for some Scalibregmatidae or Sphaerodoropsis
parva are not apparent (App.-Fig. 16-17). The Southern Ocean is of a comparably
young age. In addition, polychaetes are not very mobile; active distribution of species is
therefore very slow. Distribution by larval dispersal, in contrast, is dependent on ocean
currents. A distribution originating from the deep Southern Ocean should be traceable
by species records along large ocean currents, such as the Humboldt Current or the
Antarctic bottom water.
In contrast a possible distribution from the Southern Ocean northwards into more
temperate waters can be recognized for some species. Sphaerodoropsis parva is
supposedly invading the South Pacific via the Humboldt Current. Also a distribution
along the west coast of Africa into the Angola basin can be seen. Braniella palpata is
also recorded from sites along the Humboldt Current. The opheliid genus
Ammotrypanella is presumably of Antarctic origin as well. Although A. arctica is
reported also in the Northern Atlantic, the additional three species newly described for
that genus are only known for the Southern Ocean until now. Records from the Pacific
or Indic sector of Antarctica are not known for A. arctica. It is assumed that the species
originates from the Southern Ocean and reached into the Northern Atlantic from there.
Evidence for local radiation events is found in the families Syllidae (Sphaerosyllis),
Opheliidae (Ophelina), Terebellidae (Pista), Scalibregmatidae (Oligobregma), and
Sphaerodoridae (Sphaerodoropsis).
The presented results predict that the deep Southern Ocean polychaete fauna is
composed of relict species, species submerging from the Antarctic shelf, endemics that
evolved during local radiation events, and species secondarily invading the deep sea
from the Atlantic, Pacific, and Indic oceans. Additionally, an ongoing dispersal of deep-
DISCUSSION
157
sea species into northern deep-sea basins takes place, supported by currents and water
masses such as the Humboldt Current and the Antarctic deep water.
Based on the analysis of the global distribution patterns longitudinal gradients are not
apparent within the Southern Ocean. Although local differences in species composition
exist, a general pattern for the origin of species is not seen. The eastern Weddell Sea
stations do not contain more Atlantic species while the stations of the Antarctic
Peninsula are not apparently more related to the Pacific. However, during this study
only a low number of stations were analyzed. The distances between stations were low
in comparison to the size of the Southern Ocean. A sampling of the Pacific sector of the
Southern Ocean might result in a more differentiated biogeographic picture.
SUMMARY
158
5 Summary In course of this study the Polychaeta of the Antarctic deep sea have been analyzed. The
sample material originated from the expeditions ANDEEP I-III to the Atlantic sector of
the Southern Ocean in early spring 2002 and 2005. Samples were taken with an
epibenthic sledge constructed at the Ruhr-University of Bochum. Analyses of
biodiversity and community structure of the Polychaeta were carried out with the
program PRIMER v 6.1.6. In addition, the global distribution patterns of named species
were reconstructed to gain insight into the possible origin of the Antarctic deep-sea
benthos.
In the samples 14,176 individuals of 47 families were found. Thirteen families were
identified to species level, 155 species were distinguished. A percentage of 30 % of
species were new to science indicating an undersampling of the Antarctic deep sea.
During this study 18 species were described, three descriptions have already been
published. Additionally, identification keys for all species found were constructed.
Calculations of species richness (Margalef’s Index) and diversity (Shannon Index and
Pielou’s Evenness) indicate high polychaete diversities in the Antarctic deep sea
compared to deep-sea basins world wide. Within the sampling area the central Weddell
Sea is characterized by the lowest diversity. Possible explanation is a lack of influences
of faunal elements from adjacent deep-sea basins in comparison to the Drake Passage
and the Antarctic Peninsula (possible Pacific influences), as well as the eastern Weddell
Sea (possible Atlantic influences).
Similarity analyses result in one main cluster that only presents minor differences
between sampling areas. Within this cluster evidence for an east-west gradient in
polychaete composition is given. On a regional scale the sampled sector of the Southern
Ocean supposedly hosts a potentially homogenous polychaete community. Different
influences of adjacent deep-sea basins, however, result in a longitudinal gradient
between eastern and western stations. For local differences between stations sediment
grain size is suggested as a factor of major influence. Depth seems to play a
subordinated role. Vertical distribution patterns show a high percentage of eurybath
species that are found on the upper continental slope as well as on the abyssal plains.
Additionally, the presence of a classical deep-sea fauna becomes evident.
SUMMARY
159
The global distribution patterns of named species present distinct differences between
the analyzed families. These might be based on reproduction strategies and habitat
preferences.
Based on distribution charts dispersal ways of polychaetes along ocean currents become
apparent. The Humboldt Current supports the dispersal of species into the Pacific, the
Antarctic bottom water that into the Atlantic. The found east-west gradient in species
composition, as well as the finding of cosmopolitan species indicate that some
polychaete species sampled originate from outside the Southern Ocean deep sea. The
Antarctic deep sea is consequently not isolated, as known for the Antarctic shelf. Within
the Southern Ocean there is evidence for submergence and emergence of polychaete
species. Additionally, local radiation events originating from relict species are
suggested.
ZUSAMMENFASSUNG
160
5.1 Zusammenfassung
Im Rahmen der vorliegenden Arbeit wurden die Polychaeta der antarktischen Tiefsee
untersucht. Das analysierte Probenmaterial stammt von den Expeditionen ANDEEP I-
III, welche im Winter und Frühjahr 2002 und 2005 im atlantischen Sektor des
Südozeans durchgeführt wurden. Die Probennahme erfolgte mit einem an der Ruhr-
Universität Bochum konstruierten Epibenthosschlitten. Analysen zur Biodiversität und
Gemeinschaftsstruktur der Polychaeten wurden mit Hilfe des Programms PRIMER v
6.1.6 durchgeführt. Zudem wurden die globalen Verbreitungsmuster benannter Arten
rekonstruiert, um Rückschlüsse auf die Besiedlungsgeschichte der antarktischen Tiefsee
zu erlangen.
Die Proben umfassten 14.176 Individuen aus 47 Familien. Hiervon wurden 13 Familien
auf Artniveau bestimmt, 155 Arten wurden unterschieden. Der Anteil neuer Arten lag
bei ca. 30 %, was für eine Unterbeprobung der antarktischen Tiefsee spricht. Im Laufe
der vorliegenden Studie wurden 18 Arten neu beschrieben. Drei Neubeschreibungen
wurden vorzeitig publiziert. Zudem wurden Bestimmungsschlüssel für alle in den
Proben gefundenen Arten erstellt.
Die Berechnung des Artenreichtums (Margalef’s Index) und der Diversität (Shannon
Index und Pielou’s Evenness) weisen auf eine hohe Polychaetendiversität der
antarktischen Tiefsee im Vergleich zu Tiefseebecken weltweit hin. Innerhalb des
beprobten Areals ist das zentrale Weddell Meer durch die geringste Diversität
charakterisiert. Eine mögliche Erklärung bietet der fehlender Einfluss von
Faunenelementen aus benachbarten Tiefseebecken verglichen mit der Drake Passage
und der Antarktischen Halbinsel (potentiell pazifische Einflüsse), sowie dem östlichen
Weddell Meer (potentiell atlantische Einflüsse).
Die Ähnlichkeitsanalysen resultieren in einem großen Cluster, der nur geringe
Unterscheide zwischen den Regionen aufzeigt. Innerhalb des Clusters gibt es Hinweise
auf einen Ost-West-Gradienten in der Polychaetenzusammensetzung. Auf regionaler
Ebene ist daher zu vermuten, dass der beprobte Sektor des Südozeans eine potentiell
homogene Polychaetengemeinschaft beheimatet. Unterschiedliche Einflüsse
angrenzender Tiefseebecken des Atlantiks und Pazifiks resultieren jedoch in einem
longitudinalen Gradienten zwischen östlichen und westlichen Stationen. Für lokale
ZUSAMMENFASSUNG
161
Unterschiede zwischen Stationen konnte die Sedimentgröße der beprobten Flächen als
mögliche Erklärung ausgemacht werden. Die Tiefe scheint für die
Artenzusammensetzung der Polychaeten nur eine untergeordnete Rolle zu spielen. Die
vertikalen Verbreitungsmuster weisen auf einen hohen Anteil eurybather Arten hin, die
sowohl auf dem oberen Kontinentalabhang zu finden sind, als auch auf den
Tiefseeebenen. Zudem ist die Existenz einer klassischen Tiefseefauna nachgewiesen.
Die globalen Verbreitungsmuster benannter Arten zeigen starke Unterschiede zwischen
den analysierten Familien auf, welche auf Unterschieden in der Reproduktionsstrategie
und der Habitatspräferenz basieren könnten.
Anhand der erstellten Verbreitungskarten konnten klare Verbreitungswege einzelnen
Polychaetenarten entlang ozeanischer Strömungen nachvollzogen werden. Der
Humboldtstrom dient vermutlich der Verbreitung von Arten in den Pazifik, das
Antarktische Tiefenwasser der in den Atlantik. Sowohl der Ost-West-Gradient in der
Artenzusammensetzung als auch der Fund vieler Kosmopoliten weisen darauf hin, dass
einige Polychaeten ihren Ursprung außerhalb des Südozeans haben. Die antarktische
Tiefsee ist demnach nachweislich nicht isoliert wie der antarktische Schelf. Innerhalb
des Südozeans gibt es Hinweise auf Submergenz und Emergenz von Polychaetenarten.
Des Weiteren werden lokale Radiationsereignisse ausgehend von Reliktarten vermutet.
REFERENCES
162
6 References
Adkins, J.F., McIntyre, K. & Schrag, D.P. (2002). The salinity, temperature, and
δ18 O of the glacial deep ocean. Science Vol. 298. 1769-1773
Alongi, D.M. (1992). Bathymetric patterns of deep-sea benthic communities from
bathyal to abyssal depths in the western South Pacific (Solomon and Coral
Seas). Deep-Sea Research 39(3/4). 549-565
Arntz, W.E., Gutt, J. & Klages, M. (1997). Antarctic marine biodiversity. In:
Battaglia, B., Valencia, J. & Walton, D.W.H. (eds). Antarctic communities:
species, structure and survival. Cambridge, Cambridge University Press. 3-14
Arwidsson, I. (1899). Studien über die Familien Glyceridae und Goniadidae. Bergens
Museums Aarbog 1898. 1-69
Augener, H. (1912). Beiträge zur Kenntnis verschiedener Anneliden und Bemerkungen
über die Nephtys-Arten und deren epitoke Formen. Archiv für Naturgeschichte
Berlin, Jahrbuch, vol. 78 A, Heft 10. pp. 162-212, 2 pls.
Augener, H. (1932). Antarktische und antiboreale Polychaeten nebst einer Hirudinee.
Scientific Results of the Norwegian Antarctic Expedition 1927-28. Norske
Videnskaps Akademi i Oslo, vol. 9. pp. 1-86, 1 pl., 10 figs.
Barker, P.F. (2001). Scotia Sea regional tectonic evolution: implications for mantle
flow and palaeocirculation. Earth Science Reviews 55. 1-39
Barker, P.F. & Thomas, E. (2004). Origin, signature and palaeoclimatic influence of
the Antarctic Circumpolar Current. Earth Science Reviews 66. 143-162
Barnes, D.K.A. & de Grave, S. (2000). Biogeography of southern polar bryozoans. Vie
et Milieu 50(4). 261-273
Barnes, D.K.A. & de Grave, S. (2001). Ecological biogeography of southern polar
encrusting faunas. Journal of Biogeography 28. 359-365
Basford, D.J., Eleftheriou & A., Raffaelli, D. (1989). The epifauna of teh North Sea
(56° - 61°N). Journal of the Marine Biological Association of the United
Kingdom 69. 387-407
REFERENCES
163
Benham, W.B. (1921). Australasian Antarctic Expedition 1911-1914 under the
leadership of Sir Douglas Mawson. Scientific Reports of the Australasian
Antarctic Expedition (Series C), Zoology and Botany, vol. 6, pl. pp. 1-128, pls.
5-10
Benham, W.B. (1927). Polychaeta. British Antarctic (Terra Nova) Expedition, 1910.
History Report, Zoology, vol. VII, no. 2. pp. 47-182, 6 pls., 200 figs
Berkman, P.A. (1992). The Antarctic marine ecosystem and humankind. Reviews in
Aquatic Sciences 6(3/4). 295-333
Bilyard, G.R. & Carey, Jr., A.G. (1979). Distribution of Western Beaufort Sea
polychaetous annelids. Marine Biology 54. 329-339
Blake, J.A. (1975). The larval development of the polychaeta from the northern
California coast. III. Eighteen species of Errantia. Ophelia 14. 23-84
Blake, J.A. (1981). The Scalibregmatidae (Annelida, Polychaeta) from Southern
America and Antarctica collected chiefly during the cruises of the R/V ANTON
BRUUN, R/V HERO and USNS ELTANIN. Proceedings of the Biological
Society of Washington 94(4). 1131-1162
Blankensteyn, A. & Lana, P. da Cunha (1986). Anelidos Poliquetos des Expedicoes
Antarcticas Brasilieras de 1982/1983; 1983/1984 e 1984/1985. Neritica, Pontal
du Sol, PR 1(3). 49-77
Bleidorn, C., Vogt, L. & Bartholomaeus, T. (2003a). A contribution to sedentary
polychaete using 18S rRNA sequence data. Journal of Zoological Systematics
and Evolutionary Research 41. 186-195
Bleidorn, C., Vogt, L. & Bartholomaeus, T. (2003b). New insights into polychaete
phylogeny (Annelida) inferred from 18S rDNA sequence data. Molecular
Phylogenetics and Evolution 29. 279-288
Borowski, C. & Thiel, H. (1998). Deep-sea macrofaunal impacts of a large-scale
physical disturbance experiment in the Southeast Pacific. Deep-Sea Research II
45. 55-81
Brandt, A. (1991). Zur Besiedlungsgeschichte des antarktischen Schelfes am Beispiel
der Isopoda (Crustacea, Malacostraca). Berichte zur Polarforschung 98. 1-240
Brandt, A. (1992). Origin of Antarctic Isopoda (Crustacea, Malacostraca). Marine
Biology 113. 415-423
REFERENCES
164
Brandt, A. (1993). Composition, abundance, and diversity of peracarid crustaceans on
a transect of the Kolbeinsey Ridge, north of Iceland. Polar Biology 13. 565-576
Brandt, A., Brökeland, W., Brix, S. & Malyutina, M. (2004) Diversity of Southern
Ocean deep-sea Isopoda (Crustacea, Malacostraca)- a comparison with shelf
data. Deep-Sea Research II, 51. 1753-1768.
Brandt, A., Ellingsen, K.E., Brix, S., Brökeland, W. & Malyutina, M. (2005).
Southern Ocean deep-sea isopod species richnesss (Crustacea, Malacostraca):
influence of depth, latitude and longitude. Polar Biology 28. 284-289
Brandt, A., De Broyer, C., De Mesel, I., Ellingsen, K.E., Gooday, A.J., Hilbig, B.,
Linse, K., Thomson, M.R.A. & Tyler, P.A. (2007). The biodiversity of the
deep Southern Ocean benthos. Philosophical Transactions of the Royal Society
B 362. 39-66
Brandt, A. & Piepenburg, D. (1994). Peracarid crustacean assemblages of the
Kolbeinsey Ridge, north of Iceland. Polar Biology 14. 97-105
Brandt, A. & Schnack, K. (1999). Macrofaunal abundance at 79°N off East
Greenland: opposing data from epibenthic-sledge and box-corer samples. Polar
Biology 22. 75-81
Brattegard, T. & Fossa, J.H. (1991). Replicability of an epibenthic sampler. Journal
of the Marine Biological Association of the UK 71. 153-166
Brenke, N. (2005). An epibenthic sledge for operations on marine soft bottom and
bedrock. Marine Technology Society, 39(2). 13-24.
Breton, S., Dufresne, F., Desrosiere, G. & Blier, P.U. (2003). Population structure of
two northern hemisphere polychaetes, Neanthes virens and Hediste diversicolor
(Nereididae), with different life-history traits. Marine Biology 142(4). 707-715
Brey, T. & Clarke, A. (1993). Population dynamics of marine benthic invertebrates in
Antractic and subantarctic environments: are there unique adaptations? Antarctic
Science 5(3). 253-266
Brökeland, W., Choudhury, M. & Brandt, A. (submitted). Composition, abundance
and distribution of Peracarida from the Southern Ocean deep sea. Deep Sea
Research II.
REFERENCES
165
Campbell, B.J., Jeanthon, C., Kostka, J.E., Luther III, G.W. & Cary, S.C. (2001).
Growth and phylogenetic properties of novel bacteria belonging to the epsilon
subdivision of the Proteobacteria enriched from Alvinella pompejana and deep-
sea hydrothermal vents. Applied and Environmental Microbiology 67(10). 4566-
4572
Cazaux, C. (1969). Études morphologique du développement larvaire d’annélides
polychetes (Bassin d’Arcachon). II. Phyllodocidae, Syllidae, Nereidae. Archives
de Zoologoe expérimentale et générale 110. 145-202
Cazaux, C. (1972). Développement larvaire d’annélides polychetes (Bassin
d’Arcachon). Archives de Zoologoe expérimentale et générale 113. 71-108
Clark, R.B. (1961). The origin and formation of the Heteronereis. Biological Reviews,
Cambridge Philosophical Society 36. 199-236
Clark, R.B. (1964). Dynamics of metazoan evolution. The origin of coelom and
segments. Claredon Press, Oxford.
Clark, R.B. (1969). Systematics and Phylogeny: Annelida, Echiura, Sipuncula. In:
Chemical Zoology, Vol. 4. Annelida, Echiura, Sipuncula (eds Florkin, M. &
Scheer, B.T.). Academic Press New York. 1-68
Clark, R.B. (1956). Capitella capitata as a commensal, with a bibliography of
parasitism and commensalism in the polychaetes. Annals and Magazine of
Natural History, 9. 433-448, 2 tables
Clarke, K.R. & Warwick, R.M. (2001). Changes in marine communities: an approach
to statistical analysis and interpretation, 2nd edition. PRIMER-E Ltd: Plymouth
Conway Morris, S. & Peel, J.S. (1995). Articulated halkieriids from the lower
Cambrian of north Greenland and their role in early postome evolution.
Philosophical Transactions of the Royal Society of London. Series B 347. 305-
358
Cosso-Sarradin, N., Sibuet, M., Paterson, G.L.J. & Vangriesheim, A. (1998).
Polychaete diversity at tropical Atlantic deep-sea sites: environmental effects.
Marine Ecology Progress Series 165. 173-185
Crame, J.A. (2000). Evolution of taxonomic diversity gradients in the marine realm:
evidence from the composition of recent bivalve faunas. Palaeontology 26(2).
188-214
REFERENCES
166
Dales, R.P. (1950). The reproduction and larval development of Nereis diversicolor
O.F. Müller. Journal of the Marine Biological Association of the United
Kingdom 29. 321-360
Dales, R.P. (1951). Notes on the reproduction and early development of the cirratulid
Tharyx marioni. Journal of the Marine Biological Association of the United
Kingdom 30. 113-117
Day, J.H. (1964). A review of the family Ampharetidae (Polychaeta). Annals of the
South African Museum 48. 97-121
Dayton, P.K., Mordida, B.J. & Bacon, F. (1994). Polar marine communities.
American Zoologist 34. 90-99
Deacon, G.E.R. (1933). A general account of the hydrology of the South Atlantic
Ocean. Discovery Reports 7. 171-238
Deacon, G.E.R. (1937). The hydrology of the Southern Ocean. Discovery Reports 15.
1-24
de Broyer, C., Jazdzewski, K. & Dauby, P. (2002). Biodiversity patterns in the
Southern Ocean: Lessons from Crustacea. In: Huiskes, A.H.L., Gieskes, W.W.C.,
Rozema, J., Schorno, R.M.L., van der Vies, S.M. & Wolff, W.J. (eds). Antarctic
Biology in a global context. Backhuys Publishers, Leiden, Netherlands. 210-214
Diaz, R.J. (2004). Biological and physical processes structuring deep-sea surface
sediments in the Scotia and Weddell Seas, Antarctica. Deep-Sea Research II 51.
1515-1532
Ehlers, E. (1868). Die Borstenwürmer, nach systematischen und anatomischen
Untersuchungen dargestellt. Leipzig. 269-748
Ehlers, E. (1897). Polychaeten. Hamburg, Magalhaenische Sammelreise. Friedrichsen
& Co.1-148
Ehlers, E. (1900). Magellanische Anneliden gesammelt während der schwedischen
Expedition nach den Magellansländern. Königliche Gesellschaft der
Wissenschaft zu Göttingen. Nachrichten Mathematisch-Physikalische Klasse. pp.
1-18
REFERENCES
167
Ehlers, E. (1901). Die Polychaeten des magellanischen und chilenischen Strandes. Ein
faunistischer Versuch. Festschrift zur Feier 150jährigen Bestehens der
königlichen Gesellschaft der Wissenschaft zu Göttingen. Abhandlung Mathisch-
Physikalische Klasse, Berlin, Wiedmannsche Buchhandlung. pp. 1-232, pls. 1-25
Ehlers, E. (1908). Die bodensässigen Anneliden aus den Sammlungen der deutschen
Tiefsseeexpedition auf dem Dampfer Valdivia 1898-99. Wissenschaftliche
Ergebnisse der Deutschen Tiefsee-Expeditionen auf dem Dampfer Valdivia
1898-1899, 16(1). 1-168
Ehlers, E. (1912). National Antarctic Expedition. Natural History, vol. 6, Zoology,
British Museum, London. pp. 1-32, 3 pls.
Ehlers, E. (1913). Die Polychaetensammlung der deutschen Südpolar-Expedition 1901-
1903. Deutsche Südpolar-Expedition, vol. 13, Heft 4. pp. 379.598, pls. 26-46
Fage, L & Legendre, R. (1927). Péches planctoniques à la lumière, effectuées à
Banyuls-sur-Mer et à Concarneau. I. Annélides polychètes. Archives de Zoologie
expérimentale et générale 67. 23-222
Fauchald, K. (1972). Benthic poylchaetous annelids from deep water off western
Mexico and adjacent areas in the eastern Pacific Ocean. Allan Hancock
Monographs in Marine Biology, 7. 1-575, 69 pls.
Fauchald, K. (1974). Sphaerodoridae (Polychaeta: Errantia) from world-wide
areas. Journal of Natural History 8. 257-289
Fauchald, K. (1977). The Polychaete worms. Definitions and keys to the Orders,
Families and Genera. Natural History Museum of Los Angeles County Science
Series 28. 1-188
Fauchald, K. & Rouse, G.W. (1997). Polychaete systematics: past and present.
Zoologica Scripta 26. 71-138
Fauvel, P. (1914). Annélides Polychètes non pélagique non provenant des campagnes
de l’Hirondelle et de la Princesse-Alice (1885-1910). Résultats de campagnes
scientifiques accomplies sur son yacht par Albert 1er Prince souverain de
Monaco. Fascicule 46. 432pp, 31pl
Fauvel, P. (1916). Annélides polychètes des Iles Falkland recuellies par M. Rubert
Vallentin Esqre (1902-1910). Archives de Zoologie Expérimentalle et Génerale,
Paris, vol. 55. pp. 417-482, pls. 8, 9
REFERENCES
168
Fauvel, P. (1936). Polychètes expédition antarctique Belgica. Résultats du voyage de la
Belgica en 1897-1899, sur le commendament de A. de Gerlache de Gomery, 46
pp., 1 pl., 4 figs.
Fauvel, P. (1941). Annélides polychètes de la Mission du Cape Horn (1882-1883).
Bulletin du Muséum National d’HistoireNnaturelle Paris, Sér. 2, vol. 13. 272-
298
Fauvel, P. (1951). Mission du batiment polaire Commandant-Charcot. Recoltes faites
en Tene Adélie (1950) par Paul Tchernia. Annélides polychètes. Bulletin du
Muséum National d’Histoire Naturelle Paris, Sér. 2, vol. 22. 753-773, fig. 1
Fütterer, D.K., Brandt, A. & Kock, K-H (2003). The expedition ANTARKTIS-
XIX/3-4 of the Research Vessel POLARSTERN in 2002. Introduction and
Summary. Berichte zur Polar- und Meeresforschung 470. 5-6
Gibbs, P.E. (1971). A comparative study of reproductive cycles in four polychaete
species belonging to the family Cirratulidae. Journal of the Marine Biological
Association of the UK 51. 745-771
Gillet, P. & Dauvin, J.C. (2000). Polychaetes from the Atlantic seamounts of the
Southern Azores: biogeographical distribution and reproductive patterns.
Journal of the Marine Biological Association of the UK 80. 1019-1029
Glover, A., Paterson, G., Bett, B., Gage, J., Sibuet, M., Sheader, M. & Hawkins, L.
(2001). Patterns in polychaete abundance and diversity from the Madeira
Abyssal Plain, northeast Atlantic. Deep-Sea Research I 48. 217-236
Gnanadesikan, A. & Hallberg, R.W. (2000). On the relationships of the Circumpolar
Current to Southern Hemisphere winds in coarse-resolution ocean models.
Journal of Physical Oceanography vol. 30. 2013-2034
Grahame, J. & Branch, G.M. (1985). Reproductive patterns of marine invertebrates.
Oceanography and Marine Biology. An Annual Review 23. 373-398.
Gravier, C. (1906a). Annélides polychètes recuellies par l’expédition antarctique
francaise (Syllidiens). Bulletin du Muséum National d’Histoire Naturelle Paris,
vol. 12. pp. 283-290
Gravier, C. (1906b). Sur le Annélides polychètes recuellies par l’Expédition
Antarctique Francaise (Hesioniens, Phyllodociens, Nereidiens, Euniciens).
Bulletin du Muséum National d’Histoire Naturelle Paris, vol. 12. pp. 386-391
REFERENCES
169
Gravier, C. (1906c). Sur le Annélides polychètes recuellies par l’Expédition
Antarctique Francaise (Aphroditiens, Amphinominiens, Flabelligeriens,
Maldaniens, Ampharetiens). Bulletin du Muséum National d’Histoire Naturelle
Paris, vol. 12. pp. 535-540
Gravier, C. (1907a). Annélides polychètes expédition antarctique francaise. Paris,
Masson Cie. Pp. 1-75, 5 pls., 46 text figs.
Gravier, C. (1907b). Sur le Annélides polychètes recuellies par l’Expédition
Antarctique Francaise (Terebelliens, Serpuliens). Bulletin du Muséum National
d’Histoire Naturelle Paris, vol. 13. pp. 46-52
Gravier, C. (1911a). Annélides polychètes recuellies par la seconde expédition
antarctique francaise (1908-1910). Diexième expédition antarctique francaise,
vol. 1. pp. 1-165, 12 pls.
Gravier, C. (1911b). Expédition antarctique francaise du Pourquoi Pas?, dirigée par le
Dr. J.B. Charcot (1908-1910). Espéces nouvelles d’Annélides polychètes.
Bulletin du Muséum National d’Histoire Naturelle Paris, vol. 17. pp. 310-316
Gray, J.S. (2001). Antarctic marine benthic biodiversity in a world-wide latitudinal
context. Polar Biology 24(9). 633-641
Grube, A.E. (1877). Die von der Gazelle mitgebrachten Anneliden, zu denen noch zwei
von Dr. Buchholz gesammelte kommen. Akademie der Wissenschaften, Berlin,
Monatsberichte. 509-554
Haaland, B. & Schram, T.A. (1983). Larval development and metamorphosis of
Ophiodromus flexuosus (delle Chiaje) (Hesionidae, Polychaeta). Sarsia 67. 107-
118
Hartley, J.P. (1985). The re-establishment of Amphicteis midas (Gosse, 1855) and
redescription of the type material of A.gunneri (M. Sars, 1835) (Polychaeta:
Ampharetidae). Sarsia 70. 309-316
Hartman, O. (1936). New species of polychaetous annelids of the family Nereidae
from California. Proceedings of the United States National Museum Vol. 83 No.
2994. 467-480
Hartman, O. (1950). Polychaetous annelids. Goniadidae, Glyceridae and Nephtyidae.
Allan Hancock Pacific Expeditions 15(1). 1-181, 3 figures, plates 1-19
REFERENCES
170
Hartman, O. (1952). The marine annelids of the U.S. Navy Antarctic Expedition 1947-
48. Journal of the Washington Academy of Science, vol. 42. pp. 231-237, 1 pl.
Hartman, O. (1953). Non-pelagic Polychaeta of the Swedish Antarctic Expedition
1901-1903. Further Zoological Results of the Swedish Antarctic Expedition
1901-1903, vol. IV, no. II. Pp. 1-83, 21 figs., 1 chart
Hartman, O. (1964). Polychaeta Errantia of Antarctica. Antarctic Research Series 3. 1-
131, 39 pls., 1 map
Hartman, O. (1966). Polychaeta Myzostomida and Sedentaria of Antarctica. Antarctic
Research Series 7. 1-158, 46 pls., 5 charts
Hartman, O. (1967). Polychaetous annelids collected by the USNS ELTANIN and
Staten Island cruises, chiefly from Antarctic Seas. Allan Hancock Monographs
in Marine Biology, 2. 1-387, 51 pls.
Hartman, O. (1971). Abyssal polycahetous annelids from the Mozambique Basin off
Southeast Africa, with a compendium of abyssal polychaetous annelids from
world.wide areas. Journal of the Fisheries Research Board of Canada, 28. 1407-
1428
Hartman, O. (1978). Polychaeta from the Weddell Sea Quadrant, Antarctica. Paper 4
in: Biology of the Antarctic Seas VI, Antarctic Research Series, vol. 26. 125-223
Hartman, O. & Fauchald, K. (1971). Deep-water benthic polychaetous annelids off
New England to Bermuda and other North Atlantic areas. 2nd Allan Hancock
Monography in Marine Biology, 6. 1-327, 34 pls., 1 chart
Hartmann-Schröder, G. (1965). In Hartmann-Schröder, G. & Hartmann, g. (1965).
Zur Kenntnis des Sublitorals der chilenischen Küste unter besonderer
Berücksichtigung der Polychaeten und Ostracoden. ibd., 62. 1-384. Hamburg
Hartmann-Schröder, G. (1996). Annelida, Borstenwürmer, Polychaeta. Die Tierwelt
Deutschlands 58. 1-594, fig. 1-191
Hartmann-Schröder, G & Hartmann, G. (1980). Zur Kenntnis des Eulitorals der
australischen Küsten unter besonderer Berücksichtigung der Polychaeten und
Ostracoden (Teil 4 und 5). Mitteilungen des Hamburgerischen Zoologischen
Museums und Instituts 77. 41-110
REFERENCES
171
Hartmann-Schröder, G. & Rosenfeldt, P. (1988). Die Polychaeten der „Polarstern“-
Reise ANT III/2 in die Antarktis 1984. Teil 1: Euphrosinidae bis Chaetopteridae.
Mitteilungen des Hamburgerischen Zoologischen Museums und Instituts 85. 25-
72
Hartmann-Schröder, G. & Rosenfeldt, P. (1989). Die Polychaeten der „Polarstern“-
Reise ANT III/2 in die Antarktis 1984. Teil 2: Cirratulidae bis Serpulidae.
Mitteilungen des Hamburgerischen Zoologischen Museums und Instituts 86. 65-
106
Hartmann-Schröder, G. & Rosenfeldt, P. (1990). Die Polychaeten der „Walther
Herwig“-Reise 68/1 nach Elephant Island (Antarktis) 1985. Teil 1: Aphroditidae
bis Cirratulidae. Mitteilungen des Hamburgerischen Zoologischen Museums und
Instituts 87. 89-122
Hartmann-Schröder, G. & Rosenfeldt, P. (1991). Die Polychaeten der „Walther
Herwig“-Reise 68/1 nach Elephant Island (Antarktis) 1985. Teil 2: Acrocirridae
bis Sabllidae. Mitteilungen des Hamburgerischen Zoologischen Museums und
Instituts 88. 73-96
Hartmann-Schröder, G. & Rosenfeldt, P. (1992). Die Polychaeten der „Polarstern“-
Reise ANT V/1 in die Antarktis 1986. Teil 1: Euphrosinidae bis Iphitimidae.
Mitteilungen des Hamburgerischen Zoologischen Museums und Instituts 89. 85-
124
Held, C. (2003). Molecular evidence for cryptic speciation within the widespread
Antarctic crustacean Ceratoserolis trilobitoides (Crustacea, Isopoda). In:
Huiskes AHL, WWC Gieskes, J Rozema, RML Schorno, SM van der Vies and WJ
Wolff (eds) Antarctic biology in a global context. Backhuys Publishers, Leiden.
135-139
Held, C., Leese, F. (2007). The utility of fast evolving molecular markers for studying
speciation in the Antarctic benthos. Polar Biology 30(4). 513-521
Held, C., Wägele, JW. (2005). Cryptic speciation in the giant Antarctic isopod
Glyptonotus antarcticus (Isopoda: Valvifera: Chaetiliidae). Sci Mar 69. 175-181
Herpin, R. (1925). Recherches biologiques sur la reproduction et le développement
quelques Annélides Polychètes. Bulletin de la Société des Sciences Naturelles de
l'ouest de Frances 5. 1-250, 6 plates, 128 figs.
REFERENCES
172
Herring, P. (2002). The Biology of the Deep Ocean. Biology of Habitats (Crawley,
M.J., Little, C., Southwood, T.R.E. & Ulfstrand, S., eds). Oxfort University
Press. 314 pp
Hessle, C. (1917). Zur Jenntnis der terebellomorphen Polychaeten. Zoologiska Bidrang
fran Uppsala, vol. 5. pp. 39-248, pls. 1-5, 66 figs.
Hessler, R.R. & Jumars, P.A. (1974). Abyssal community analysis from replicate box
cores in the central north Pacific. Deep-Sea Research 21. 185-209
Hessler, R.R. & Sanders, H.L. (1967). Faunal diversity in the deep-sea. Deep-Sea
Research 14. 65-78
Hilbig, B. (2004). Polychaetes of the deep Weddell and Scotia Seas- composition and
zoogeographical links. Deep-Sea Research II 51. 1817-1825
Hilbig, B. & Blake, J.A. (2006). Deep-sea Polychaete communities in the northeast
Pacific ocean off the Gulf of the Farallones, California. Bulletin of Marine
Science 78(2). 243-269
Hilbig, B., Gerdes, D. & Montiel, A. (2006). Distributional patterns and biodiversity in
polychaete communities of the Weddell Sea and Antarctic Peninsula area
(Southern Ocean). Journal of the Marine Biological Association of the United
Kingdom 86. 711-725
Holthe, T. (1986). Polychaeta terebellomorpha from the northern Norwegian sea and
the polar sea, with descriptions of Mugga bathyalis sp.n. and Ymerana
pteropoda gen. and sp.n. Sarsia 71(3-4). 227-234
Howe, J.A., Shimmield, T.M. & Diaz, R. (2004). Deep-water sedimentary
environments of the northwestern Weddell Sea and South Sandwich Islands,
Antarctica. Deep-Sea Research II 51. 1489-1514
Hutchings, P.A. (1998). Biodiversity and functioning of polychaetes in benthic
sediments. Biodiversity and Conservation 7. 1133-1145
Hutchings, P.A., Ward, T.J., Waterhouse, J.H. & Walker, L. (1993). Infauna of
marine sediments and seagrass beds of Upper Spencer Gulf near Port Pirie,
South Australia. Transactions of the Royal Society of Australia 117. 1-15
REFERENCES
173
Jördens, J., Struck, T. & Purschke, G. (2004). Phylogenetic inference regarding
Parergordilidae and Hrabeiella periglandulata (‚Polychaeta’, Annelida) based
on 18S rDNA, 28S rDNA and COI sequences. Journal of Zoological Systematics
and Evolutionary Research 42. 270-280
Jumars, P.A. & Hessler, R.R. (1976). Hadal community structure: implications from
the Aleutian Trench. Journal of Marine Research 34(4). 547-560
Kinberg, J.G.H. (1858-1910). Annulater. En: Kongliga Svenska Fregatten Eugenies
Resa omkring jorden under befäl af C. A. Virgin aren 1851-1853. Vetenskapliga
lakttagelser pa Konung Oscar den Förstes befallning utgifna af K. Svenska
Vetenskapsakademien. Almquist & Wiksells, Stockholm, Zoologi, 3: 1-78.
Kinberg, J.G.H. (1866). Annulata Nova. Öfversigt af Förhandlingar Konglia
Vetenskaps Akademiens, 22. 167-179
Knox, G.A. (1962). Antarctic polychaetes from Mawson, Mac Robertson Land.
Transactions of the Royal Society of New Zealand, Zoology, vol. 1, no. 27. 343-
348
Knox, G.A. & Lowry, J.K. (1977). A comparison between the benthos of the Southern
Ocean and the North Polar Ocean with special reference to the Amphipoda and
the Polychaeta. Proceedings of the Polar Oceans Conference Montreal, 1974.
423-462
Kojima, S. (1998). Paraphyletic status of Polychaeta suggested by phylogenetic
analysis based on amino acid sequences of elongation factor-1 alpha. Molecular
Phylogenetics and Evolution 9(2). 255-261
Kröncke, I. & Türkay, M. (2003). Structural and functional aspects of the benthic
communities in the deep Angola Basin. Marine Ecology Progress Series, 260.
43-53
Kruskal, J.B. (1964). Multidimensioning scaling by optimizing goodness of fit to a
nonmetric hypothesis. Psychometrika 29. 1-27
Lawver, L.A. & Gahagan, L.M. (2003). Evolution of Cenozoic seaways in the circum-
Antarctic region. Palaeogeography, Palaeoclimatology and Palaeoecology 198.
11-38
REFERENCES
174
Lawver, L.A., Gahagan, L.M. & Coffin, M.F. (1992). The development of
palaeoseaways around Antarctica. In: Kennett, J.P & Warnke, D.A. (Eds). The
Antarctic Palaeoenvironment: A perspective on global change. AGU Antarctic
Research Series Vol. 56. 7-33
Lear, C.H., Elderfield, H. & Wilson, P.H. (2000). Cenozoic deep-sea temperatures
and global ice volumes from Mg/Ca in foraminiferal calcite. Science 287. 269-
272
Levenstein, R. (1964). Polychaetous annelids of the families Terebellidae and
Trichobranchidae from the Antarctic and Subantarctic. Biological results of the
Sovetskoi Antarktisheskoi Expeditii (1955-1958). Exploration of the Marine
Fauna II. Pp. 168-184, figs. 1-5 [in Russian]
Levenstein, R. (1978). Annelida (Polychaeta) from the deep waters of the Pacific
region of the Antarctic. Trudy Instituta Okeanologii Imeni P.P. ivova Akademik
Nauk 113. 73-87
Licher, F. (1999). Revision der Gattung Typosyllis Langerhans, 1879 (Polychaeta:
Syllidae). Morphologie, Taxonomie und Phylogenie. Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft 551. 1-336
Linse, K., Brandt, A., Bohn, J., Danis, B., de Broyer, C., Heterier, V., Hilbig, B.,
Janussen, D., López González, P.J., Schüller, M., Schwabe, E., Thomson,
M.R.A. (submitted). Macro- and megabenthic communities in the abyssal
Weddell Sea (South Atlantic). Deep-Sea Research II
Linse, K., Cope, T., Lörz, A-N & Sands, C. (2007). Is the Scotia Sea a centre of
Antarctic marine diversification? Some evidence of cryptic speciation in the
circum-Antarctic bivalve Lissarca notorcadensis (Arcoidea: Philobryidae).
Polar Biology 30(8). 1059-1068
Mackie, A.S.Y. & Oliver, G. (1993). The macrobenthic infauna of Hoi Ha Wan and
Tolo Channel, Hong Kong. In: Proceedings of the first International Marine
Biology Workshop: The marine fauna and flora of Hong Kong and Southern
China. (B. Morton ed). Hong Kong University Press. 657-674
Mackintosh, N.A. (1946). The Antarctic convergence and the distribution of surface
temperatures in Antarctic waters. Discovery Reports XXIII. 177-212
REFERENCES
175
Matsumotu, K., Lynch-Stieglitz, J. & Anderson, R.F. (2001). Similar glacial and
holocene Southern Ocean hydrography. Paleoceanography vol. 16. 1-10
Mc Hugh, D. (1997). Molecular evidence that echiurans and pogonophorans are
derived annelids. Proceedings of the National Academy of Science of the United
States of America 94. 8006-8009
McIntosh, W.C. (1879). An account of the petrological, botanical and zoological
collections made in Kerguelen’s Land and Rondriguez during the transit of
Venus expeditions carried out by order of her Majesty’s government in the years
1874-75. Marine Annelida. Royal Society of London, Transactions on
Philosophy, vol. 168. pp. 258-263, pls. 15
McIntosh, W.C. (1885). Report on the Annelida Polychaeta collected by H.M.S.
Challenger during the years 1873-76. Challenger Reports 12. 1-554, pls. 1-55,
1A-39A
McIntosh, W.C. (1910). A monograph of the British annelids. Polychaeta. Syllidae to
Ariciidae. Ray Society of London, 2(2). 233-524, pl. 51-56, 71-87
Monro, C.C.A. (1930). Polychaete worms. Discovery Reports, vol. 2. pp. 1-222, figs.
1-91
Monro, C.C.A. (1936). Polychaete worms II. Discovery Reports, vol. 12. pp. 59-198,
figs. 1-33
Monro, C.C.A. (1939). Polychaeta. B.A.N.Z. Antarctic Research Expedition, 1929-
1931. Reports – Report Series of Biology (Zoology and Botany),B.A.N.Z
Antarctic Research Expedition vol. 4, pt. 4. pp. 87-156, 28 figs.
Moore, J.K., Abbott, M.R. & Richman, J.G. (1999). Location and dynamics of the
Antarctic polar front from satellite sea surface temperature data. Journal of
Geophysical Research vol. 104. 3059-3073
Parapar, J. & San Martín, G. (1997). “Sedentary” polychaetes of the Livingston
Island shelf (South Shetlands, Antarctica), with the description of a new species.
Polar Biology. 502-514
Pearse, J.S., McClintock, J.B. & Bosch, I. (1991). Reproduction of the Antarctic
benthic marine invertebrates: tempo, modes and timing. American Zoologist 31.
65-80
REFERENCES
176
Persson, J. & Pleijel, F. (2005). On the phylogenetic relationships of Axiokebuita,
Travisia and Scalibregmatidae (Polychaeta). Zootaxa 998. 1-14
Pettibone, M.H. (1957). Endoparasitic polychaetous annelids of the family Arabellidae
with descriptions of new species. Biological Bulletin. Marine Biological
Laboratory, Woods Hole, 113(1). 170-187, 5 figures
Pettibone, M.H. (1961). New species of polychaete worms from the Atlantic Ocean,
with a revision of the Dorvilleidae. Proceedings of the Biological Society of
Washington, Vol. 74. 167-186
Pleijel, F. (2001). Revision of Amphiduros (Gyptini, Hesionidae, Polychaeta). Ophelia
54. 15-27
Pleijel, F. & Rouse, G. (2005). A revision of Micropodarke (Psamanthini, Hesionidae,
Polychaeta). Journal of Natural History 39(17). 1313-1325
Pocklington, P. & Fournier, J.A. (1987). Axiokebuita millsi, new genus, new species,
(Polychaeta, Scalibregmatidae) from Eastern Canada. Bulletin of the Biological
Society of Washington,7. 108-113
Ramsay, L.N.G. (1914). Polychaeta of the family Nereidae, collected by the Scottish
National Antarctic Expedition (1902-1904). Transactions of the Royal Society of
Edinburgh, vol. 50. pp. 41-48, pl. 3
Raupach, M.J. & Wägele, J.W. (2006). Distinguishing cryptic species in Antarctic
Asselota (Crustacea: Isopoda)- a preliminary study of mitochonrial DNA in
Acanthaspidia drygalskii. Antarctic Science 18(2). 191-198
Reish, D.J. (1954). The life history and ecology of the polychaetous annelid Nereis
grubei (Kinberg). Allan Hancock Foundation Publications. Occasional Paper
14. 1-75
Reish, D.J. (1957). The lide history of the polychaetous annels Neanthes caudata (delle
Chiaje), including a summary of development in the family Nereidae. Pacific
Science 11. 216-228
Riser, N.W. (1987). Observations on the genus Ophelia (Polychaeta: Opheliidae) with
the descriptions of a new species. Ophelia, 28(1). 11-29
Rosbaczylo, N. & Canete, J.I. (1993). A new species of scale-worms, Harmothoe
commensalis (Polychaeta: Polynoidea), from mantle cavities of two Chilean
clams. Proceedings of the Biological Society of Washington 106(4). 666-672
REFERENCES
177
Rouse, G.W. & Fauchald, K. (1995). The articulation of annelids. Zoologica Scripta
24. 269-301
Rouse, G.W. & Fauchald, K. (1997). Cladistics and polychaetes. Zoological Scripta
26. 139-204
Rouse, G. & Pleijel, F. (2003). Problems in polychaete systematics. Hydrobiologia
496. 175-189
Rouse, G. & Pleijel, F. (vol. eds.) (2006). Reproductive Biology and Phylogeny of the
Annelida. Vol. 4. Science Publishers, Enfield, NH, USA. 688 pp
San Martín, G. & Parapar, J. (1997). „Errant“ polychaetes of the Livingston Island
shelf (South Shetland Islands, Antarctica), with description of a new species.
Polar Biology 17. 285-295
Schäfer, W. (1972). Ecology and palaeoecology of marine environments. The
University of Chicago Press, Chicago. 1-568
Scheltema, R.S. (1994). Adaptations for reproduction among deep-sea benthic
molluscs: an appraisal for the existing evidence. In: Young, C.M., Eckelbarger,
K.J. (eds), Reproduction, Larval Biology, and Recruitment of the Deep-Sea
Benthos. 44-75
Schüller, M. & Hilbig, B. (2007). Three new species of the genus Oligobregma
(Polychaeta, Scalibregmatidae) from the Scotia and Weddell Seas (Antarctica).
Zootaxa 1391. 35-45
Smith, K.L., Kaufmann, R.S. & Baldwin, R.J. (1994). Coupling of near-bottom
pelagic and benthic processes atabyssal depth in the eastern North Pacific.
Oceanology, Limnology and Oceanography 39. 1101-1118
Stolte, H.A. (1932). Untersuchungen über Bau und Funktion der Sinnesorgane der
Polychaetengattung Glycera Sav. Zeitschrift für Naturwissenschaften, Jena, 140.
421-538, 10 plates
Stössel, A. & Kim, S.-J. (1998). The impact of Southern Ocean ice in a global ocean
model. Journal of Physical Oceanography vol. 28. 1999-2018
Struck, T., Hessling, R. & Purschke G. (2002). The phylogenetic position of the
Aeoloosomatidae and Parergodrilidae, two enigmatic oligochaete-like taxa of the
‘Polychaeta’, based on molecular data from 18S rDNA sequences. Journal of
Zoological Systematics and Evolutionary Research 40. 155-163
REFERENCES
178
Svavarsson, J., Brattegard, T. & Strömberg, J.O. (1990). Distribution and diversity
pattern of asselote isopods (Crustacea) in the deep Norwegian and Greenland
Seas. Progress in Oceanography 24. 297-310
Thatje, S., Hillenbrand, C-D & Larter, R. (2005). On the origin of Antarctic marine
benthic community structure. Trends in Ecology and Evolution 20(10). 534-540
Thistle, D., Yingst, J.Y. & Fauchald, K. (1985). A deep-sea benthic community
exposed to strong bottom currents on the Scotia Rise (Western Atlantic). Marine
Geology 66. 91-112
Thorson, G. (1946). Reproduction and larval development of Danish marine bottom
invertebrates with special reference to planctonic larvae in the Sound (Øresund).
Meddelelser fra Kommissionen for Danmarks Fiskeri- Og Havundersøgelser,
Serie: Plankton 4. 1-523
Treadwell, L.A. (1898). The cell lineage of Podarke obscura. Zoological Bulletin,
Boston, 1(4). 195-203
Ugland, K.I., Gray, J.G. & Ellingsen, K. (2003). The species-accumulation curve and
estimation of species richness. Journal of Animal Ecology 72. 888-897
Uschakov, P.V. (1952). Bathypelagische und Tiefsee-Polychaeten vor Kamtschatka
(Pazifischer Ozean). Exploration of the Marine Fauna, 1(9). 129-189 [in
Russian]
Ushakov, P.V. (1974). Fauna of the USSR. Polychaetes. Volume I. Polychaetes of the
suborder Phyllodocidiformia of the Polar Basin and the Northwestern part of the
Pacific. Families Phyllodocidae, Alciopidae, Tomopteridae, Typhloscolecidae
and Lacydoniidae. Akademiya Nauk SSSR. 1-259 [translation from russian]
Wesenberg-Lund, E. (1949). Polychaetes of the Iranean Gulf. Danish scientific
Investigations in Iran 1944/49, 4. 247-400
Wesenberg-Lund, E. (1961). Polychaeta, Errantia. Reports of the Lund University
Chile. Expedition 1948-49, 43. Lunds universitets årsskrift. Andra avdelningen,
Medicin samt matematiska och naturvetenskapliga ämnen, N.F., 57(12). 1-137
Westheide, W. (1967). Monographie der Gattungen Hesionides Friedrich und
Microphthalmus Meczinkow (Polychaeta, Hesionidae). Zeitschrift für
Morphologie und Ökologie der Tiere 61. 1-159
REFERENCES
179
Westheide, W. (1970). Zur Organisation, Biologie und Ökologie des interstitiellen
Polychaeten Hesionides gohari Hartmann-Schröder (Hesionidae). Mikrofauna
des Meeresbodens 3. 101-135
Westheide, W. (1974). Interstitielle Fauna von Galapagos. XI. Pisionidae, Hesionidae,
Pilargidae, Syllidae (Polychaeta). Mikrofauna des Meeresbodens 44. 1-146, 63
figures
Westheide, W. (1997). The direction of evolution within the Polychaeta. Journal of
Natural History 31. 1-15
Westheide, W., McHugh, D., Purschke, G. & Rouse, G. (1999). Systematization of
the Annelida: different approaches. Hydrobiologia, 402. 291-307
White, W.B. & Peterson, R.G. (1996). An Antarctic circumpolar wave in surface
pressure, wind, temperature and sea-ice extend. Reprinted from Nature vol. 380.
11pp.
Willey, A. (1902). Polychaeta. Report on the collections of natural history made in the
Antarctic regions during the voyage of the Southern Cross. London, vol. 12.
262-283, pls. 41-46
Wilson, D.P. (1932). The development of Nereis pelagica Linnaeus. Journal of the
Marine Biological Association of the United Kingdom 18. 203-217, 12 figures
Wilson, W.H. (1991). Sexual reproductive modes in polychaetes: classification and
diversity. Bulletin of Marine Science 48. 500-516
APPENDIX- FIGURES
180
7 Appendix
7.1 Figures
App.-Fig. 1: Basic classification of Articulata (A/Pwr analysis) after Rouse & Fauchald, 1997
A I/II
553
499497
263
193
PholoididaeSphaerodoridaeSpionidaeOpheliidaeScalibregmat idae
AIII
1399
13071294
1016
668
SpionidaeHesionidaeSyllidaePolynoidaeTerebellidae
A B
App.-Fig. 2A-B: Most abundant families of ANDEEP I-III stations
APPENDIX- FIGURES
181
st 41-3
13
10
9
32
OnuphidaeM aldanidaePholoididaeAmpharetidaeSphaerodoridae
st 42-2
207
84
78
64
42
SpionidaeSphaerodoridaePholoididaeM aldanidaeCirratulidae
C D
st 46-7
367
336
97
8273
SphaerodoridaePholoididaeOpheliidaeLumbrineridaeCirratulidae
st 131-1
11124
19
11 10
OpheliidaeScalibregmat idaeSpionidaeCirratulidaeHesionidae
E F
st 132-2
166
3
3
PolynoidaeScalibregmat idaeAcrocirridaeHesionidae
st 133-3
3
1
1
1
PolynoidaeAmpharetidaeScalibregmatidaeSyllidae
G H
App.-Fig. 2C-H: Most abundant families of ANDEEP I-III stations
APPENDIX- FIGURES
182
st 134-4
8
7
7
7
6
Ampharet idaeHesionidaeOpheliidaeScalibregmat idaeSpionidae
st 135-4
3517
64
SpionidaeGlyceridaePolynoidaeFauveliopsidae
I J
st 141-10
130
58
57
39
25
PholoididaeSpionidaePolynoidaeHesionidaeFauveliopsidae
st 143-1
161
76
72
3122
SpionidaeLumbrineridaeScalibregmat idaeGlyceridaeSphaerodoridae
K L
st 16-10
36
2924
12
12
12
SpionidaeOpheliidaeAmpharet idaeFauveliopsidaeHesionidaePolynoidae
st 21-7
33
2017
11
9
9
SpionidaeFauveliopsidaeAmpharet idaeHesionidaeFlabelligeridaeParaonidae
M N
App.-Fig. 2I-N: Most abundant families of ANDEEP I-III stations
APPENDIX- FIGURES
183
st 59-5
68
2424
17
14
1414
SpionidaeCirratulidaeOpheliidaeParaonidaeGlyceridaeHesionidaeNephtyidae
st 74-6
476
399
212
143
132
PolynoidaeSpionidaeSyllidaeHesionidaeAmpharetidae
O P
st 78-9
123
97
53
47
39
SpionidaePholoididaeOpheliidaeGlyceridaeAmpharetidae
st 80-9
45
3729
23
17
GlyceridaeSpionidaeHesionidaeFauveliopsidaePolynoidae
Q R
st 81-8
37
3428
19
18
SpionidaeAmpharet idaeSphaerodoridaeParaonidaeCirratulidae
st 88-8
150
49
29
2523
PolygordiidaeSpionidaeGlyceridaePolynoidaeCirratulidae
S T
App.-Fig. 2O-T: Most abundant families of ANDEEP I-III stations
APPENDIX- FIGURES
184
st 94-14
41
19
19
65 5
PolygordiidaeGlyceridaeSpionidaeCirratulidaeHesionidaePolynoidae
st 102-13
15
1210
8
6
SpionidaePolygordiidaeOpheliidaeGlyceridaeAmpharetidae
U V
st 110-8
76
30
29
12 7
SpionidaeOpheliidaeGlyceridaeFauveliopsidaeHesionidae
st 121-11
98
49
33
1515
OpheliidaeAmpharet idaeOnuphidaeScalibregmat idaeTrichobranchidae
W X
st 133-2
1000
984
577
403192
SyllidaeHesionidaeTerebellidaePolynoidaeSphaerodoridae
st 142-5
16
1310
9
6
6
FauveliopsidaeSpionidaeCirratulidaeOnuphidaeAmpharetidaeTrichobranchidae
Y Z
App.-Fig. 2U-Z: Most abundant families of ANDEEP I-III stations
APPENDIX- FIGURES
185
st 150-6
117
113
37
35
34
PholoididaeSpionidaeOpheliidaeLumbrineridaeAmpharetidae
st 151-7
65
49
28
22
22
SpionidaeParaonidaeHesionidaeSternaspididaeSyllidae
AA AB
st 152-6
76
2 111
CirratulidaeHesionidaeNephtyidaeSphaerodoridaeSpionidae
st 153-7
130
122112
71
44
SpionidaePholoididaeOnuphidaeOpheliidaeSyllidae
AC AD
st 154-9
43
10
6
6
55
OpheliidaeFlabelligeridaeAmpharetidaeSigalionidaeGlyceridaeSpionidae
AE
App.-Fig. 2AA-AE: Most abundant families of ANDEEP I-III stations
APPENDIX- FIGURES
186
App.-Fig. 3: Schematic figures for identifiction of species
APPENDIX- FIGURES
187
App.-Fig. 4: Schematic figures for identifiction of species
APPENDIX- FIGURES
188
A
Melinna cristataMicronephtys sp. 1
Micropodarke cylindripalpata sp.n.Mugga sp. 1BH
Muggoides cf. cinctusMuggoides sp. 1BH
Neosabellides elongatusNereis eugeniae
Octobranchus antarcticusOligobregma blakei
Oligobregma collareOligobregma hartmanae
Oligobregma notialeOligobregma pseudocollareOligobregma quadrispinosa
Oligobregma sp. 1Ophelina ammotrypanella sp.n.
Ophelina breviataOphelina gymnopygeOphelina nematoides
Ophelina robusta sp.n.Ophelina scaphigera
Ophelina setigeraOphelina sp. 3Ophelina sp. 4Ophelina sp. 5
Ophiodromus calligocervix sp.n.Ophiodromus comatus
Parasyllidea delicata sp.n.Phalacrostemma elegans
Phisidia rubrolineataPhisidia sp. 1BH
Phyllocomus croceaPionosyllis cf. maxima
Pionosyllis comosaPionosyllis epipharynx
Pionosyllis sp. 1Pista corrientis
Pista cristataPista spinifera
Polycirrus cf. antarcticusPolycirrus insignis
Polycirrus sp. 1Polycirrus sp. 2
Proceraea cf. mclearanusProclea cf. graffii
Pseudoscalibregma bransfieldiumPseudoscalibregma papilia sp.n.
Pseudoscalibregma ursapiumSamytha sp. 1
Scalibregma inflatumSclerocheilus cf. antarcticus
Sosanopsis kerguelensisSosanopsis sp. 1
Sphaerodoropsis parvaSphaerodoropsis polypapillata
Sphaerodoropsis simplex sp.n.Sphaerodoropsis distincta sp.n.
Sphaerodoropsis maculata sp.n.Sphaerosyllis antarctica
Sphaerosyllis joinvillensisSphaerosyllis lateropapillataStreblosoma variouncinatum
Syllides articulosusTerebellidae sp. 1Terebellidae sp. 2Terebellidae sp. 3Terebellidae sp. 4
Terebellides stroemiiThelepides koehleri
Thelepides venustusThelepodinae sp. 1
Travisia cf. lithophilaTravisia kerguelensis
Trichobranchidae sp. 1Trichobranchidae sp. 2
Typosyllis cf. hyalinaTyposyllis variegata
0 1000 2000 3000 4000 5000
APPENDIX- FIGURES
189
0 1000 2000 3000 4000 5000
aff. HesionidesAglaophamus paramalmgreni
Aglaophamus trissophyllusAmage sculpta
Ammotrypanella arcticaAmmotrypanella cirrosa sp.n.
Ammotrypanella princessa sp.n.Ammotrypanella mcintoshi sp.n.
Ampharete kerguelensisAmpharetidae sp. 1Ampharetidae sp. 2Ampharetidae sp. 3Ampharetidae sp. 3Ampharetidae sp. 4Ampharetidae sp. 5Ampharetidae sp. 6Ampharetidae sp. 7Amphicteis gunneriAmphicteis juvenile
Amphicteis sp. 1Amphicteis sp. 2Amphicteis sp. 3
Amphiduros serratus sp.n.Amphiduros sp. 1
Amythas membraniferaAnobothrella antarctica
Anobothrella sp. 1Anobothrus gracilis
Anobothrus pseudoampharete sp.n.Asclerocheilus ashworthi
Autolytus gibberAxiokebuita millsii
Axiokebuita minutaBathyglycinde sp. 1 BH
Bathyglycinde sp. 2Brania sp. 1 BH
Braniella palpataCeratocephale sp. 1BH
cf. Amphisamythacf. Autolytus simplex stolon
cf. Neosabellidescf. Nicon
cf. Terebellides sp. 1BHcf. Thelepides venustuscf. Thelepus cincinnatusClavodorum antarcticum
Ephesiella antarcticaEphesiella hartmanae sp.n.
Eupistella grubeiEupistella sp. 1
Eusamythella sp. 1Exogone heterosetosa
Exogone minusculaExogone sp. 2 BH
Exogone sp. 5Exogone sp. 6Exogone sp. 7
Glycera kerguelensisGlyphanostomum scotiarum
Goniada maculataGrubianella antarctica
Grubianella sp. 1Gyptis incompta
Hauchiella tribullataHesionidae sp. 1
Hyboscolex equatorialisKefersteinia fauveliKesun abyssorum
Laphania cf. boeckiLeaena antarcticaLeaena arenilega
Leaena cf. collarisLeaena collaris
Leaena pseudobranchiaLeaena sp. 4
Leaena wandelensisLysilla sp. 1 BH
B
App.-Fig. 5: Vertical distribution ranges for species found during ANDEEP I-III, A- species M-Z, B- species A-L
APPENDIX- FIGURES
190
App.-Fig. 6: Global distribution of 84 named species found during the expeditions ANDEEP I-III
(record list see chapter 2.4.2)
APPENDIX- FIGURES
191
Ampharetidae
1
83
4
Opheliidae
2
5
1
Scalibregmatidae
2
81
2
3
Sphaerodoridae
1
11
1
Syllidae
6
4
2
4
Terebellidae
3
11
2
5
LR
A B
C D
E F
SU
SA
SP
SH
C
App.-Fig. 7A-F: Global distribution of southern ocean deep-sea polychaete species collected during
the expeditions ANDEEP I-III (total numbers; C- cosmopolitan, SH- southern
hemisphere, SP- South Pacific, SA- South Atlantic, SU- subantarctic, LR- locally
restricted within the Southern Ocean)
APPENDIX- FIGURES
192
App.-Fig. 8: Global records of the Ampharetidae- Anobothrus gracilis, Station 16-10
App.-Fig. 9: Global records of the Glyceridae
APPENDIX- FIGURES
193
App.-Fig. 10: Global records of the Goniadidae
App.-Fig. 11: Global records of the Hesionidae
APPENDIX- FIGURES
194
App.-Fig. 12: Global records of the Nereididae
App.-Fig. 13: Global records of the Nephtyidae
APPENDIX- FIGURES
195
App.-Fig. 14: Global records of the Opheliidae- Ammotrypanella arctica
App.-Fig. 15: Global records of the Sabellariidae
APPENDIX- FIGURES
196
App.-Fig. 16: Global records of the Scalibregmatidae- Axiokebuita millsi, A. minuta, Scalibregma
inflatum
App.-Fig. 17: Global records of the Sphaerodoridae- Ephesiella antarctica, Clavodorum
antarcticum
APPENDIX- FIGURES
197
App.-Fig. 18: Global records of the Syllidae- Autolytus gibber, Proceraea mclearanus,
Braniella palpata, Exogone heterosetosa, E. minuscula, Typosyllis variegata, T. hyalina
App.-Fig. 19: Global patterns of the Terebellidae- species of the genus Pista, Thelepus
cincinnatus, Hauchiella tribullata, Proclea graffiti, Laphania boeckii
APPENDIX- FIGURES
198
App.-Fig. 20: Global patterns of the Trichobranchidae
App.-Fig. 21: Global patterns of polychaete species found at Southern Atlantic sites (SA, st. 16-10, 21-7)
APPENDIX- FIGURES
199
App.-Fig. 22: Global patterns of polychaete species found at Eastern Southern Atlantic sites (EA, st. 59-5)
App.-Fig. 23: Global patterns of polychaete species found at Eastern Weddell Sea sites (EWS, st. 74-6,
78-9, 80-9, 88-8, 94-14, 102-13)
App.-Fig. 24: Global patterns of polychaete species found at central Weddell Sea sites (WS, st. 110-8,
121-11, 131-3, 132-2, 133-2, 133-3, 134-4, 135-4, 142-5)
APPENDIX- FIGURES
200
App.-Fig. 25: Global patterns of polychaete species found off the South Orkney Islands (SOI, st. 150-6,
151-7)
App.-Fig. 26: Global patterns of polychaete species found in the South Sandwich Trench (SST, st. 141-
10, 143-1)
App.-Fig. 27: Global patterns of polychaete species found off Elephant Island (EI, st. 152-6)
APPENDIX- FIGURES
201
App.-Fig. 28: Global patterns of polychaete species found in the Drake Passage (DP, st. 41-3,
42-2, 46-7)
App.-Fig. 29: Global patterns of polychaete species found off King George Island (KGI, st. 153-7,
154-9)
App.-Fig. 30: Global patterns of polychaete species found between 0 – 1000 m water depth
APPENDIX- FIGURES
202
App.-Fig. 31: Global patterns of polychaete species found between 1000 – 2000 m water depth
App.-Fig. 32: Global patterns of polychaete species found between 2000 – 3000 m water depth
App.-Fig. 33: Global patterns of polychaete species found between 3000 – 4000 m water depth
APPENDIX- FIGURES
203
App.-Fig. 34: Global patterns of polychaete species found between 4000 – 5000 m water depth
APPENDIX- TABLES
204
7.2 Tables
App.-Tab. 1: Family assemblage matrix for EBS-stations of the expeditions ANDEEP I/II
Family 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1 total Acrocirridae 1 5 0 2 3 0 0 0 4 1 16
Ampharetidae 3 13 10 3 0 1 8 0 2 8 48 Amphinomidae 2 3 20 0 0 0 0 0 1 2 28
Apistobranchidae 0 0 1 0 0 0 0 0 0 0 1 Arabellidae 0 0 0 0 0 0 0 0 1 0 1 Capitellidae 0 0 0 0 0 0 0 0 1 0 1
Chaetopteriae 0 1 9 0 0 0 0 0 0 0 10 Cirratulidae 1 42 73 11 0 0 5 2 12 4 150 Cossuridae 0 1 6 0 0 0 0 0 0 0 7 Dorvilleidae 1 1 13 8 0 0 0 0 2 1 26
Euphrosinidae 1 0 1 0 0 0 0 0 0 0 2 Fauveliopsidae 0 0 72 0 0 0 0 4 25 0 101 Flabelligeridae 0 8 27 6 1 0 2 0 4 0 48
Glyceridae 1 2 3 0 0 0 0 17 23 31 77 Goniadidae 0 4 0 0 0 0 0 0 0 0 4 Hesionidae 0 37 16 10 3 0 7 1 39 1 114
Lumbrineridae 1 8 82 0 0 0 1 0 0 76 168 Maldanidae 10 64 7 2 1 0 1 0 16 2 103 Nephtyidae 0 7 34 0 0 0 0 0 0 0 41
Nereidae 0 1 1 0 1 0 0 0 1 0 4 Onuphidae 13 6 65 0 0 0 0 0 3 0 87 Opheliidae 1 26 97 111 0 0 7 1 14 6 263 Orbiniidae 0 12 11 2 1 0 0 2 7 12 47
Paralacydonidae 0 0 1 0 0 0 0 0 0 0 1 Paraonidae 2 12 11 0 0 0 3 0 4 16 48 Pholoididae 9 78 336 0 0 0 0 0 130 0 553
Phyllodocidae 1 3 9 2 0 0 0 0 0 0 15 Polygordiidae 0 0 0 0 0 0 0 0 1 0 1
Polynoidae 0 0 6 5 16 3 3 6 57 20 116 Sabellaridae 0 0 1 0 0 0 0 0 0 0 1 Sabellidae 0 2 57 6 1 0 1 0 13 0 80
Scalibregmatidae 0 31 36 24 6 1 7 0 16 72 193 Sigalionidae 1 26 15 0 0 0 0 0 1 0 43
Sphaerodoridae 2 84 367 2 0 0 0 0 22 22 499 Spionidae 0 207 10 19 1 0 6 35 58 161 497 Syllidae 0 1 3 2 1 1 0 0 17 13 38
Terebellidae 0 1 14 1 0 0 0 0 2 1 19 Trichobranchidae 0 1 4 1 0 0 0 0 2 0 8
indet 0 7 4 17 3 0 21 30 0 0 82 total 3541
APPENDIX- TABLES
205
App.-Tab. 2: Family assemblage matrix for EBS-stations of the expeditions ANDEEP III
family 16-10-S
16-10-E
21-7-S
21-7-E
59-5-S
59-5-E
59-5-Ü
74-6-S
74-6-E
78-9-E
78-9-S
80-9-E
Acrocirridae 2 0 1 1 6 1 0 4 9 0 0 0 Alciopidae 0 0 0 0 0 0 0 0 0 0 0 0
Ampharetidae 15 9 11 6 4 2 1 31 101 21 18 8 Amphinomidae 0 0 0 0 0 0 0 13 54 10 23 0 Aphroditidae 0 0 0 0 0 0 0 1 2 0 0 0
Apistobranchidae 0 0 1 0 0 0 0 0 0 0 0 0 Arabellidae 0 0 0 0 0 0 0 0 0 1 0 0 Capitellidae 1 1 1 0 2 2 0 0 1 1 0 0
Chaetopteridae 0 0 0 0 0 0 0 0 0 1 0 0 Chrysopetaliidae 0 0 0 0 0 0 0 0 2 0 0 0
Cirratulidae 1 2 1 4 9 15 0 16 16 25 13 5 Cossuridae 0 0 0 0 0 0 0 0 0 0 0 0 Dorvilleidae 0 0 0 0 0 0 0 0 3 0 0 0
Euphrosinidae 0 0 0 0 0 0 0 1 0 0 0 3 Fauveliopsidae 11 1 17 3 0 3 0 0 2 5 0 18 Flabelligeridae 9 2 6 3 1 5 2 2 5 11 5 4
Glyceridae 4 0 2 0 10 4 0 6 37 38 9 28 Goniadidae 2 0 0 0 0 0 0 0 0 0 0 0 Hesionidae 10 2 8 3 4 10 0 19 124 11 3 20
Lacydoniidae 0 0 0 0 0 0 0 0 1 0 0 1 Lumbrineridae 0 0 2 3 0 1 0 2 7 2 3 0
Lysaretidae 0 0 0 0 0 0 0 0 0 0 0 0 Maldanidae 1 0 0 0 0 2 0 1 0 4 1 0 Nephtyidae 0 0 0 0 10 4 0 12 61 11 8 2 Nereididae 0 0 0 0 0 0 0 0 0 1 0 0 Onuphidae 2 0 0 0 2 2 0 0 2 0 1 0 Opheliidae 25 4 2 0 16 7 1 11 29 31 22 8 Orbiniidae 1 1 3 1 0 0 0 2 10 7 3 0 Oweniidae 0 0 0 0 1 0 0 0 9 3 3 0 Paraonidae 6 3 4 5 10 5 2 4 30 11 8 3 Pholoididae 0 0 0 0 0 0 0 1 2 80 17 9
Phyllodocidae 2 0 0 0 0 0 0 2 11 2 0 1 Pilargiidae 4 1 0 1 0 0 0 0 0 0 0 0
Polygordiidae 0 0 0 0 1 0 0 1 0 3 0 0 Polynoidae 10 2 4 0 5 4 0 70 406 8 12 12
Sabellariidae 0 0 0 0 0 0 0 0 0 0 0 1 Sabellidae 5 0 4 4 2 2 0 10 32 0 1 2
Scalibregmatidae 0 0 0 1 8 2 0 3 8 7 0 14 Sigalionidae 2 0 4 1 5 0 1 0 1 3 5 1
Sphaerodoridae 2 0 0 1 1 4 1 14 101 3 0 8 Spionidae 20 16 20 13 17 50 1 68 331 81 42 32
Sternaspididae 0 0 0 0 0 0 0 3 5 0 0 0 Syllidae 0 0 1 0 0 1 0 44 168 0 1 1
Terebellidae 2 1 1 2 1 0 0 19 37 1 2 2 Trichobranchidae 1 0 0 0 0 1 0 8 17 10 2 6 Trochochaetidae 1 0 0 0 1 0 0 0 0 0 0 0
APPENDIX- TABLES
206
family 80-9-S
80-9-Ü
81-8-E
81-8-S
81-8-Ü
88-8-E
88-8-S
88-8-Ü
94-14-S
94-14-Ü
94-14-E
102-13-E
Acrocirridae 0 0 0 2 0 0 0 0 0 0 0 0 Alciopidae 0 0 0 0 0 0 0 0 0 0 0 0
Ampharetidae 2 3 8 23 3 1 0 0 1 0 2 1 Amphinomidae 1 1 0 0 0 0 0 0 0 0 0 0 Aphroditidae 0 0 0 0 0 0 0 0 0 0 0 0
Apistobranchidae 0 0 0 0 0 0 0 0 0 0 0 0 Arabellidae 0 0 0 0 0 0 0 0 0 0 0 0 Capitellidae 0 0 0 1 0 2 0 0 0 0 0 0
Chaetopteridae 0 0 0 0 0 0 0 0 0 0 0 0 Chrysopetaliidae 0 0 0 0 0 0 0 0 0 0 0 0
Cirratulidae 0 2 11 5 2 18 2 3 1 0 5 0 Cossuridae 0 0 0 0 0 0 0 0 0 0 0 0 Dorvilleidae 0 0 0 0 0 0 0 0 0 0 0 0
Euphrosinidae 0 2 6 5 1 0 0 0 0 0 0 0 Fauveliopsidae 3 2 9 1 1 0 0 0 0 0 0 0 Flabelligeridae 3 0 2 0 0 9 0 0 0 0 1 0
Glyceridae 5 12 0 1 1 18 7 4 9 1 9 6 Goniadidae 0 0 0 0 0 0 0 0 0 0 0 0 Hesionidae 9 0 7 0 0 12 2 1 3 0 2 1
Lacydoniidae 0 0 0 0 0 0 0 0 0 0 0 0 Lumbrineridae 0 0 2 4 0 0 0 0 0 0 0 0
Lysaretidae 0 0 0 0 0 0 0 0 0 0 0 0 Maldanidae 0 0 11 4 0 0 0 0 0 0 0 0 Nephtyidae 0 0 0 0 0 0 0 0 0 0 0 0 Nereididae 0 0 0 0 0 0 1 0 0 0 0 0 Onuphidae 0 0 0 7 1 0 0 0 0 0 0 0 Opheliidae 1 2 2 7 0 0 0 0 1 0 0 9 Orbiniidae 0 0 4 0 0 0 0 0 0 0 0 0 Oweniidae 0 0 0 1 0 0 0 0 0 0 0 0 Paraonidae 1 3 7 12 0 0 0 0 0 0 0 0 Pholoididae 3 2 0 0 0 1 0 0 0 0 0 0
Phyllodocidae 0 0 0 0 0 0 0 0 0 0 0 0 Pilargiidae 0 0 0 0 0 0 0 0 0 0 0 0
Polygordiidae 0 0 0 0 7 129 21 0 21 0 20 12 Polynoidae 4 1 4 1 0 15 8 2 2 0 3 1
Sabellariidae 0 0 0 0 0 0 0 0 0 0 0 0 Sabellidae 0 0 0 0 0 0 0 0 0 0 0 0
Scalibregmatidae 0 1 0 1 0 2 0 0 2 0 0 0 Sigalionidae 0 0 0 0 0 1 1 0 2 0 0 0
Sphaerodoridae 1 3 14 12 2 0 0 0 0 0 0 0 Spionidae 2 3 27 9 1 33 14 2 12 0 7 14
Sternaspididae 0 0 0 0 0 0 0 0 0 0 0 0 Syllidae 0 1 1 0 0 0 0 0 0 0 0 0
Terebellidae 1 2 2 1 0 0 0 0 0 0 0 1 Trichobranchidae 2 1 1 1 0 0 0 0 2 0 0 1 Trochochaetidae 0 0 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
207
family 102-13-Ü
102-13-S
110-8-S
110-8-E
110-8-Ü
121-11-E
121-11-S
133-2-Ü
133-2-E
133-2-S
142-5-E
142-5-Ü
Acrocirridae 0 0 0 1 0 0 0 0 1 2 0 0 Alciopidae 0 0 0 0 0 0 0 0 0 1 0 0
Ampharetidae 5 0 0 2 0 30 19 1 14 11 1 1 Amphinomidae 0 0 0 0 0 1 0 5 64 10 0 0 Aphroditidae 0 0 0 0 0 0 0 0 0 0 0 0
Apistobranchidae 0 0 0 0 0 0 0 0 0 0 0 0 Arabellidae 0 0 0 0 0 0 0 0 0 0 0 0 Capitellidae 0 0 0 0 0 1 0 0 1 0 0 0
Chaetopteridae 0 0 0 0 0 0 0 0 0 0 0 0 Chrysopetaliidae 0 0 0 0 0 0 0 0 0 0 0 0
Cirratulidae 0 0 1 6 1 9 4 7 81 9 0 0 Cossuridae 0 0 0 0 0 0 0 0 0 0 0 0 Dorvilleidae 1 0 0 0 0 0 1 3 0 0 0 0
Euphrosinidae 0 0 0 0 0 0 0 1 22 3 0 0 Fauveliopsidae 0 0 4 5 3 0 0 0 0 0 3 3 Flabelligeridae 0 0 1 2 0 2 0 0 4 2 0 0
Glyceridae 1 1 8 16 5 0 0 3 13 3 0 0 Goniadidae 0 0 0 0 0 0 0 0 0 0 0 0 Hesionidae 0 0 0 7 0 2 4 7 837 140 1 0
Lacydoniidae 0 0 0 0 0 0 0 0 0 0 0 0 Lumbrineridae 0 0 0 0 0 0 0 0 0 0 0 0
Lysaretidae 0 0 0 0 0 0 0 3 13 4 0 0 Maldanidae 0 0 0 0 0 1 0 5 123 37 0 1 Nephtyidae 0 0 1 1 1 0 0 0 0 0 0 0 Nereididae 0 0 0 1 0 0 0 0 0 0 0 0 Onuphidae 1 1 0 0 0 24 9 1 0 0 5 0 Opheliidae 0 1 12 11 7 63 35 0 47 2 0 0 Orbiniidae 0 0 1 5 0 0 0 0 1 0 2 0 Oweniidae 0 0 0 0 0 1 1 0 1 0 1 1 Paraonidae 0 0 0 0 0 1 0 13 106 27 0 0 Pholoididae 0 0 0 0 0 0 0 0 18 0 0 0
Phyllodocidae 0 0 0 0 0 0 0 6 31 23 0 0 Pilargiidae 0 0 0 0 0 0 0 0 0 0 0 0
Polygordiidae 0 0 0 2 0 3 0 0 0 0 0 0 Polynoidae 0 0 1 3 0 1 4 0 337 66 1 1
Sabellariidae 0 0 0 0 0 0 0 0 0 0 0 0 Sabellidae 0 0 0 0 0 0 0 3 11 2 0 0
Scalibregmatidae 0 0 2 0 0 11 4 1 30 8 1 0 Sigalionidae 0 0 0 2 0 2 1 1 0 0 0 0
Sphaerodoridae 0 0 0 0 0 5 0 5 161 26 0 0 Spionidae 1 0 11 58 7 6 2 19 131 22 2 1
Sternaspididae 0 0 0 0 0 0 0 0 0 0 0 0 Syllidae 0 0 0 0 0 1 1 5 887 108 0 0
Terebellidae 0 0 0 0 0 0 0 39 427 111 0 1 Trichobranchidae 0 0 0 3 0 12 3 0 93 27 3 0 Trochochaetidae 0 0 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
208
family 142-5S
150-6-E
150-6-Ü
150-6-S
151-7-E
151-7-S
152-6-S
152-6-E
153-7-S
153-7-E
154-9-E
154-9-S total
Acrocirridae 1 0 0 0 0 0 0 0 0 0 2 0 33 Alciopidae 0 0 0 0 0 0 0 0 0 0 0 0 1
Ampharetidae 4 29 0 5 2 2 0 0 13 6 4 2 422 Amphinomidae 2 23 0 3 7 0 0 0 9 0 0 0 226 Aphroditidae 0 0 0 0 0 0 0 0 0 0 0 0 3
Apistobranchidae 0 0 0 0 0 0 0 0 0 0 0 0 1 Arabellidae 0 1 0 0 0 0 0 0 2 1 0 4 9 Capitellidae 0 0 0 0 11 2 0 0 3 0 1 0 31
Chaetopteridae 0 0 0 0 0 0 0 0 0 0 0 0 1 Chrysopetaliidae 0 0 0 0 0 0 0 0 0 1 0 0 3
Cirratulidae 10 13 5 0 11 8 1 75 8 17 2 2 426 Cossuridae 0 0 1 0 0 0 0 0 4 6 0 0 11 Dorvilleidae 0 0 0 0 9 2 0 0 0 14 0 0 33
Euphrosinidae 0 1 0 0 0 0 0 0 2 4 0 0 51 Fauveliopsidae 10 3 0 1 4 1 0 0 6 8 0 0 127 Flabelligeridae 0 0 0 1 1 1 0 0 4 2 8 2 100
Glyceridae 0 6 0 0 0 0 0 0 2 2 3 2 276 Goniadidae 0 0 0 0 0 0 0 0 0 0 0 0 2 Hesionidae 2 10 0 3 19 9 2 0 2 7 3 1 1307
Lacydoniidae 0 0 0 0 0 0 0 0 0 0 0 0 2 Lumbrineridae 0 32 0 3 4 1 0 0 14 23 0 1 104
Lysaretidae 0 0 0 0 0 0 0 0 0 0 0 0 20 Maldanidae 2 4 0 0 0 0 0 0 2 10 1 0 210 Nephtyidae 0 2 1 0 0 2 0 1 8 7 2 2 136 Nereididae 0 0 0 0 0 0 0 0 0 1 1 1 6 Onuphidae 4 1 0 0 1 1 0 0 92 20 1 2 180 Opheliidae 4 35 1 1 2 0 0 0 54 17 13 30 513 Orbiniidae 0 0 0 0 1 0 0 0 0 0 2 1 45 Oweniidae 1 0 0 0 0 0 0 0 0 0 0 0 23 Paraonidae 0 6 0 0 28 21 0 0 4 11 2 0 333 Pholoididae 0 101 0 16 0 1 0 0 87 35 4 0 377
Phyllodocidae 0 4 0 0 3 2 0 0 4 18 2 0 111 Pilargiidae 0 0 0 0 0 0 0 0 0 0 0 0 6
Polygordiidae 0 0 0 0 0 0 0 0 0 5 0 0 225 Polynoidae 3 7 0 2 1 0 0 0 11 4 0 0 1016
Sabellariidae 0 0 0 0 0 0 0 0 0 0 0 0 1 Sabellidae 1 2 0 0 0 0 0 0 2 1 1 1 86
Scalibregmatidae 1 11 1 1 0 5 0 0 21 3 1 0 150 Sigalionidae 0 0 0 0 0 0 0 0 2 0 1 5 41
Sphaerodoridae 2 5 0 0 6 5 0 1 4 3 2 0 392 Spionidae 10 95 0 18 40 25 1 0 43 87 5 0 1399
Sternaspididae 0 0 0 0 20 2 0 0 0 0 0 0 30 Syllidae 0 5 1 0 10 12 0 0 10 34 2 0 1294
Terebellidae 1 1 0 1 0 0 0 0 4 4 3 1 668 Trichobranchidae 3 4 0 0 0 0 0 0 1 0 0 0 202 Trochochaetidae 0 0 0 0 0 0 0 0 0 0 0 0 2
total 10635
APPENDIX- TABLES
209
App.-Tab. 3: Family assemblage matrix for EBS-stations of the expeditions ANDEEPI-III
family 41-3 42-2 46-7 131-1 132-2 133-3 134-4 135-4 141-10 143-1 Acrocirridae 1 5 0 2 3 0 0 0 4 1 Alciopidae 0 0 0 0 0 0 0 0 0 0
Ampharetidae 3 13 10 3 0 1 8 0 2 8 Amphinomidae 2 3 20 0 0 0 0 0 1 2 Aphroditidae 0 0 0 0 0 0 0 0 0 0
Apistobranchidae 0 0 1 0 0 0 0 0 0 0 Arabellidae 0 0 0 0 0 0 0 0 1 0 Capitellidae 0 0 0 0 0 0 0 0 1 0
Chaetopteridae 0 1 9 0 0 0 0 0 0 0 Chrysopetaliidae 0 0 0 0 0 0 0 0 0 0
Cirratulidae 1 42 73 11 0 0 5 2 12 4 Cossuridae 0 1 6 0 0 0 0 0 0 0 Dorvilleidae 1 1 13 8 0 0 0 0 2 1
Euphrosinidae 1 0 1 0 0 0 0 0 0 0 Fauveliopsidae 0 0 72 0 0 0 0 4 25 0 Flabelligeridae 0 8 27 6 1 0 2 0 4 0
Glyceridae 1 2 3 0 0 0 0 17 23 31 Goniadidae 0 4 0 0 0 0 0 0 0 0 Hesionidae 0 37 16 10 3 0 7 1 39 1
Lacydoniidae 0 0 0 0 0 0 0 0 0 0 Lumbrineridae 1 8 82 0 0 0 1 0 0 76
Lysaretidae 0 0 0 0 0 0 0 0 0 0 Maldanidae 10 64 7 2 1 0 1 0 16 2 Nephtyidae 0 7 34 0 0 0 0 0 0 0 Nereididae 0 1 1 0 1 0 0 0 1 0 Onuphidae 13 6 65 0 0 0 0 0 3 0 Opheliidae 1 26 97 111 0 0 7 1 14 6 Orbiniidae 0 12 11 2 1 0 0 2 7 12 Oweniidae 0 0 0 0 0 0 0 0 0 0
Paralacydonidae 0 0 1 0 0 0 0 0 0 0 Paraonidae 2 12 11 0 0 0 3 0 4 16 Pholoididae 9 78 336 0 0 0 0 0 130 0
Phyllodocidae 1 3 9 2 0 0 0 0 0 0 Pilargiidae 0 0 0 0 0 0 0 0 0 0
Polygordiidae 0 0 0 0 0 0 0 0 1 0 Polynoidae 0 0 6 5 16 3 3 6 57 20
Sabellariidae 0 0 1 0 0 0 0 0 0 0 Sabellidae 0 2 57 6 1 0 1 0 13 0
Scalibregmatidae 0 31 36 24 6 1 7 0 16 72 Sigalionidae 1 26 15 0 0 0 0 0 1 0
Sphaerodoridae 2 84 367 2 0 0 0 0 22 22 Spionidae 0 207 10 19 1 0 6 35 58 161
Sternaspididae 0 0 0 0 0 0 0 0 0 0 Syllidae 0 1 3 2 1 1 0 0 17 13
Terebellidae 0 1 14 1 0 0 0 0 2 1 Trichobranchidae 0 1 4 1 0 0 0 0 2 0 Trochochaetidae 0 0 0 0 0 0 0 0 0 0
indet 0 7 4 17 3 0 21 30 0 0
APPENDIX- TABLES
210
family 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13 Acrocirridae 2 2 7 13 0 0 2 0 0 0 Alciopidae 0 0 0 0 0 0 0 0 0 0
Ampharetidae 24 17 7 132 39 13 34 1 3 6 Amphinomidae 0 0 0 67 33 2 0 0 0 0 Aphroditidae 0 0 0 3 0 0 0 0 0 0
Apistobranchidae 0 1 0 0 0 0 0 0 0 0 Arabellidae 0 0 0 0 1 0 0 0 0 0 Capitellidae 2 1 4 1 1 0 1 2 0 0
Chaetopteridae 0 0 0 0 1 0 0 0 0 0 Chrysopetaliidae 0 0 0 2 0 0 0 0 0 0
Cirratulidae 3 5 24 32 38 7 18 23 6 0 Cossuridae 0 0 0 0 0 0 0 0 0 0 Dorvilleidae 0 0 0 3 0 0 0 0 0 1
Euphrosinidae 0 0 0 1 0 5 12 0 0 0 Fauveliopsidae 12 20 3 2 5 23 11 0 0 0 Flabelligeridae 11 9 8 7 16 7 2 9 1 0
Glyceridae 4 2 14 43 47 45 2 29 19 8 Goniadidae 2 0 0 0 0 0 0 0 0 0 Hesionidae 12 11 14 143 14 29 7 15 5 1
Lacydoniidae 0 0 0 1 0 1 0 0 0 0 Lumbrineridae 0 5 1 9 5 0 6 0 0 0
Lysaretidae 0 0 0 0 0 0 0 0 0 0 Maldanidae 1 0 2 1 5 0 15 0 0 0 Nephtyidae 0 0 14 73 19 2 0 0 0 0 Nereididae 0 0 0 0 1 0 0 1 0 0 Onuphidae 2 0 4 2 1 0 8 0 0 2 Opheliidae 29 2 24 40 53 11 9 0 1 10 Orbiniidae 2 4 0 12 10 0 4 0 0 0 Oweniidae 0 0 1 9 6 0 1 0 0 0
Paralacydonidae 0 0 0 0 0 0 0 0 0 0 Paraonidae 9 9 17 34 19 7 19 0 0 0 Pholoididae 0 0 0 3 97 14 0 1 0 0
Phyllodocidae 2 0 0 13 2 1 0 0 0 0 Pilargiidae 5 1 0 0 0 0 0 0 0 0
Polygordiidae 0 0 1 1 3 0 7 150 41 12 Polynoidae 12 4 9 476 20 17 5 25 5 1
Sabellariidae 0 0 0 0 0 1 0 0 0 0 Sabellidae 5 8 4 42 1 2 0 0 0 0
Scalibregmatidae 0 1 10 11 7 15 1 2 2 0 Sigalionidae 2 5 6 1 8 1 0 2 2 0
Sphaerodoridae 2 1 6 115 3 12 28 0 0 0 Spionidae 36 33 68 399 123 37 37 49 19 15
Sternaspididae 0 0 0 8 0 0 0 0 0 0 Syllidae 0 1 1 212 1 2 1 0 0 0
Terebellidae 3 3 1 56 3 5 3 0 0 1 Trichobranchidae 1 0 1 25 12 9 2 0 2 1 Trochochaetidae 1 0 1 0 0 0 0 0 0 0
indet 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
211
family 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total Acrocirridae 1 0 3 1 0 0 0 0 2 49 Alciopidae 0 0 1 0 0 0 0 0 0 1
Ampharetidae 2 49 26 6 34 4 0 19 6 470 Amphinomidae 0 1 79 2 26 7 0 9 0 254 Aphroditidae 0 0 0 0 0 0 0 0 0 3
Apistobranchidae 0 0 0 0 0 0 0 0 0 2 Arabellidae 0 0 0 0 1 0 0 3 4 10 Capitellidae 0 1 1 0 0 13 0 3 1 32
Chaetopteridae 0 0 0 0 0 0 0 0 0 11 Chrysopetaliidae 0 0 0 0 0 0 0 1 0 3
Cirratulidae 8 13 97 10 18 19 76 25 4 576 Cossuridae 0 0 0 0 1 0 0 10 0 18 Dorvilleidae 0 1 3 0 0 11 0 14 0 59
Euphrosinidae 0 0 26 0 1 0 0 6 0 53 Fauveliopsidae 12 0 0 16 4 5 0 14 0 228 Flabelligeridae 3 2 6 0 1 2 0 6 10 148
Glyceridae 29 0 19 0 6 0 0 4 5 353 Goniadidae 0 0 0 0 0 0 0 0 0 6 Hesionidae 7 6 984 3 13 28 2 9 4 1421
Lacydoniidae 0 0 0 0 0 0 0 0 0 2 Lumbrineridae 0 0 0 0 35 5 0 37 1 272
Lysaretidae 0 0 20 0 0 0 0 0 0 20 Maldanidae 0 1 165 3 4 0 0 12 1 313 Nephtyidae 3 0 0 0 3 2 1 15 4 177 Nereididae 1 0 0 0 0 0 0 1 2 10 Onuphidae 0 33 1 9 1 2 0 112 3 267 Opheliidae 30 98 49 4 37 2 0 71 43 776 Orbiniidae 6 0 1 2 0 1 0 0 3 92 Oweniidae 0 2 1 3 0 0 0 0 0 23
Paralacydonidae 0 0 0 0 0 0 0 0 0 1 Paraonidae 0 1 146 0 6 49 0 15 2 381 Pholoididae 0 0 18 0 117 1 0 122 4 930
Phyllodocidae 0 0 60 0 4 5 0 22 2 126 Pilargiidae 0 0 0 0 0 0 0 0 0 6
Polygordiidae 2 3 0 0 0 0 0 5 0 226 Polynoidae 4 5 403 5 9 1 0 15 0 1132
Sabellariidae 0 0 0 0 0 0 0 0 0 2 Sabellidae 0 0 16 1 2 0 0 3 2 166
Scalibregmatidae 2 15 39 2 13 5 0 24 1 343 Sigalionidae 2 3 1 0 0 0 0 2 6 84
Sphaerodoridae 0 5 192 2 5 11 1 7 2 891 Spionidae 76 8 172 13 113 65 1 130 5 1896
Sternaspididae 0 0 0 0 0 22 0 0 0 30 Syllidae 0 2 1000 0 6 22 0 44 2 1332
Terebellidae 0 0 577 2 2 0 0 8 4 687 Trichobranchidae 3 15 120 6 4 0 0 1 0 210 Trochochaetidae 0 0 0 0 0 0 0 0 0 2
indet 0 0 0 0 0 0 0 0 0 82 total 14176
APPENDIX- TABLES
212
App.-Tab. 4: Species assemblage matrix for EBS-stations of the expeditions ANDEEPI/II
species 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1 total
Aglaophamus paramalmgreni 0 5 6 0 0 0 0 0 0 0 11 Aglaophamus trissophyllus 0 2 27 0 0 0 0 0 0 0 29
Amaga sculpta 0 1 0 0 0 0 0 0 0 0 1 Ammotrypanella arctica 0 2 0 7 0 0 1 0 0 0 10
Ammotrypanella cirrosa sp.n. 0 4 0 64 0 0 2 0 1 0 71 Ammotrypanella princessa sp.n. 0 1 0 0 0 0 1 1 0 0 3
Ampharete kerguelensis 2 0 7 0 0 0 0 0 0 0 9 Ampharetidae sp. 1 0 0 0 0 0 0 0 0 1 0 1 Ampharetidae sp. 2 0 0 0 0 0 0 2 0 0 0 2 Ampharetidae sp. 3 0 0 0 0 0 0 2 0 0 0 2 Amphicteis gunneri 0 0 1 0 0 0 0 0 0 0 1
Amphicteis sp. 1 1 0 0 0 0 0 0 0 0 0 1 Amphiduros serratus sp.n. 0 0 0 0 2 0 1 0 0 0 3
Amphiduros sp. 1 0 0 12 1 0 0 0 0 1 0 14 Anobothrella antarctica 0 0 0 0 0 0 0 0 0 1 1
Anobothrus pseudoampharete sp.n. 0 1 0 0 0 1 0 0 0 7 9 Ascleirocheilus ashworthi 0 0 1 0 0 0 0 0 0 0 1
Axiokebuita minuta 0 0 1 6 0 1 3 0 11 0 22 Axiokebuita millsi 0 0 0 0 0 0 0 0 0 62 62
Bathyglycinde sp. 1 BH 0 3 0 0 0 0 0 0 0 0 3 Bathyglycinde sp. 2 0 1 0 0 0 0 0 0 0 0 1
Brania sp. 1 BH 0 0 0 2 1 0 0 0 0 0 3 Braniella palpata 0 0 0 0 0 0 0 0 2 0 2
cf. Autolytus simplex stolon 0 0 1 0 0 0 0 0 0 0 1 cf. Muggoides sp. 1 0 1 0 0 0 0 0 0 0 0 1
Clavodorum antarcticum 0 0 1 0 0 0 0 0 0 0 1 Ephesiella antarctica 0 1 5 0 0 0 0 0 0 8 14
Ephesiella hartmanae sp.n. 0 0 0 0 0 0 0 0 0 3 3 Eusamythella sp. 1 0 4 0 0 0 0 0 0 0 0 4
Exogone heterosetosa 0 1 0 0 0 0 0 0 0 0 1 Exogone sp. 2 BH 0 0 1 0 0 0 0 0 0 0 1
Glycera kerguelensis 1 2 3 0 0 0 0 17 23 31 77 Glyphanostomum scotiarum 0 3 0 0 0 0 0 1 0 0 4
Grubianella antarctica 0 2 0 0 0 0 0 0 0 0 2 Hauchiella tribullata 0 1 0 0 0 0 0 0 0 0 1
Hesionidae sp. 1 0 0 0 3 0 0 0 0 0 0 3 Hyboscolex equatorialis 0 0 0 0 0 0 0 0 1 0 1
Kefersteinia fauveli 0 0 0 0 0 0 0 0 35 0 35 Kesun abyssorum 0 0 86 1 0 0 3 0 8 0 98
Laphania cf. boeckeri 0 0 2 0 0 0 0 0 0 0 2 Lysilla sp. 1BH 0 0 1 0 0 0 0 0 0 0 1
Micronephtys sp. 1 0 0 1 0 0 0 0 0 0 0 1 Micropodarke cylindripalpata sp.n. 0 9 4 2 0 0 0 0 2 0 17
Mugga sp. 1BH 0 0 0 0 0 0 0 0 1 0 1
APPENDIX- TABLES
213
species 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1 total
Neosabellides elongatus 0 0 2 0 0 0 1 0 0 0 3 Nereis eugeniae 0 1 1 0 1 0 0 0 1 0 4
Octobranchus antarcticus 0 0 1 0 0 0 0 0 1 0 2 Oligobregma blakei 0 0 1 0 0 0 0 0 0 0 1 Oligobregma collare 0 4 3 8 0 0 2 0 0 0 17 Oligobregma notiale 0 7 0 0 0 0 0 0 0 0 7
Oligobregma pseudocollare 0 0 20 1 0 0 0 0 0 8 29 Oligobregma quadrispinosa 0 5 0 8 0 0 2 0 1 0 16
Ophelina ammotrypanella sp.n. 0 0 0 2 0 0 0 0 0 0 2 Ophelina breviata 0 1 2 16 0 0 0 0 0 0 19
Ophelina gymnopyge 0 3 0 15 0 0 0 0 0 0 18 Ophelina nematoides 1 14 7 2 0 0 0 0 5 0 29
Ophelina robusta sp.n. 0 0 0 1 0 0 0 0 0 0 1 Ophelina setigera 0 0 1 1 0 0 0 0 0 0 2
Ophelina sp. 3 0 0 0 2 0 0 0 0 0 0 2 Ophelina sp. 4 0 1 0 0 0 0 0 0 0 0 1
Ophiodromus calligocervix sp.n. 0 28 0 3 1 0 6 1 1 1 41 Parasyllidea delicata sp.n. 0 0 0 1 0 0 0 0 0 0 1 Phalacromstemma elegans 0 0 1 0 0 0 0 0 0 0 1
Pionosyllis epipharynx 0 0 1 0 0 1 0 0 1 0 3 Pionosyllis sp. 1 0 0 0 0 0 0 0 0 1 0 1
Polycirrus insignis 0 0 1 0 0 0 0 0 0 1 2 Proclea graffii 0 0 1 0 0 0 0 0 0 0 1
Pseudoscalibregma bransfieldium 0 8 5 0 6 0 0 0 0 2 21 Pseudoscalibregma papilia sp.n. 0 3 2 0 0 0 0 0 3 0 8
Pseudoscalibregma usarpium 0 1 1 1 0 0 0 0 0 0 3 Scalibregma inflatum 0 3 2 0 0 0 0 0 0 0 5
Sosanopsis kerguelensis 0 0 0 3 0 0 2 0 0 0 5 Sphaerodoropsis distincta sp.n. 0 0 6 0 0 0 0 0 0 0 6
Sphaerodoropsis parva 2 82 333 0 0 0 0 0 21 11 449 Sphaerodoropsis polypapillata 0 1 0 2 0 0 0 0 0 0 3 Sphaerodoropsis simplex sp.n. 0 0 22 0 0 0 0 0 1 0 23
Sphaerosyllis joinvillensis 0 0 0 0 0 0 0 0 1 0 1 Sphaerosyllis lateropapillata uteae 0 0 0 0 0 0 0 0 12 1 13
Streblosoma variouncinatum 0 0 0 0 0 0 0 0 2 0 2 Terebellidae sp. 1 0 0 0 1 0 0 0 0 0 0 1 Terebellidae sp. 2 0 0 1 0 0 0 0 0 0 0 1 Terebellidae sp. 4 0 0 1 0 0 0 0 0 0 0 1
Terebellides stroemii 0 1 2 1 0 0 0 0 0 0 4 Thelepodinae sp. 1 0 0 7 0 0 0 0 0 0 0 7
Travisia kerguelensis 0 0 1 0 0 0 0 0 0 6 7 Trichobranchidae sp. 1 0 0 1 0 0 0 0 0 0 0 1 Trichobranchidae sp. 2 0 0 0 0 0 0 0 0 1 0 1
Typosyllis hyalina 0 0 0 0 0 0 0 0 0 8 8 Typosyllis variegata 0 0 0 0 0 0 0 0 0 4 4
total 1308
APPENDIX- TABLES
214
App.-Tab. 5: Species assemblage matrix for EBS-stations of the expeditions ANDEEPIII
Species 16-10-S 16-10-E 21-7-S 21-7-E 59-5-S 59-5-E 59-5-Ü 74-6-S 74-6-E 78-9-E
aff. Hesionides 2 2 2 1 0 1 0 0 0 0 Aglaophamus paramalmgreni 0 0 0 0 10 4 0 0 0 1
Aglaophamus trissophyllus 0 0 0 0 0 0 0 12 61 10 Amage sculpta 0 0 5 3 0 0 0 0 1 0
Ammotrypanella arctica 6 0 0 0 4 0 0 0 0 1 Ammotrypanella cirrosa sp.n. 6 0 0 0 0 2 0 0 0 2
Ammotrypanella mcintoshi sp.n. 1 0 0 0 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 1 2 1 0 1 0 0 0 2 0
Ampharete kerguelensis 9 4 0 0 1 0 0 0 0 1 Ampharetidae sp. 2 0 0 0 1 0 0 0 0 0 2 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 0 0 Amphicteis gunneri 0 0 0 0 0 0 0 0 0 1 Amphicteis juvenile 0 0 0 0 0 0 0 0 0 0
Amphicteis sp. 1 0 0 0 0 0 0 0 0 0 14 Amphicteis sp. 2 0 0 0 0 0 0 0 0 0 0 Amphicteis sp. 3 0 0 0 0 0 0 0 3 2 0
Amphiduros serratus sp.n. 0 0 0 0 0 0 0 0 0 0 Amythas membranifera 0 0 0 0 0 0 0 0 0 0 Anobothrella antarctica 0 0 0 0 0 0 0 2 1 0
Anobothrella sp. 1 0 0 0 0 0 0 0 0 0 0 Anobothrus gracilis 0 0 0 0 0 0 0 0 0 0
Anobothrus pseudoampharete sp.n. 0 0 0 1 0 0 0 20 84 0 Autolytus gibber 0 0 0 0 0 0 0 0 1 0
Axiokebuita minuta 0 0 0 0 0 0 0 2 6 0 Bathyglycinde sp. 1 BH 1 0 0 0 0 0 0 0 0 0
Braniella palpata 0 0 1 0 0 1 0 0 3 0 Ceratocephale sp. 1BH 0 0 0 0 0 0 0 0 0 1
cf. Amphisamytha 0 0 0 0 0 0 0 1 0 0 cf. Neosabellides 0 0 0 0 0 0 0 1 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 0 0 cf. Thelepides venustus 0 0 0 0 0 0 0 1 0 0 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 1 0 Clavodorum antarcticum 0 0 0 0 0 0 0 0 0 0
Ephesiella antarctica 0 0 0 0 0 0 0 0 0 0 Ephesiella hartmanae sp.n. 1 0 0 0 0 0 0 0 2 0
Eupistella grubei 0 0 0 0 0 0 0 5 4 0 Eupistella sp. 1 2 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 0 0 0 0 0 1 0 Exogone heterosetosa 0 0 0 0 0 0 0 0 1 0
Exogone minuscula 0 0 0 0 0 0 0 2 6 0 Exogone sp. 5 0 0 0 0 0 0 0 1 4 0 Exogone sp. 6 0 0 0 0 0 0 0 0 0 0 Exogone sp. 7 0 0 0 0 0 0 0 0 0 0
Glycera kerguelensis 4 0 2 0 10 4 0 6 37 38 Glyphanostomum scotiarum 0 0 0 0 0 0 0 2 1 1
Goniada maculata 1 0 0 0 0 0 0 0 0 0 Grubianella antarctica 2 0 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 0 0 0 0 0 1 0 Gyptis incompta 0 0 0 0 0 0 0 5 43 0
Hauchiella tribullata 0 0 0 0 0 0 0 2 0 0 Kefersteinia fauveli 3 0 0 0 4 2 0 11 59 0 Kesun abyssorum 0 0 1 0 10 4 0 0 0 11 Leaena antarctica 0 0 0 0 0 0 0 0 0 0 Leaena arenilega 0 0 0 0 0 0 0 0 5 0 Leaena cf. collaris 0 1 0 0 0 0 0 0 3 0
Leaena collaris 0 0 0 0 0 0 0 0 2 0 Leaena pseudobranchia 0 0 0 0 0 0 0 1 2 0
Leaena sp. 4 0 0 0 0 0 0 0 0 1 0 Leaena wandelensis 0 0 0 1 0 0 0 0 0 0
APPENDIX- TABLES
215
species 78-9-S 80-9-E 80-9-S 80-9-Ü 81-8-E 81-8-S 81-8-Ü 88-8-E 88-8-S 88-8-Ü
aff. Hesionides 0 0 0 0 0 0 0 0 0 1 Aglaophamus paramalmgreni 0 0 0 0 0 0 0 0 0 0
Aglaophamus trissophyllus 8 2 0 0 0 0 0 0 0 0 Amage sculpta 0 2 0 1 0 0 0 0 0 0
Ammotrypanella arctica 0 0 0 0 0 0 0 0 0 0 Ammotrypanella cirrosa sp.n. 0 0 0 0 0 0 0 0 0 0
Ammotrypanella mcintoshi sp.n. 1 0 0 0 1 0 0 0 0 0 Ammotrypanella princessa sp.n. 0 0 0 0 0 0 0 0 0 0
Ampharete kerguelensis 6 0 0 0 3 3 0 0 0 0 Ampharetidae sp. 2 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 6 2 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 7 0 1 0 0 1 0 0 0 0 0 Ampharetidae sp. 8 0 0 0 0 1 1 0 0 0 0 Amphicteis gunneri 0 1 0 0 0 0 0 0 0 0 Amphicteis juvenile 1 0 0 0 0 0 0 0 0 0
Amphicteis sp. 1 1 0 0 0 0 1 0 0 0 0 Amphicteis sp. 2 7 0 0 0 0 0 0 0 0 0 Amphicteis sp. 3 0 0 1 0 0 0 0 0 0 0
Amphiduros serratus sp.n. 0 0 0 0 0 0 0 0 0 0 Amythas membranifera 0 0 0 0 0 0 0 0 0 0 Anobothrella antarctica 0 0 0 0 0 1 0 0 0 0
Anobothrella sp. 1 0 0 0 0 1 0 0 0 0 0 Anobothrus gracilis 0 0 0 0 0 2 0 0 0 0
Anobothrus pseudoampharete sp.n. 0 0 0 0 0 3 0 0 0 0 Autolytus gibber 0 0 0 0 0 0 0 0 0 0
Axiokebuita minuta 0 7 0 0 0 1 0 0 0 0 Bathyglycinde sp. 1 BH 0 0 0 0 0 0 0 0 0 0
Braniella palpata 0 1 0 1 1 0 0 0 0 0 Ceratocephale sp. 1BH 0 0 0 0 0 0 0 0 1 0
cf. Amphisamytha 0 0 0 0 0 0 0 0 0 0 cf. Neosabellides 0 0 0 0 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 1 0 0 0 0 0 0 0 0 cf. Thelepides venustus 0 0 0 0 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 0 0 Clavodorum antarcticum 0 0 0 0 0 0 0 0 0 0
Ephesiella antarctica 0 0 0 0 0 0 0 0 0 0 Ephesiella hartmanae sp.n. 0 1 0 0 0 0 0 0 0 0
Eupistella grubei 0 1 1 1 0 0 0 0 0 0 Eupistella sp. 1 0 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 0 0 2 0 0 0 0 Exogone heterosetosa 0 0 0 0 0 0 0 0 0 0
Exogone minuscula 0 0 0 0 0 0 0 0 0 0 Exogone sp. 5 0 0 0 0 0 0 0 0 0 0 Exogone sp. 6 0 0 0 0 0 0 0 0 0 0 Exogone sp. 7 0 0 0 0 0 0 0 0 0 0
Glycera kerguelensis 9 28 5 12 0 1 1 18 7 4 Glyphanostomum scotiarum 0 0 0 0 0 0 0 0 0 0
Goniada maculata 0 0 0 0 0 0 0 0 0 0 Grubianella antarctica 0 0 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 0 0 0 0 0 0 0 Gyptis incompta 0 0 0 0 0 0 0 0 0 0
Hauchiella tribullata 0 0 0 0 0 0 0 0 0 0 Kefersteinia fauveli 0 0 0 0 0 0 0 0 0 0 Kesun abyssorum 13 5 1 2 0 1 0 0 0 0 Leaena antarctica 0 0 0 0 0 1 0 0 0 0 Leaena arenilega 0 0 0 0 0 0 0 0 0 0 Leaena cf. collaris 0 0 0 0 0 0 0 0 0 0
Leaena collaris 0 0 0 0 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 0 0 0 0 0 0 0
Leaena sp. 4 0 0 0 0 0 0 0 0 0 0 Leaena wandelensis 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
216
species 94-14-S 94-14-Ü 94-14-E 102-13-E
102-13-Ü
102-13-S 110-8-S 110-8-E 110-8-Ü 121-11-
E aff. Hesionides 0 0 0 0 0 0 0 0 0 0
Aglaophamus paramalmgreni 0 0 0 0 0 0 1 0 1 0 Aglaophamus trissophyllus 0 0 0 0 0 0 0 1 0 0
Amage sculpta 0 0 0 0 0 0 0 0 0 0 Ammotrypanella arctica 0 0 0 2 0 0 5 5 4 10
Ammotrypanella cirrosa sp.n. 0 0 0 3 0 1 0 1 0 30 Ammotrypanella mcintoshi sp.n. 0 0 0 0 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 0 0 0 0 0 0 2 1 0 0
Ampharete kerguelensis 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 2 0 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 0 0 Amphicteis gunneri 0 0 0 0 0 0 0 0 0 1 Amphicteis juvenile 0 0 0 0 0 0 0 0 0 0
Amphicteis sp. 1 0 0 0 0 0 0 0 0 0 21 Amphicteis sp. 2 0 0 0 0 0 0 0 0 0 0 Amphicteis sp. 3 0 0 0 0 0 0 0 0 0 0
Amphiduros serratus sp.n. 0 0 0 0 0 0 0 0 0 0 Amythas membranifera 0 0 0 0 0 0 0 0 0 0 Anobothrella antarctica 0 0 0 0 0 0 0 0 0 0
Anobothrella sp. 1 0 0 0 0 0 0 0 0 0 0 Anobothrus gracilis 0 0 0 0 1 0 0 0 0 0
Anobothrus pseudoampharete sp.n. 0 0 0 0 0 0 0 0 0 6 Autolytus gibber 0 0 0 0 0 0 0 0 0 0
Axiokebuita minuta 0 0 0 0 0 0 0 0 0 0 Bathyglycinde sp. 1 BH 0 0 0 0 0 0 0 0 0 0
Braniella palpata 0 0 0 0 0 0 0 0 0 1 Ceratocephale sp. 1BH 0 0 0 0 0 0 0 1 0 0
cf. Amphisamytha 0 0 0 0 0 0 0 0 0 0 cf. Neosabellides 0 0 0 0 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 0 0 cf. Thelepides venustus 0 0 0 0 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 0 0 Clavodorum antarcticum 0 0 0 0 0 0 0 0 0 0
Ephesiella antarctica 0 0 0 0 0 0 0 0 0 0 Ephesiella hartmanae sp.n. 0 0 0 0 0 0 0 0 0 0
Eupistella grubei 0 0 0 0 0 0 0 0 0 0 Eupistella sp. 1 0 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 0 0 0 0 0 0 0 Exogone heterosetosa 0 0 0 0 0 0 0 0 0 0
Exogone minuscula 0 0 0 0 0 0 0 0 0 0 Exogone sp. 5 0 0 0 0 0 0 0 0 0 0 Exogone sp. 6 0 0 0 0 0 0 0 0 0 0 Exogone sp. 7 0 0 0 0 0 0 0 0 0 0
Glycera kerguelensis 9 1 9 6 1 1 8 16 5 0 Glyphanostomum scotiarum 0 0 0 0 0 0 0 0 0 0
Goniada maculata 0 0 0 0 0 0 0 0 0 0 Grubianella antarctica 0 0 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 0 0 0 0 0 0 0 Gyptis incompta 0 0 0 0 0 0 0 0 0 0
Hauchiella tribullata 0 0 0 0 0 0 0 0 0 0 Kefersteinia fauveli 0 0 0 0 0 0 0 0 0 0 Kesun abyssorum 1 0 0 3 0 0 3 0 1 10 Leaena antarctica 0 0 0 1 0 0 0 0 0 0 Leaena arenilega 0 0 0 0 0 0 0 0 0 0 Leaena cf. collaris 0 0 0 0 0 0 0 0 0 0
Leaena collaris 0 0 0 0 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 0 0 0 0 0 0 0
Leaena sp. 4 0 0 0 0 0 0 0 0 0 0 Leaena wandelensis 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
217
species 121-11-S 133-2-Ü 133-2-E 133-2-S 142-5-E 142-5-Ü 142-5-S 150-6-E 150-6-Ü 150-6-S
aff. Hesionides 0 0 1 1 0 0 0 0 0 0 Aglaophamus paramalmgreni 0 0 0 0 0 0 0 0 1 0
Aglaophamus trissophyllus 0 0 0 0 0 0 0 2 0 0 Amage sculpta 1 0 1 1 0 0 0 0 0 0
Ammotrypanella arctica 12 0 0 0 0 0 0 0 0 0 Ammotrypanella cirrosa sp.n. 10 0 0 0 0 0 0 0 0 0
Ammotrypanella mcintoshi sp.n. 1 0 0 0 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 1 0 0 0 0 0 0 0 0 0
Ampharete kerguelensis 0 0 1 0 0 0 0 0 0 0 Ampharetidae sp. 2 0 0 0 0 1 0 0 0 0 0 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 0 0 Amphicteis gunneri 0 0 0 0 0 0 0 0 0 0 Amphicteis juvenile 0 0 0 0 0 0 0 0 0 0
Amphicteis sp. 1 11 0 0 1 0 0 0 8 0 0 Amphicteis sp. 2 0 0 0 0 0 0 0 0 0 0 Amphicteis sp. 3 0 0 0 0 0 0 1 1 0 1
Amphiduros serratus sp.n. 0 0 0 0 0 0 0 0 0 0 Amythas membranifera 0 0 0 1 0 0 3 0 0 0 Anobothrella antarctica 0 0 0 0 0 0 0 1 0 0
Anobothrella sp. 1 0 0 0 0 0 0 0 0 0 0 Anobothrus gracilis 0 1 4 2 0 0 0 0 0 0
Anobothrus pseudoampharete sp.n. 3 0 0 4 0 1 0 11 0 1 Autolytus gibber 0 0 0 0 0 0 0 0 0 0
Axiokebuita minuta 0 1 12 2 0 0 0 2 0 0 Bathyglycinde sp. 1 BH 0 0 0 0 0 0 0 0 0 0
Braniella palpata 1 0 29 3 0 0 0 0 0 0 Ceratocephale sp. 1BH 0 0 0 0 0 0 0 0 0 0
cf. Amphisamytha 0 0 0 0 0 0 0 0 0 0 cf. Neosabellides 0 0 0 0 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 0 0 cf. Thelepides venustus 0 0 0 0 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 0 0 Clavodorum antarcticum 0 0 0 0 0 0 1 1 0 0
Ephesiella antarctica 0 0 0 0 0 0 1 0 0 0 Ephesiella hartmanae sp.n. 0 0 3 0 0 0 0 1 0 0
Eupistella grubei 0 0 4 0 0 0 0 0 0 0 Eupistella sp. 1 0 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 0 0 0 0 0 0 0 Exogone heterosetosa 0 0 0 0 0 0 0 0 0 0
Exogone minuscula 0 0 1 0 0 0 0 0 0 0 Exogone sp. 5 0 0 0 0 0 0 0 0 0 0 Exogone sp. 6 0 0 15 1 0 0 0 0 0 0 Exogone sp. 7 0 0 37 5 0 0 0 0 0 0
Glycera kerguelensis 0 3 13 3 0 0 0 6 0 0 Glyphanostomum scotiarum 0 0 0 0 0 0 0 0 0 0
Goniada maculata 0 0 0 0 0 0 0 0 0 0 Grubianella antarctica 0 0 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 0 0 0 0 0 0 0 Gyptis incompta 0 1 639 96 0 0 0 0 0 0
Hauchiella tribullata 0 0 0 0 0 0 0 0 0 0 Kefersteinia fauveli 0 6 160 37 0 0 0 0 0 0 Kesun abyssorum 4 0 1 1 0 0 4 24 1 1 Leaena antarctica 0 1 1 0 0 0 0 0 0 0 Leaena arenilega 0 0 0 0 0 0 0 0 0 0 Leaena cf. collaris 0 0 0 0 0 0 0 0 0 0
Leaena collaris 0 1 46 17 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 0 0 0 0 0 0 0
Leaena sp. 4 0 0 0 0 0 0 0 0 0 0 Leaena wandelensis 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
218
species 151-7-E 151-7-S 152-6-S 152-6-E 153-7-S 153-7-E 154-9-E 154-9-S total
aff. Hesionides 0 0 0 0 0 0 0 0 11 Aglaophamus paramalmgreni 0 2 0 1 8 7 0 0 36
Aglaophamus trissophyllus 0 0 0 0 0 0 2 2 100 Amage sculpta 0 0 0 0 0 0 0 0 15
Ammotrypanella arctica 0 0 0 0 2 0 0 3 54 Ammotrypanella cirrosa sp.n. 0 0 0 0 2 2 0 1 60
Ammotrypanella mcintoshi sp.n. 0 0 0 0 1 0 0 0 5 Ammotrypanella princessa sp.n. 0 0 0 0 0 0 0 0 11
Ampharete kerguelensis 0 0 0 0 0 0 0 0 28 Ampharetidae sp. 2 0 0 0 0 5 0 0 0 11 Ampharetidae sp. 4 1 0 0 0 0 0 0 0 1 Ampharetidae sp. 5 0 1 0 0 0 0 0 0 1 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 2 Amphicteis gunneri 0 0 0 0 0 0 0 0 3 Amphicteis juvenile 0 0 0 0 0 0 0 0 1
Amphicteis sp. 1 0 0 0 0 3 0 0 0 60 Amphicteis sp. 2 0 0 0 0 0 0 0 0 7 Amphicteis sp. 3 0 0 0 0 0 0 0 1 10
Amphiduros serratus sp.n. 0 0 2 0 0 0 0 0 2 Amythas membranifera 0 0 0 0 0 0 0 0 4 Anobothrella antarctica 0 0 0 0 0 0 0 0 5
Anobothrella sp. 1 0 0 0 0 0 0 0 0 1 Anobothrus gracilis 0 0 0 0 0 0 2 0 12
Anobothrus pseudoampharete sp.n. 0 0 0 0 0 0 0 1 135 Autolytus gibber 0 0 0 0 0 0 0 0 1
Axiokebuita minuta 0 0 0 0 0 0 0 0 33 Bathyglycinde sp. 1 BH 0 0 0 0 0 0 0 0 1
Braniella palpata 0 0 0 0 0 0 0 0 42 Ceratocephale sp. 1BH 0 0 0 0 0 1 1 0 5
cf. Amphisamytha 0 0 0 0 0 0 0 0 1 cf. Neosabellides 0 0 0 0 0 0 0 0 1
cf. Nicon 0 0 0 0 0 0 0 1 1 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 1 cf. Thelepides venustus 0 0 0 0 0 0 0 0 1 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 1 Clavodorum antarcticum 0 0 0 0 0 0 0 0 2
Ephesiella antarctica 0 0 0 0 1 1 0 0 3 Ephesiella hartmanae sp.n. 0 0 0 0 0 0 0 0 8
Eupistella grubei 0 0 0 0 0 0 0 0 16 Eupistella sp. 1 0 0 0 0 0 0 0 0 2
Eusamythella sp. 1 0 0 0 0 0 0 0 0 3 Exogone heterosetosa 0 0 0 0 0 0 0 0 1
Exogone minuscula 7 11 0 0 1 0 0 0 28 Exogone sp. 5 0 0 0 0 0 0 0 0 5 Exogone sp. 6 0 0 0 0 0 0 0 0 16 Exogone sp. 7 0 0 0 0 1 3 0 0 46
Glycera kerguelensis 0 0 0 0 2 2 3 2 276 Glyphanostomum scotiarum 0 0 0 0 0 0 0 0 4
Goniada maculata 0 0 0 0 0 0 0 0 1 Grubianella antarctica 0 0 0 0 1 0 0 0 3
Grubianella sp. 1 0 0 0 0 0 0 0 0 1 Gyptis incompta 0 0 0 0 0 0 0 0 784
Hauchiella tribullata 0 0 0 0 0 0 0 0 2 Kefersteinia fauveli 7 4 0 0 0 0 0 0 293 Kesun abyssorum 0 0 0 0 41 13 2 9 167 Leaena antarctica 0 0 0 0 4 3 0 0 11 Leaena arenilega 0 0 0 0 0 1 0 0 6 Leaena cf. collaris 0 0 0 0 0 0 0 0 4
Leaena collaris 0 0 0 0 0 0 0 0 66 Leaena pseudobranchia 0 0 0 0 0 0 0 0 3
Leaena sp. 4 0 0 0 0 0 0 0 0 1 Leaena wandelensis 0 0 0 0 0 0 3 0 4
APPENDIX- TABLES
219
species 16-10-S 16-10-E 21-7-S 21-7-E 59-5-S 59-5-E 59-5-Ü 74-6-S 74-6-E 78-9-E
Melinna cristata 0 0 0 0 0 0 0 0 0 0 Micropodarke cylindripalpata sp.n. 1 0 2 0 0 5 0 0 0 0
Mugga sp. 1BH 1 2 0 0 0 0 0 0 0 0 Muggoides cf. cinctus 0 0 0 0 0 0 0 0 0 0 Muggoides sp. 1BH 0 2 0 0 0 0 0 0 0 0
Neosabellides elongatus 0 0 0 1 0 0 0 1 3 2 Octobranchus antarcticus 0 0 0 0 0 0 0 8 11 0
Oligobregma blakei 0 0 0 0 0 0 0 0 0 0 Oligobregma collare 0 0 0 0 0 1 0 1 0 7
Oligobregma hartmanae 0 0 0 0 3 0 0 0 0 0 Oligobregma pseudocollare 0 0 0 0 0 0 0 0 0 0 Oligobregma quadrispinosa 0 0 0 1 5 1 0 0 0 0
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 4 0 0 0 0 0 0 0 0 3
Ophelina breviata 0 1 0 0 0 0 0 6 10 1 Ophelina gymnopyge 2 0 0 0 1 0 0 0 0 0 Ophelina nematoides 1 1 0 0 0 0 0 2 16 0
Ophelina robusta sp.n. 0 0 0 0 0 0 0 0 0 10 Ophelina scaphigera 4 0 0 0 0 1 0 0 0 0
Ophelina setigera 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 0 0 0 0 0 0 0 0
Ophiodromus calligocervix sp.n. 2 0 1 2 0 2 0 0 0 11 Ophiodromus comatus 0 0 0 0 0 0 0 3 22 0
Parasyllidea delicata sp.n. 2 0 3 0 0 0 0 0 0 0 Phalacrostemma elegans 0 0 0 0 0 0 0 0 0 0
Phisidia rubrolineata 0 0 0 0 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 0 0
Phyllocomus crocea 0 0 0 0 1 1 0 0 2 0 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 1 0
Pionosyllis comosa 0 0 0 0 0 0 0 5 2 0 Pionosyllis epipharynx 0 0 0 0 0 0 0 4 10 0
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 0 0 Pista corrientis 0 0 0 0 0 0 0 1 0 0 Pista cristata 0 0 0 0 0 0 0 0 2 0 Pista spinifera 0 0 0 0 0 0 0 2 3 0
Polycirrus cf. antarcticus 0 0 0 0 0 0 0 1 0 0 Polycirrus insignis 0 0 0 0 0 0 0 4 9 1
Polycirrus sp. 1 0 0 0 0 1 0 0 0 0 0 Polycirrus sp. 2 0 0 0 0 0 0 0 0 1 0
Proceaea cf. mclearanus 0 0 0 0 0 0 0 0 1 0 Proclea cf. graffii 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma bransfieldium 0 0 0 0 0 0 0 0 2 0 Pseudoscalibregma papilia sp.n. 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma ursapium 0 0 0 0 0 0 0 0 0 0 Samytha sp. 1 0 0 0 0 0 0 0 0 1 0
Scalibregma inflatum 0 0 0 0 0 0 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 0 0
Sosanopsis kerguelensis 3 1 6 0 2 1 1 1 2 0 Sosanopsis sp. 1 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 3 20 0 Sphaerodoropsis parva 1 0 0 1 1 4 1 11 76 3
Sphaerodoropsis polypapillata 0 0 0 0 0 0 0 0 0 0 Sphaerodoropsis simplex sp.n. 0 0 0 0 0 0 0 0 3 0
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 1 0 Sphaerosyllis joinvillensis 0 0 0 0 0 0 0 2 10 0
Sphaerosyllis lateropapillata uteae 0 0 0 0 0 0 0 30 126 0 Syllides articulosus 0 0 0 0 0 0 0 0 1 0 Terebellidae sp. 3 0 0 0 1 0 0 0 0 0 0
Terebellides stroemii 1 0 0 0 0 1 0 0 6 10 Thelepides koehleri 0 0 0 0 0 0 0 0 2 0 Thelepides venustus 0 0 0 0 0 0 0 1 2 0 Thelepodinae sp. 1 0 0 0 0 0 0 0 1 0 0
Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 0 0 0 0 0 0 1 3 3 3 Typosyllis cf. hyalina 0 0 0 0 0 0 0 0 1 0 Typosyllis variegata 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
220
species 78-9-S 80-9-E 80-9-S 80-9-Ü 81-8-E 81-8-S 81-8-Ü 88-8-E 88-8-S 88-8-Ü
Melinna cristata 0 2 1 1 0 0 0 0 0 0 Micropodarke cylindripalpata sp.n. 0 3 2 0 4 0 0 6 0 0
Mugga sp. 1BH 0 0 0 0 0 0 0 0 0 0 Muggoides cf. cinctus 0 0 0 0 0 8 3 0 0 0 Muggoides sp. 1BH 0 0 0 0 0 0 0 0 0 0
Neosabellides elongatus 0 1 0 0 1 0 0 0 0 0 Octobranchus antarcticus 0 3 0 1 0 0 0 0 0 0
Oligobregma blakei 0 0 0 0 0 0 0 0 0 0 Oligobregma collare 0 1 0 0 0 0 0 0 0 0
Oligobregma hartmanae 0 0 0 0 0 0 0 0 0 0 Oligobregma pseudocollare 0 2 0 0 0 0 0 0 0 0 Oligobregma quadrispinosa 0 0 0 0 0 0 0 1 0 0
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 7 0 0 0 0 4 0 0 0 0
Ophelina breviata 0 0 0 0 0 0 0 0 0 0 Ophelina gymnopyge 0 0 0 0 1 2 0 0 0 0 Ophelina nematoides 0 1 0 0 0 0 0 0 0 0
Ophelina robusta sp.n. 0 2 0 0 0 0 0 0 0 0 Ophelina scaphigera 0 0 0 0 0 0 0 0 0 0
Ophelina setigera 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 0 0 0 0 0 0 0 0
Ophiodromus calligocervix sp.n. 3 10 2 0 0 0 0 6 2 0 Ophiodromus comatus 0 3 4 0 3 0 0 0 0 0
Parasyllidea delicata sp.n. 0 3 1 0 0 0 0 0 0 0 Phalacrostemma elegans 0 1 0 0 0 0 0 0 0 0
Phisidia rubrolineata 0 0 0 0 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 0 0
Phyllocomus crocea 0 0 0 0 0 2 0 0 0 0 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 0 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 0 0 0 0 0 0 0 0
Pionosyllis sp. 1 1 0 0 0 0 0 0 0 0 0 Pista corrientis 0 0 0 0 0 0 0 0 0 0 Pista cristata 0 0 0 0 0 0 0 0 0 0 Pista spinifera 0 0 0 0 0 0 0 0 0 0
Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 0 0 Polycirrus insignis 2 0 0 0 0 0 0 0 0 0
Polycirrus sp. 1 0 0 0 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 0 0 0 0 0 0 0
Proceaea cf. mclearanus 0 0 0 0 0 0 0 0 0 0 Proclea cf. graffii 0 1 0 0 0 0 0 0 0 0
Pseudoscalibregma bransfieldium 0 1 0 0 0 0 0 1 0 0 Pseudoscalibregma papilia sp.n. 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma ursapium 0 3 0 0 0 0 0 0 0 0 Samytha sp. 1 0 0 0 0 0 0 0 0 0 0
Scalibregma inflatum 0 0 0 0 0 0 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 1 0 0 0 0 0 0
Sosanopsis kerguelensis 1 1 0 1 0 0 0 1 0 0 Sosanopsis sp. 1 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 0 0 Sphaerodoropsis parva 0 3 1 2 14 12 2 0 0 0
Sphaerodoropsis polypapillata 0 3 0 1 0 0 0 0 0 0 Sphaerodoropsis simplex sp.n. 0 1 0 0 0 0 0 0 0 0
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 0 0 0 0 0 0 0 0
Sphaerosyllis lateropapillata uteae 0 0 0 0 0 0 0 0 0 0 Syllides articulosus 0 0 0 0 0 0 0 0 0 0 Terebellidae sp. 3 0 0 0 0 0 0 0 0 0 0
Terebellides stroemii 2 2 2 1 1 1 0 0 0 0 Thelepides koehleri 0 0 0 0 0 0 0 0 0 0 Thelepides venustus 0 0 0 0 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 0 0 0 0 0 0 0 0
Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 1 0 0 0 0 0 0 0 0 0 Typosyllis cf. hyalina 0 0 0 0 0 0 0 0 0 0 Typosyllis variegata 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
221
species 94-14-S 94-14-Ü 94-14-E 102-13-E
102-13-Ü
102-13-S 110-8-S 110-8-E 110-8-Ü 121-11-
E Melinna cristata 0 0 0 0 0 0 0 0 0 0
Micropodarke cylindripalpata sp.n. 1 0 0 0 0 0 0 2 0 1 Mugga sp. 1BH 0 0 0 0 0 0 0 0 0 0
Muggoides cf. cinctus 0 0 0 0 0 0 0 0 0 0 Muggoides sp. 1BH 0 0 0 0 4 0 0 0 0 0
Neosabellides elongatus 0 0 0 0 0 0 0 0 0 0 Octobranchus antarcticus 0 0 0 0 0 0 0 0 0 0
Oligobregma blakei 0 0 0 0 0 0 0 0 0 1 Oligobregma collare 0 0 0 0 0 0 1 0 0 7
Oligobregma hartmanae 0 0 0 0 0 0 0 0 0 0 Oligobregma pseudocollare 0 0 0 0 0 0 0 0 0 1 Oligobregma quadrispinosa 2 0 0 0 0 0 1 0 0 1
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 0 0 0 0 0 0 2 2 0 2
Ophelina breviata 0 0 0 1 0 0 0 1 0 6 Ophelina gymnopyge 0 0 0 0 0 0 0 1 2 1 Ophelina nematoides 0 0 0 0 0 0 0 0 0 0
Ophelina robusta sp.n. 0 0 0 0 0 0 0 0 0 3 Ophelina scaphigera 0 0 0 0 0 0 0 0 0 0
Ophelina setigera 0 0 0 0 0 0 0 0 0 1 Ophelina sp. 5 0 0 0 0 0 0 0 0 0 0
Ophiodromus calligocervix sp.n. 2 0 2 1 0 0 0 3 0 0 Ophiodromus comatus 0 0 0 0 0 0 0 0 0 1
Parasyllidea delicata sp.n. 0 0 0 0 0 0 0 2 0 0 Phalacrostemma elegans 0 0 0 0 0 0 0 0 0 0
Phisidia rubrolineata 0 0 0 0 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 0 0
Phyllocomus crocea 0 0 0 0 0 0 0 0 0 0 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 0 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 0 0 0 0 0 0 0 0
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 0 0 Pista corrientis 0 0 0 0 0 0 0 0 0 0 Pista cristata 0 0 0 0 0 0 0 0 0 0 Pista spinifera 0 0 0 0 0 0 0 0 0 0
Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 0 0 Polycirrus insignis 0 0 0 0 0 0 0 0 0 0
Polycirrus sp. 1 0 0 0 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 0 0 0 0 0 0 0
Proceaea cf. mclearanus 0 0 0 0 0 0 0 0 0 0 Proclea cf. graffii 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma bransfieldium 0 0 0 0 0 0 0 0 0 0 Pseudoscalibregma papilia sp.n. 0 0 0 0 0 0 0 0 0 1
Pseudoscalibregma ursapium 0 0 0 0 0 0 0 0 0 0 Samytha sp. 1 0 0 0 0 0 0 0 0 0 0
Scalibregma inflatum 0 0 0 0 0 0 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 0 0
Sosanopsis kerguelensis 1 0 2 1 0 0 0 2 0 0 Sosanopsis sp. 1 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 0 0 Sphaerodoropsis parva 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis polypapillata 0 0 0 0 0 0 0 0 0 5 Sphaerodoropsis simplex sp.n. 0 0 0 0 0 0 0 0 0 0
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 0 0 0 0 0 0 0 0
Sphaerosyllis lateropapillata uteae 0 0 0 0 0 0 0 0 0 0 Syllides articulosus 0 0 0 0 0 0 0 0 0 0 Terebellidae sp. 3 0 0 0 0 0 0 0 0 0 0
Terebellides stroemii 2 0 0 1 0 0 0 3 0 12 Thelepides koehleri 0 0 0 0 0 0 0 0 0 0 Thelepides venustus 0 0 0 0 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 0 0 0 0 0 0 0 0
Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 0 0 0 0 0 0 0 0 0 0 Typosyllis cf. hyalina 0 0 0 0 0 0 0 0 0 0 Typosyllis variegata 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
222
species 121-11-S 133-2-Ü 133-2-E 133-2-S 142-5-E 142-5-Ü 142-5-S 150-6-E 150-6-Ü 150-6-S
Melinna cristata 0 0 0 0 0 0 0 3 0 1 Micropodarke cylindripalpata sp.n. 0 0 0 0 0 0 0 3 0 0
Mugga sp. 1BH 0 0 0 0 0 0 0 0 0 0 Muggoides cf. cinctus 0 0 0 0 0 0 0 0 0 0 Muggoides sp. 1BH 0 0 0 0 0 0 0 0 0 0
Neosabellides elongatus 0 0 1 0 0 0 0 0 0 0 Octobranchus antarcticus 0 0 92 27 0 0 0 0 0 0
Oligobregma blakei 0 0 1 1 0 0 0 0 0 0 Oligobregma collare 4 0 2 1 1 0 0 1 0 0
Oligobregma hartmanae 0 0 1 1 0 0 0 0 0 1 Oligobregma pseudocollare 0 0 7 1 0 0 0 1 0 0 Oligobregma quadrispinosa 0 0 0 1 0 0 0 0 0 0
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 5 0 0 0 0 0 0 0 0 0
Ophelina breviata 0 0 18 1 0 0 0 1 0 0 Ophelina gymnopyge 1 0 14 0 0 0 0 0 0 0 Ophelina nematoides 0 0 0 0 0 0 0 0 0 0
Ophelina robusta sp.n. 1 0 0 0 0 0 0 5 0 0 Ophelina scaphigera 0 0 2 0 0 0 0 0 0 0
Ophelina setigera 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 10 0 0 0 0 2 0 0
Ophiodromus calligocervix sp.n. 1 0 0 0 0 0 0 0 0 0 Ophiodromus comatus 3 0 37 6 0 0 0 2 0 1
Parasyllidea delicata sp.n. 0 0 0 0 1 0 2 5 0 2 Phalacrostemma elegans 0 0 0 0 0 0 0 0 0 0
Phisidia rubrolineata 0 32 184 50 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 1 0 0
Phyllocomus crocea 0 0 0 0 0 0 0 0 0 0 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 0 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 20 4 0 0 0 1 0 0
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 0 0 Pista corrientis 0 0 0 0 0 0 0 0 0 0 Pista cristata 0 0 0 0 0 0 0 0 0 0 Pista spinifera 0 0 0 0 0 0 0 0 0 0
Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 0 0 Polycirrus insignis 0 5 192 44 0 1 1 0 0 1
Polycirrus sp. 1 0 0 0 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 0 0 0 0 0 0 0
Proceaea cf. mclearanus 0 0 0 0 0 0 0 0 0 0 Proclea cf. graffii 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma bransfieldium 0 0 2 0 0 0 0 0 1 0 Pseudoscalibregma papilia sp.n. 0 0 0 0 0 0 1 5 0 0
Pseudoscalibregma ursapium 0 0 4 1 0 0 0 0 0 0 Samytha sp. 1 0 0 0 0 0 0 0 0 0 0
Scalibregma inflatum 0 0 1 0 0 0 0 2 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 0 0
Sosanopsis kerguelensis 3 0 6 2 0 0 0 1 0 1 Sosanopsis sp. 1 0 0 0 0 0 0 0 4 0 1
Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 0 0 Sphaerodoropsis parva 0 5 158 26 0 0 0 2 0 0
Sphaerodoropsis polypapillata 0 0 0 0 0 0 0 0 0 0 Sphaerodoropsis simplex sp.n. 0 0 0 0 0 0 0 1 0 0
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 243 42 0 0 0 0 0 0
Sphaerosyllis lateropapillata uteae 0 0 200 17 0 0 0 4 1 0 Syllides articulosus 0 0 288 29 0 0 0 0 0 0 Terebellidae sp. 3 0 0 0 0 0 0 0 0 0 0
Terebellides stroemii 3 0 1 0 3 0 3 4 0 0 Thelepides koehleri 0 0 0 0 0 0 0 0 0 0 Thelepides venustus 0 0 0 0 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 0 0 0 0 0 0 0 0
Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 0 0 2 0 0 0 0 3 0 0 Typosyllis cf. hyalina 0 0 5 0 0 0 0 0 0 0 Typosyllis variegata 0 4 49 7 0 0 0 0 0 0
APPENDIX- TABLES
223
species 151-7-E 151-7-S 152-6-S 152-6-E 153-7-S 153-7-E 154-9-E 154-9-S total
Melinna cristata 0 0 0 0 0 0 0 0 8 Micropodarke cylindripalpata sp.n. 0 0 0 0 0 0 2 0 32
Mugga sp. 1BH 0 0 0 0 0 0 1 0 4 Muggoides cf. cinctus 0 0 0 0 0 0 0 0 11 Muggoides sp. 1BH 0 0 0 0 0 0 0 0 6
Neosabellides elongatus 0 0 0 0 0 2 0 0 12 Octobranchus antarcticus 0 0 0 0 1 0 0 0 143
Oligobregma blakei 0 0 0 0 0 0 0 0 3 Oligobregma collare 0 1 0 0 10 0 1 0 39
Oligobregma hartmanae 0 0 0 0 0 0 0 0 6 Oligobregma pseudocollare 0 0 0 0 1 1 0 0 14 Oligobregma quadrispinosa 0 1 0 0 9 1 0 0 24
Oligobregma sp. 1 0 1 0 0 0 0 0 0 1 Ophelina ammotrypanella sp.n. 0 0 0 0 0 0 4 4 37
Ophelina breviata 0 0 0 0 1 0 1 0 48 Ophelina gymnopyge 0 0 0 0 0 0 0 4 29 Ophelina nematoides 0 0 0 0 0 1 0 0 22
Ophelina robusta sp.n. 2 0 0 0 6 1 0 0 30 Ophelina scaphigera 0 0 0 0 0 0 0 0 7
Ophelina setigera 0 0 0 0 0 0 0 0 1 Ophelina sp. 5 0 0 0 0 0 0 0 5 17
Ophiodromus calligocervix sp.n. 0 0 0 0 2 5 1 1 59 Ophiodromus comatus 12 5 0 0 0 1 0 0 103
Parasyllidea delicata sp.n. 0 0 0 0 0 1 0 0 22 Phalacrostemma elegans 0 0 0 0 0 0 0 0 1
Phisidia rubrolineata 0 0 0 0 0 0 0 0 266 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 1
Phyllocomus crocea 1 1 0 0 0 0 0 0 8 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 1
Pionosyllis comosa 0 0 0 0 0 0 0 0 7 Pionosyllis epipharynx 0 0 0 0 0 1 2 0 42
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 1 Pista corrientis 0 0 0 0 0 0 0 0 1 Pista cristata 0 0 0 0 0 0 0 0 2 Pista spinifera 0 0 0 0 0 0 0 0 5
Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 1 Polycirrus insignis 0 0 0 0 0 0 0 1 261
Polycirrus sp. 1 0 0 0 0 0 0 0 0 1 Polycirrus sp. 2 0 0 0 0 0 0 0 0 1
Proceaea cf. mclearanus 0 0 0 0 0 0 0 0 1 Proclea cf. graffii 0 0 0 0 0 0 0 0 1
Pseudoscalibregma bransfieldium 0 1 0 0 0 1 0 0 9 Pseudoscalibregma papilia sp.n. 0 0 0 0 1 0 0 0 8
Pseudoscalibregma ursapium 0 1 0 0 0 0 0 0 9 Samytha sp. 1 0 0 0 0 0 0 0 0 1
Scalibregma inflatum 0 0 0 0 0 0 0 0 3 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 1
Sosanopsis kerguelensis 0 0 0 0 2 4 1 0 47 Sosanopsis sp. 1 0 0 0 0 2 0 0 0 7
Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 23 Sphaerodoropsis parva 6 5 0 1 3 0 2 0 340
Sphaerodoropsis polypapillata 0 0 0 0 0 2 0 0 11 Sphaerodoropsis simplex sp.n. 0 0 0 0 0 0 0 0 5
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 1 Sphaerosyllis joinvillensis 0 0 0 0 0 0 0 0 297
Sphaerosyllis lateropapillata uteae 3 1 0 0 8 30 0 0 420 Syllides articulosus 0 0 0 0 0 0 0 0 318 Terebellidae sp. 3 0 0 0 0 0 0 0 0 1
Terebellides stroemii 0 0 0 0 0 0 0 0 59 Thelepides koehleri 0 0 0 0 0 0 0 0 2 Thelepides venustus 0 0 0 0 0 0 0 0 3 Thelepodinae sp. 1 0 0 0 0 0 0 0 0 1
Travisia cf. lithophila 0 0 0 0 0 0 0 1 1 Travisia kerguelensis 0 0 0 0 1 0 6 3 26 Typosyllis cf. hyalina 0 0 0 0 0 0 0 0 6 Typosyllis variegata 0 0 0 0 0 0 0 0 60
APPENDIX- TABLES
224
App.-Tab. 6: Species assemblage matrix for EBS-stations of the expeditions ANDEEP I-III
species 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1
aff. Hesionides 0 0 0 0 0 0 0 0 0 0 Aglaophamus paramalmgreni 0 5 6 0 0 0 0 0 0 0
Aglaophamus trissophyllus 0 2 27 0 0 0 0 0 0 0 Amage sculpta 0 1 0 0 0 0 0 0 0 0
Ammotrypanella arctica 0 2 0 7 0 0 1 0 0 0 Ammotrypanella cirrosa sp.n, 0 4 0 64 0 0 2 0 1 0
Ammotrypanella mcintoshi sp.n. 0 0 0 0 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 0 1 0 0 0 0 1 1 0 0
Ampharete kerguelensis 2 0 7 0 0 0 0 0 0 0 Ampharetidae sp. 1 0 0 0 0 0 0 0 0 1 0 Ampharetidae sp. 2 0 0 0 0 0 0 2 0 0 0 Ampharetidae sp. 3 0 0 0 0 0 0 2 0 0 0 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 0 0 Amphicteis gunneri 0 0 1 0 0 0 0 0 0 0 Amphicteis juvenile 0 0 0 0 0 0 0 0 0 0
Amphicteis sp. 1 1 0 0 0 0 0 0 0 0 0 Amphicteis sp. 2 0 0 0 0 0 0 0 0 0 0 Amphicteis sp. 3 0 0 0 0 0 0 0 0 0 0
Amphiduros serratus sp.n. 0 0 0 0 2 0 1 0 0 0 Amphiduros sp. 1 0 0 12 1 0 0 0 0 1 0
Amythas membranifera 0 0 0 0 0 0 0 0 0 0 Anobothrella antarctica 0 0 0 0 0 0 0 0 0 1
Anobothrella sp. 1 0 0 0 0 0 0 0 0 0 0 Anobothrus gracilis 0 0 0 0 0 0 0 0 0 0
Anobothrus pseudoampharete sp.n. 0 1 0 0 0 1 0 0 0 7 Asclerocheilus ashworthi 0 0 1 0 0 0 0 0 0 0
Autolytus gibber 0 0 0 0 0 0 0 0 0 0 Axiokebuita millsii 0 0 0 0 0 0 0 0 0 62
Axiokebuita minuta 0 0 1 6 0 1 3 0 11 0 Bathyglycinde sp. 1 BH 0 3 0 0 0 0 0 0 0 0
Bathyglycinde sp. 2 0 1 0 0 0 0 0 0 0 0 Brania sp. 1 BH 0 0 0 2 1 0 0 0 0 0 Braniella palpata 0 0 0 0 0 0 0 0 2 0
Ceratocephale sp. 1BH 0 0 0 0 0 0 0 0 0 0 cf. Amphisamytha 0 0 0 0 0 0 0 0 0 0
cf. Autolytus simplex stolon 0 0 1 0 0 0 0 0 0 0 cf. Neosabellides 0 0 0 0 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 0 0 cf. Thelepides venustus 0 0 0 0 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 0 0 Clavodorum antarcticum 0 0 1 0 0 0 0 0 0 0
Ephesiella antarctica 0 1 5 0 0 0 0 0 0 8 Ephesiella hartmanae sp.n. 0 0 0 0 0 0 0 0 0 3
Eupistella grubei 0 0 0 0 0 0 0 0 0 0 Eupistella sp. 1 0 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 4 0 0 0 0 0 0 0 0 Exogone heterosetosa 0 1 0 0 0 0 0 0 0 0
Exogone minuscula 0 0 0 0 0 0 0 0 0 0 Exogone sp. 2 BH 0 0 1 0 0 0 0 0 0 0
APPENDIX- TABLES
225
species 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13
aff. Hesionides 4 3 1 0 0 0 0 1 0 0 Aglaophamus paramalmgreni 0 0 14 0 1 0 0 0 0 0
Aglaophamus trissophyllus 0 0 0 73 18 2 0 0 0 0 Amage sculpta 0 8 0 1 0 3 0 0 0 0
Ammotrypanella arctica 6 0 4 0 1 0 0 0 0 2 Ammotrypanella cirrosa sp.n, 6 0 2 0 2 0 0 0 0 4
Ammotrypanella mcintoshi sp.n. 3 1 1 2 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 1 0 0 0 1 0 1 0 0 0
Ampharete kerguelensis 13 0 1 0 7 0 6 0 0 0 Ampharetidae sp. 1 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 2 0 1 0 0 2 0 0 0 0 0 Ampharetidae sp. 3 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 2 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 1 1 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 2 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 0 0 Amphicteis gunneri 0 0 0 0 1 1 0 0 0 0 Amphicteis juvenile 0 0 0 0 1 0 0 0 0 0
Amphicteis sp. 1 0 0 0 0 15 0 1 0 0 0 Amphicteis sp. 2 0 0 0 0 7 0 0 0 0 0 Amphicteis sp. 3 0 0 0 5 0 1 0 0 0 0
Amphiduros serratus sp.n. 0 0 0 0 0 0 0 0 0 0 Amphiduros sp. 1 0 0 0 0 0 0 0 0 0 0
Amythas membranifera 0 0 0 0 0 0 0 0 0 0 Anobothrella antarctica 0 0 0 3 0 0 1 0 0 0
Anobothrella sp. 1 0 0 0 0 0 0 1 0 0 0 Anobothrus gracilis 0 0 0 0 0 0 2 0 0 1
Anobothrus pseudoampharete sp.n. 0 1 0 104 0 0 3 0 0 0 Asclerocheilus ashworthi 0 0 0 0 0 0 0 0 0 0
Autolytus gibber 0 0 0 1 0 0 0 0 0 0 Axiokebuita millsii 0 0 0 0 0 0 0 0 0 0
Axiokebuita minuta 0 0 0 8 0 7 1 0 0 0 Bathyglycinde sp. 1 BH 1 0 0 0 0 0 0 0 0 0
Bathyglycinde sp. 2 0 0 0 0 0 0 0 0 0 0 Brania sp. 1 BH 0 0 0 0 0 0 0 0 0 0 Braniella palpata 0 1 1 3 0 2 1 0 0 0
Ceratocephale sp. 1BH 0 0 0 0 1 0 0 1 0 0 cf. Amphisamytha 0 0 0 1 0 0 0 0 0 0
cf. Autolytus simplex stolon 0 0 0 0 0 0 0 0 0 0 cf. Neosabellides 0 0 0 1 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 0 0 0 0 1 0 0 0 0 cf. Thelepides venustus 0 0 0 1 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 1 0 0 0 0 0 0 Clavodorum antarcticum 0 0 0 0 0 0 0 0 0 0
Ephesiella antarctica 0 0 0 0 0 0 0 0 0 0 Ephesiella hartmanae sp.n. 1 0 0 2 0 1 0 0 0 0
Eupistella grubei 0 0 0 9 0 3 0 0 0 0 Eupistella sp. 1 2 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 1 0 0 2 0 0 0 Exogone heterosetosa 0 0 0 1 0 0 0 0 0 0
Exogone minuscula 0 0 0 8 0 0 0 0 0 0 Exogone sp. 2 BH 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
226
species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total
aff. Hesionides 0 0 2 0 0 0 0 0 0 11 Aglaophamus paramalmgreni 2 0 0 0 1 2 1 15 0 47
Aglaophamus trissophyllus 1 0 0 0 2 0 0 0 4 129 Amage sculpta 0 1 2 0 0 0 0 0 0 16
Ammotrypanella arctica 14 22 0 0 0 0 0 2 3 64 Ammotrypanella cirrosa sp.n, 1 40 0 0 0 0 0 4 1 131
Ammotrypanella mcintoshi sp.n. 3 1 0 0 0 0 0 0 0 11 Ammotrypanella princessa sp.n. 0 1 0 0 0 0 0 1 0 8
Ampharete kerguelensis 0 0 1 0 0 0 0 0 0 37 Ampharetidae sp. 1 0 0 0 0 0 0 0 0 0 1 Ampharetidae sp. 2 0 2 0 1 0 0 0 5 0 13 Ampharetidae sp. 3 0 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 4 0 0 0 0 0 1 0 0 0 1 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 0 2 Ampharetidae sp. 8 0 0 0 0 0 1 0 0 0 1 Amphicteis gunneri 0 1 0 0 0 0 0 0 0 4 Amphicteis juvenile 0 0 0 0 0 0 0 0 0 1
Amphicteis sp. 1 0 32 1 0 8 0 0 3 0 61 Amphicteis sp. 2 0 0 0 0 0 0 0 0 0 7 Amphicteis sp. 3 0 0 0 1 2 0 0 0 1 10
Amphiduros serratus sp.n. 0 0 0 0 0 0 2 0 0 5 Amphiduros sp. 1 0 0 0 0 0 0 0 0 0 14
Amythas membranifera 0 0 1 3 0 0 0 0 0 4 Anobothrella antarctica 0 0 0 0 1 0 0 0 0 6
Anobothrella sp. 1 0 0 0 0 0 0 0 0 0 1 Anobothrus gracilis 0 0 7 0 0 0 0 0 2 12
Anobothrus pseudoampharete sp.n. 0 9 4 1 12 0 0 0 1 144 Asclerocheilus ashworthi 0 0 0 0 0 0 0 0 0 1
Autolytus gibber 0 0 0 0 0 0 0 0 0 1 Axiokebuita millsii 0 0 0 0 0 0 0 0 0 62
Axiokebuita minuta 0 0 15 0 2 0 0 0 0 55 Bathyglycinde sp. 1 BH 0 0 0 0 0 0 0 0 0 4
Bathyglycinde sp. 2 0 0 0 0 0 0 0 0 0 1 Brania sp. 1 BH 0 0 0 0 0 0 0 0 0 3 Braniella palpata 0 2 32 0 0 0 0 0 0 44
Ceratocephale sp. 1BH 1 0 0 0 0 0 0 1 1 5 cf. Amphisamytha 0 0 0 0 0 0 0 0 0 1
cf. Autolytus simplex stolon 0 0 0 0 0 0 0 0 0 1 cf. Neosabellides 0 0 0 0 0 0 0 0 0 1
cf. Nicon 0 0 0 0 0 0 0 0 1 1 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 0 1 cf. Thelepides venustus 0 0 0 0 0 0 0 0 0 1 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 0 1 Clavodorum antarcticum 0 0 0 1 1 0 0 0 0 3
Ephesiella antarctica 0 0 0 1 0 0 0 2 0 17 Ephesiella hartmanae sp.n. 0 0 3 0 1 0 0 0 0 11
Eupistella grubei 0 0 4 0 0 0 0 0 0 16 Eupistella sp. 1 0 0 0 0 0 0 0 0 0 2
Eusamythella sp. 1 0 0 0 0 0 0 0 0 0 7 Exogone heterosetosa 0 0 0 0 0 0 0 0 0 2
Exogone minuscula 0 0 1 0 0 18 0 1 0 28 Exogone sp. 2 BH 0 0 0 0 0 0 0 0 0 1
APPENDIX- TABLES
227
species 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1
Exogone sp. 5 0 0 0 0 0 0 0 0 0 0 Exogone sp. 6 0 0 0 0 0 0 0 0 0 0 Exogone sp. 7 0 0 0 0 0 0 0 0 0 0
Glycera kerguelensis 1 2 3 0 0 0 0 17 23 31 Glyphanostomum scotiarum 0 3 0 0 0 0 0 1 0 0
Goniada maculata 0 0 0 0 0 0 0 0 0 0 Grubianella antarctica 0 2 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 0 0 0 0 0 0 0 Gyptis incompta 0 0 0 0 0 0 0 0 0 0
Hauchiella tribullata 0 1 0 0 0 0 0 0 0 0 Hesionidae sp. 1 0 0 0 3 0 0 0 0 0 0
Hyboscolex equatorialis 0 0 0 0 0 0 0 0 1 0 Kefersteinia fauveli 0 0 0 0 0 0 0 0 35 0 Kesun abyssorum 0 0 86 1 0 0 3 0 8 0
Laphania cf. boecki 0 0 2 0 0 0 0 0 0 0 Leaena antarctica 0 0 0 0 0 0 0 0 0 0 Leaena arenilega 0 0 0 0 0 0 0 0 0 0 Leaena cf. collaris 0 0 0 0 0 0 0 0 0 0
Leaena collaris 0 0 0 0 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 0 0 0 0 0 0 0
Leaena sp. 4 0 0 0 0 0 0 0 0 0 0 Leaena wandelensis 0 0 0 0 0 0 0 0 0 0
Lysilla sp. 1 BH 0 0 1 0 0 0 0 0 0 0 Melinna cristata 0 0 0 0 0 0 0 0 0 0
Micronephtys sp. 1 0 0 1 0 0 0 0 0 0 0 Micropodarke cylindripalpata sp.n. 0 9 4 2 0 0 0 0 2 0
Mugga sp. 1BH 0 0 0 0 0 0 0 0 1 0 Muggoides cf. cinctus 0 0 0 0 0 0 0 0 0 0 Muggoides sp. 1BH 0 1 0 0 0 0 0 0 0 0
Neosabellides elongatus 0 0 2 0 0 0 1 0 0 0 Nereis eugeniae 0 1 1 0 1 0 0 0 1 0
Octobranchus antarcticus 0 0 1 0 0 0 0 0 1 0 Oligobregma blakei 0 0 1 0 0 0 0 0 0 0 Oligobregma collare 0 4 3 8 0 0 2 0 0 0
Oligobregma hartmanae 0 0 0 0 0 0 0 0 0 0 Oligobregma notiale 0 7 0 0 0 0 0 0 0 0
Oligobregma pseudocollare 0 0 20 1 0 0 0 0 0 8 Oligobregma quadrispinosa 0 5 0 8 0 0 2 0 1 0
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 0 0 0 2 0 0 0 0 0 0
Ophelina breviata 0 1 2 16 0 0 0 0 0 0 Ophelina gymnopyge 0 3 0 15 0 0 0 0 0 0 Ophelina nematoides 1 14 7 2 0 0 0 0 5 0
Ophelina robusta sp.n. 0 0 0 1 0 0 0 0 0 0 Ophelina scaphigera 0 0 0 0 0 0 0 0 0 0
Ophelina setigera 0 0 1 1 0 0 0 0 0 0 Ophelina sp. 3 0 0 0 2 0 0 0 0 0 0 Ophelina sp. 4 0 1 0 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 0 0 0 0 0 0 0 0
Ophiodromus calligocervix sp.n. 0 28 0 3 1 0 6 1 1 1 Ophiodromus comatus 0 0 0 0 0 0 0 0 0 0
Parasyllidea delicata sp.n. 0 0 0 1 0 0 0 0 0 0 Phalacrostemma elegans 0 0 1 0 0 0 0 0 0 0
APPENDIX- TABLES
228
species 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13
Exogone sp. 5 0 0 0 5 0 0 0 0 0 0 Exogone sp. 6 0 0 0 0 0 0 0 0 0 0 Exogone sp. 7 0 0 0 0 0 0 0 0 0 0
Glycera kerguelensis 4 2 14 43 47 45 2 29 19 8 Glyphanostomum scotiarum 0 0 0 3 1 0 0 0 0 0
Goniada maculata 1 0 0 0 0 0 0 0 0 0 Grubianella antarctica 2 0 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 1 0 0 0 0 0 0 Gyptis incompta 0 0 0 48 0 0 0 0 0 0
Hauchiella tribullata 0 0 0 2 0 0 0 0 0 0 Hesionidae sp. 1 0 0 0 0 0 0 0 0 0 0
Hyboscolex equatorialis 0 0 0 0 0 0 0 0 0 0 Kefersteinia fauveli 3 0 6 70 0 0 0 0 0 0 Kesun abyssorum 0 1 14 0 24 8 1 0 1 3
Laphania cf. boecki 0 0 0 0 0 0 0 0 0 0 Leaena antarctica 0 0 0 0 0 0 1 0 0 1 Leaena arenilega 0 0 0 5 0 0 0 0 0 0 Leaena cf. collaris 1 0 0 3 0 0 0 0 0 0
Leaena collaris 0 0 0 2 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 3 0 0 0 0 0 0
Leaena sp. 4 0 0 0 1 0 0 0 0 0 0 Leaena wandelensis 0 1 0 0 0 0 0 0 0 0
Lysilla sp. 1 BH 0 0 0 0 0 0 0 0 0 0 Melinna cristata 0 0 0 0 0 4 0 0 0 0
Micronephtys sp. 1 0 0 0 0 0 0 0 0 0 0 Micropodarke cylindripalpata sp.n. 1 2 5 0 0 5 4 6 1 0
Mugga sp. 1BH 3 0 0 0 0 0 0 0 0 0 Muggoides cf. cinctus 0 0 0 0 0 0 11 0 0 0 Muggoides sp. 1BH 2 0 0 0 0 0 0 0 0 4
Neosabellides elongatus 0 1 0 4 2 1 1 0 0 0 Nereis eugeniae 0 0 0 0 0 0 0 0 0 0
Octobranchus antarcticus 0 0 0 19 0 4 0 0 0 0 Oligobregma blakei 0 0 0 0 0 0 0 0 0 0 Oligobregma collare 0 0 1 1 7 1 0 0 0 0
Oligobregma hartmanae 0 0 3 0 0 0 0 0 0 0 Oligobregma notiale 0 0 0 0 0 0 0 0 0 0
Oligobregma pseudocollare 0 0 0 0 0 2 0 0 0 0 Oligobregma quadrispinosa 0 1 6 0 0 0 0 1 2 0
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 4 0 0 0 10 0 4 0 0 0
Ophelina breviata 1 0 0 16 1 0 0 0 0 1 Ophelina gymnopyge 2 0 1 0 0 0 3 0 0 0 Ophelina nematoides 2 0 0 18 0 1 0 0 0 0
Ophelina robusta sp.n. 0 0 0 0 10 2 0 0 0 0 Ophelina scaphigera 4 0 1 0 0 0 0 0 0 0
Ophelina setigera 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 3 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 4 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 0 0 0 0 0 0 0 0
Ophiodromus calligocervix sp.n. 2 3 2 0 14 12 0 8 4 1 Ophiodromus comatus 0 0 0 25 0 7 3 0 0 0
Parasyllidea delicata sp.n. 2 3 0 0 0 4 0 0 0 0 Phalacrostemma elegans 0 0 0 0 0 1 0 0 0 0
APPENDIX- TABLES
229
species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total
Exogone sp. 5 0 0 0 0 0 0 0 0 0 5 Exogone sp. 6 0 0 16 0 0 0 0 0 0 16 Exogone sp. 7 0 0 42 0 0 0 0 4 0 46
Glycera kerguelensis 29 0 19 0 6 0 0 4 5 353 Glyphanostomum scotiarum 0 0 0 0 0 0 0 0 0 8
Goniada maculata 0 0 0 0 0 0 0 0 0 1 Grubianella antarctica 0 0 0 0 0 0 0 1 0 5
Grubianella sp. 1 0 0 0 0 0 0 0 0 0 1 Gyptis incompta 0 0 736 0 0 0 0 0 0 784
Hauchiella tribullata 0 0 0 0 0 0 0 0 0 3 Hesionidae sp. 1 0 0 0 0 0 0 0 0 0 3
Hyboscolex equatorialis 0 0 0 0 0 0 0 0 0 1 Kefersteinia fauveli 0 0 203 0 0 11 0 0 0 328 Kesun abyssorum 4 14 2 4 26 0 0 54 11 265
Laphania cf. boecki 0 0 0 0 0 0 0 0 0 2 Leaena antarctica 0 0 2 0 0 0 0 7 0 11 Leaena arenilega 0 0 0 0 0 0 0 1 0 6 Leaena cf. collaris 0 0 0 0 0 0 0 0 0 4
Leaena collaris 0 0 64 0 0 0 0 0 0 66 Leaena pseudobranchia 0 0 0 0 0 0 0 0 0 3
Leaena sp. 4 0 0 0 0 0 0 0 0 0 1 Leaena wandelensis 0 0 0 0 0 0 0 0 3 4
Lysilla sp. 1 BH 0 0 0 0 0 0 0 0 0 1 Melinna cristata 0 0 0 0 4 0 0 0 0 8
Micronephtys sp. 1 0 0 0 0 0 0 0 0 0 1 Micropodarke cylindripalpata sp.n. 2 1 0 0 3 0 0 0 2 49
Mugga sp. 1BH 0 0 0 0 0 0 0 0 1 5 Muggoides cf. cinctus 0 0 0 0 0 0 0 0 0 11 Muggoides sp. 1BH 0 0 0 0 0 0 0 0 0 7
Neosabellides elongatus 0 0 1 0 0 0 0 2 0 15 Nereis eugeniae 0 0 0 0 0 0 0 0 0 4
Octobranchus antarcticus 0 0 119 0 0 0 0 1 0 145 Oligobregma blakei 0 1 2 0 0 0 0 0 0 4 Oligobregma collare 1 11 3 1 1 1 0 10 1 56
Oligobregma hartmanae 0 0 2 0 1 0 0 0 0 6 Oligobregma notiale 0 0 0 0 0 0 0 0 0 7
Oligobregma pseudocollare 0 1 8 0 1 0 0 2 0 43 Oligobregma quadrispinosa 1 1 1 0 0 1 0 10 0 40
Oligobregma sp. 1 0 0 0 0 0 1 0 0 0 1 Ophelina ammotrypanella sp.n. 4 7 0 0 0 0 0 0 8 39
Ophelina breviata 1 6 19 0 1 0 0 1 1 67 Ophelina gymnopyge 3 2 14 0 0 0 0 0 4 47 Ophelina nematoides 0 0 0 0 0 0 0 1 0 51
Ophelina robusta sp.n. 0 4 0 0 5 2 0 7 0 31 Ophelina scaphigera 0 0 2 0 0 0 0 0 0 7
Ophelina setigera 0 1 0 0 0 0 0 0 0 3 Ophelina sp. 3 0 0 0 0 0 0 0 0 0 2 Ophelina sp. 4 0 0 0 0 0 0 0 0 0 1 Ophelina sp. 5 0 0 10 0 2 0 0 0 5 17
Ophiodromus calligocervix sp.n. 3 1 0 0 0 0 0 7 2 100 Ophiodromus comatus 0 4 43 0 3 17 0 1 0 103
Parasyllidea delicata sp.n. 2 0 0 3 7 0 0 1 0 23 Phalacrostemma elegans 0 0 0 0 0 0 0 0 0 2
APPENDIX- TABLES
230
species 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1
Phisidia rubrolineata 0 0 0 0 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 0 0
Phyllocomus crocea 0 0 0 0 0 0 0 0 0 0 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 0 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 1 0 0 1 0 0 1 0
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 1 0 Pista corrientis 0 0 0 0 0 0 0 0 0 0 Pista cristata 0 0 0 0 0 0 0 0 0 0
Pista spinifera 0 0 0 0 0 0 0 0 0 0 Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 0 0
Polycirrus insignis 0 0 1 0 0 0 0 0 0 1 Polycirrus sp. 1 0 0 0 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 0 0 0 0 0 0 0
Proceraea cf. mclearanus 0 0 0 0 0 0 0 0 0 0 Proclea cf. graffii 0 0 1 0 0 0 0 0 0 0
Pseudoscalibregma bransfieldium 0 8 5 0 6 0 0 0 0 2 Pseudoscalibregma papilia sp.n. 0 3 2 0 0 0 0 0 3 0
Pseudoscalibregma ursapium 0 1 1 1 0 0 0 0 0 0 Samytha sp. 1 0 0 0 0 0 0 0 0 0 0
Scalibregma inflatum 0 3 2 0 0 0 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 0 0
Sosanopsis kerguelensis 0 0 0 3 0 0 2 0 0 0 Sosanopsis sp. 1 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis distincta sp.n. 0 0 6 0 0 0 0 0 0 0 Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis parva 2 82 333 0 0 0 0 0 21 11 Sphaerodoropsis polypapillata 0 1 0 2 0 0 0 0 0 0 Sphaerodoropsis simplex sp.n. 0 0 22 0 0 0 0 0 1 0
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 0 0 0 0 0 0 1 0
Sphaerosyllis lateropapillata uteae 0 0 0 0 0 0 0 0 12 1 Streblosoma variouncinatum 0 0 0 0 0 0 0 0 2 0
Syllides articulosus 0 0 0 0 0 0 0 0 0 0 Terebellidae sp. 1 0 0 0 1 0 0 0 0 0 0 Terebellidae sp. 2 0 0 1 0 0 0 0 0 0 0 Terebellidae sp. 3 0 0 0 0 0 0 0 0 0 0 Terebellidae sp. 4 0 0 1 0 0 0 0 0 0 0
Terebellides stroemii 0 1 2 1 0 0 0 0 0 0 Thelepides koehleri 0 0 0 0 0 0 0 0 0 0 Thelepides venustus 0 0 0 0 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 7 0 0 0 0 0 0 0 Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 0 0 1 0 0 0 0 0 0 6
Trichobranchidae sp. 1 0 0 1 0 0 0 0 0 0 0 Trichobranchidae sp. 2 0 0 0 0 0 0 0 0 1 0 Typosyllis cf. hyalina 0 0 0 0 0 0 0 0 0 8 Typosyllis variegata 0 0 0 0 0 0 0 0 0 4
APPENDIX- TABLES
231
species 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13
Phisidia rubrolineata 0 0 0 0 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 0 0
Phyllocomus crocea 0 0 2 2 0 0 2 0 0 0 Pionosyllis cf. maxima 0 0 0 1 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 7 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 0 14 0 0 0 0 0 0
Pionosyllis sp. 1 0 0 0 0 1 0 0 0 0 0 Pista corrientis 0 0 0 1 0 0 0 0 0 0 Pista cristata 0 0 0 2 0 0 0 0 0 0
Pista spinifera 0 0 0 5 0 0 0 0 0 0 Polycirrus cf. antarcticus 0 0 0 1 0 0 0 0 0 0
Polycirrus insignis 0 0 0 13 3 0 0 0 0 0 Polycirrus sp. 1 0 0 1 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 1 0 0 0 0 0 0
Proceraea cf. mclearanus 0 0 0 1 0 0 0 0 0 0 Proclea cf. graffii 0 0 0 0 0 1 0 0 0 0
Pseudoscalibregma bransfieldium 0 0 0 2 0 1 0 1 0 0 Pseudoscalibregma papilia sp.n. 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma ursapium 0 0 0 0 0 3 0 0 0 0 Samytha sp. 1 0 0 0 1 0 0 0 0 0 0
Scalibregma inflatum 0 0 0 0 0 0 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 1 0 0 0 0
Sosanopsis kerguelensis 4 6 4 3 1 2 0 1 3 1 Sosanopsis sp. 1 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis distincta sp.n. 0 0 0 0 0 0 0 0 0 0 Sphaerodoropsis maculata sp.n. 0 0 0 23 0 0 0 0 0 0
Sphaerodoropsis parva 1 1 6 87 3 6 28 0 0 0 Sphaerodoropsis polypapillata 0 0 0 0 0 4 0 0 0 0 Sphaerodoropsis simplex sp.n. 0 0 0 3 0 1 0 0 0 0
Sphaerosyllis antarctica 0 0 0 1 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 0 12 0 0 0 0 0 0
Sphaerosyllis lateropapillata uteae 0 0 0 156 0 0 0 0 0 0 Streblosoma variouncinatum 0 0 0 0 0 0 0 0 0 0
Syllides articulosus 0 0 0 1 0 0 0 0 0 0 Terebellidae sp. 1 0 0 0 0 0 0 0 0 0 0 Terebellidae sp. 2 0 0 0 0 0 0 0 0 0 0 Terebellidae sp. 3 0 1 0 0 0 0 0 0 0 0 Terebellidae sp. 4 0 0 0 0 0 0 0 0 0 0
Terebellides stroemii 1 0 1 6 12 5 2 0 2 1 Thelepides koehleri 0 0 0 2 0 0 0 0 0 0 Thelepides venustus 0 0 0 3 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 0 1 0 0 0 0 0 0 Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 0 0 1 6 4 0 0 0 0 0
Trichobranchidae sp. 1 0 0 0 0 0 0 0 0 0 0 Trichobranchidae sp. 2 0 0 0 0 0 0 0 0 0 0 Typosyllis cf. hyalina 0 0 0 1 0 0 0 0 0 0 Typosyllis variegata 0 0 0 0 0 0 0 0 0 0
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species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total
Phisidia rubrolineata 0 0 266 0 0 0 0 0 0 266 Phisidia sp. 1BH 0 0 0 0 1 0 0 0 0 1
Phyllocomus crocea 0 0 0 0 0 2 0 0 0 8 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 0 1
Pionosyllis comosa 0 0 0 0 0 0 0 0 0 7 Pionosyllis epipharynx 0 0 24 0 1 0 0 1 2 45
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 0 2 Pista corrientis 0 0 0 0 0 0 0 0 0 1 Pista cristata 0 0 0 0 0 0 0 0 0 2
Pista spinifera 0 0 0 0 0 0 0 0 0 5 Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 0 1
Polycirrus insignis 0 0 241 2 1 0 0 0 1 263 Polycirrus sp. 1 0 0 0 0 0 0 0 0 0 1 Polycirrus sp. 2 0 0 0 0 0 0 0 0 0 1
Proceraea cf. mclearanus 0 0 0 0 0 0 0 0 0 1 Proclea cf. graffii 0 0 0 0 0 0 0 0 0 2
Pseudoscalibregma bransfieldium 0 0 2 0 1 1 0 1 0 30 Pseudoscalibregma papilia sp.n. 0 1 0 1 5 0 0 1 0 16
Pseudoscalibregma ursapium 0 0 5 0 0 1 0 0 0 12 Samytha sp. 1 0 0 0 0 0 0 0 0 0 1
Scalibregma inflatum 0 0 1 0 2 0 0 0 0 8 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 0 1
Sosanopsis kerguelensis 2 3 8 0 2 0 0 6 1 52 Sosanopsis sp. 1 0 0 0 0 5 0 0 2 0 7
Sphaerodoropsis distincta sp.n. 0 0 0 0 0 0 0 0 0 6 Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 0 23
Sphaerodoropsis parva 0 0 189 0 2 11 1 3 2 789 Sphaerodoropsis polypapillata 0 5 0 0 0 0 0 2 0 14 Sphaerodoropsis simplex sp.n. 0 0 0 0 1 0 0 0 0 28
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 0 1 Sphaerosyllis joinvillensis 0 0 285 0 0 0 0 0 0 298
Sphaerosyllis lateropapillata uteae 0 0 217 0 5 4 0 38 0 433 Streblosoma variouncinatum 0 0 0 0 0 0 0 0 0 2
Syllides articulosus 0 0 317 0 0 0 0 0 0 318 Terebellidae sp. 1 0 0 0 0 0 0 0 0 0 1 Terebellidae sp. 2 0 0 0 0 0 0 0 0 0 1 Terebellidae sp. 3 0 0 0 0 0 0 0 0 0 1 Terebellidae sp. 4 0 0 0 0 0 0 0 0 0 1
Terebellides stroemii 3 15 1 6 4 0 0 0 0 63 Thelepides koehleri 0 0 0 0 0 0 0 0 0 2 Thelepides venustus 0 0 0 0 0 0 0 0 0 3 Thelepodinae sp. 1 0 0 0 0 0 0 0 0 0 8 Travisia cf. lithophila 0 0 0 0 0 0 0 0 1 1 Travisia kerguelensis 0 0 2 0 3 0 0 1 9 33
Trichobranchidae sp. 1 0 0 0 0 0 0 0 0 0 1 Trichobranchidae sp. 2 0 0 0 0 0 0 0 0 0 1 Typosyllis cf. hyalina 0 0 5 0 0 0 0 0 0 14 Typosyllis variegata 0 0 60 0 0 0 0 0 0 64
total 6669
APPENDIX- TABLES
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App.-Tab. 7: Standardized species assemblage matrix: ANDEEPIII (1000 m trawling distance)
species 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13aff. Hesionides 1,25078 1,02634 0,34746 0 0 0 0 0,2867 0 0
Aglaophamus paramalmgreni 0 0 4,86449 0 0,42088 0 0 0 0 0 Aglaophamus trissophyllus 0 0 0 102,672 7,57576 1,12486 0 0 0 0
Amage sculpta 0 2,73691 0 1,40647 0 1,68729 0 0 0 0 Ammotrypanella arctica 1,87617 0 1,38985 0 0,42088 0 0 0 0 0,6092
Ammotrypanella cirrosa sp.n. 1,87617 0 0,69493 0 0,84175 0 0 0 0 1,2184 Ammotrypanella mcintoshi sp.n. 0,93809 0,34211 0,34746 2,81294 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 0,3127 0 0 0 0,42088 0 0,34072 0 0 0
Ampharete kerguelensis 4,06504 0 0,34746 0 2,94613 0 2,04429 0 0 0 Ampharetidae sp. 2 0 0,34211 0 0 0,84175 0 0 0 0 0 Ampharetidae sp. 4 0 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 5 0 0 0 0 0,84175 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 0,56243 0,34072 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 0,68143 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0 0 0 0 0 Amphicteis gunneri 0 0 0 0 0,42088 0,56243 0 0 0 0 Amphicteis juvenile 0 0 0 0 0,42088 0 0 0 0 0
Amphicteis sp. 1 0 0 0 0 6,31313 0 0,34072 0 0 0 Amphicteis sp. 2 0 0 0 0 2,94613 0 0 0 0 0 Amphicteis sp. 3 0 0 0 7,03235 0 0,56243 0 0 0 0
Amphiduros serratus sp.n. 0 0 0 0 0 0 0 0 0 0 Amythas membranifera 0 0 0 0 0 0 0 0 0 0 Anobothrella antarctica 0 0 0 4,21941 0 0 0,34072 0 0 0
Anobothrella sp. 1 0 0 0 0 0 0 0,34072 0 0 0 Anobothrus gracilis 0 0 0 0 0 0 0,68143 0 0 0,3046
Anobothrus pseudoampharete sp.n. 0 0,34211 0 146,273 0 0 1,02215 0 0 0 Autolytus gibber 0 0 0 1,40647 0 0 0 0 0 0
Axiokebuita minuta 0 0 0 11,2518 0 3,93701 0,34072 0 0 0 Bathyglycinde sp. 1 BH 0,3127 0 0 0 0 0 0 0 0 0
Braniella palpata 0 0,34211 0,34746 4,21941 0 1,12486 0,34072 0 0 0 Ceratocephale sp. 1BH 0 0 0 0 0,42088 0 0 0,2867 0 0
cf. Amphisamytha 0 0 0 1,40647 0 0 0 0 0 0 cf. Neosabellides 0 0 0 1,40647 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0 0 cf. Terebellides sp. 1BH 0 0 0 0 0 0,56243 0 0 0 0 cf. Thelepides venustus 0 0 0 1,40647 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 1,40647 0 0 0 0 0 0 Clavodorum antarcticum 0 0 0 0 0 0 0 0 0 0
Ephesiella antarctica 0 0 0 0 0 0 0 0 0 0 Ephesiella hartmanae sp.n. 0,3127 0 0 2,81294 0 0,56243 0 0 0 0
Eupistella grubei 0 0 0 12,6582 0 1,68729 0 0 0 0 Eupistella sp. 1 0,62539 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 1,40647 0 0 0,68143 0 0 0 Exogone heterosetosa 0 0 0 1,40647 0 0 0 0 0 0
Exogone minuscula 0 0 0 11,2518 0 0 0 0 0 0 Exogone sp. 5 0 0 0 7,03235 0 0 0 0 0 0 Exogone sp. 6 0 0 0 0 0 0 0 0 0 0 Exogone sp. 7 0 0 0 0 0 0 0 0 0 0
Glycera kerguelensis 1,25078 0,68423 4,86449 60,4782 19,7811 25,3093 0,68143 8,31422 5,46605 2,4368 Glyphanostomum scotiarum 0 0 0 4,21941 0,42088 0 0 0 0 0
Goniada maculata 0,3127 0 0 0 0 0 0 0 0 0 Grubianella antarctica 0,62539 0 0 0 0 0 0 0 0 0
Grubianella sp. 1 0 0 0 1,40647 0 0 0 0 0 0 Gyptis incompta 0 0 0 67,5105 0 0 0 0 0 0
Hauchiella tribullata 0 0 0 2,81294 0 0 0 0 0 0 Kefersteinia fauveli 0,93809 0 2,08478 98,4529 0 0 0 0 0 0 Kesun abyssorum 0 0,34211 4,86449 0 10,101 4,49944 0,34072 0 0,28769 0,9138 Leaena antarctica 0 0 0 0 0 0 0,34072 0 0 0,3046 Leaena arenilega 0 0 0 7,03235 0 0 0 0 0 0 Leaena cf. collaris 0,3127 0 0 4,21941 0 0 0 0 0 0
Leaena collaris 0 0 0 2,81294 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 4,21941 0 0 0 0 0 0
Leaena sp. 4 0 0 0 1,40647 0 0 0 0 0 0 Leaena wandelensis 0 0,34211 0 0 0 0 0 0 0 0
APPENDIX- TABLES
234
species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 aff. Hesionides 0 0 1,71821 0 0 0 0 0 0
Aglaophamus paramalmgreni 0,68871 0 0 0 0,63816 1,44613 0,47326 7,67656 0 Aglaophamus trissophyllus 0,34435 0 0 0 1,27632 0 0 0 1,58416
Amage sculpta 0 0,51414 1,71821 0 0 0 0 0 0 Ammotrypanella arctica 4,82094 11,3111 0 0 0 0 0 1,02354 1,18812
Ammotrypanella cirrosa sp.n. 0,34435 20,5656 0 0 0 0 0 2,04708 0,39604 Ammotrypanella mcintoshi sp.n. 1,03306 0,51414 0 0 0 0 0 0 0 Ammotrypanella princessa sp.n. 0 0,51414 0 0 0 0 0 0,51177 0
Ampharete kerguelensis 0 0 0,85911 0 0 0 0 0 0 Ampharetidae sp. 2 0 1,02828 0 0,44425 0 0 0 2,55885 0 Ampharetidae sp. 4 0 0 0 0 0 0,72307 0 0 0 Ampharetidae sp. 5 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 6 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 7 0 0 0 0 0 0 0 0 0 Ampharetidae sp. 8 0 0 0 0 0 0,72307 0 0 0 Amphicteis gunneri 0 0,51414 0 0 0 0 0 0 0 Amphicteis juvenile 0 0 0 0 0 0 0 0 0
Amphicteis sp. 1 0 16,4524 0,85911 0 5,1053 0 0 1,53531 0 Amphicteis sp. 2 0 0 0 0 0 0 0 0 0 Amphicteis sp. 3 0 0 0 0,44425 1,27632 0 0 0 0,39604
Amphiduros serratus sp.n. 0 0 0 0 0 0 0,94652 0 0 Amythas membranifera 0 0 0,85911 1,33274 0 0 0 0 0 Anobothrella antarctica 0 0 0 0 0,63816 0 0 0 0
Anobothrella sp. 1 0 0 0 0 0 0 0 0 0 Anobothrus gracilis 0 0 6,01375 0 0 0 0 0 0,79208
Anobothrus pseudoampharete sp.n. 0 4,62725 3,43643 0,44425 7,65795 0 0 0 0,39604 Autolytus gibber 0 0 0 0 0 0 0 0 0
Axiokebuita minuta 0 0 12,8866 0 1,27632 0 0 0 0 Bathyglycinde sp. 1 BH 0 0 0 0 0 0 0 0 0
Braniella palpata 0 1,02828 27,4914 0 0 0 0 0 0 Ceratocephale sp. 1BH 0,34435 0 0 0 0 0 0 0,51177 0,39604
cf. Amphisamytha 0 0 0 0 0 0 0 0 0 cf. Neosabellides 0 0 0 0 0 0 0 0 0
cf. Nicon 0 0 0 0 0 0 0 0 0,39604 cf. Terebellides sp. 1BH 0 0 0 0 0 0 0 0 0 cf. Thelepides venustus 0 0 0 0 0 0 0 0 0 cf. Thelepus cincinnatus 0 0 0 0 0 0 0 0 0 Clavodorum antarcticum 0 0 0 0,44425 0,63816 0 0 0 0
Ephesiella antarctica 0 0 0 0,44425 0 0 0 1,02354 0 Ephesiella hartmanae sp.n. 0 0 2,57732 0 0,63816 0 0 0 0
Eupistella grubei 0 0 3,43643 0 0 0 0 0 0 Eupistella sp. 1 0 0 0 0 0 0 0 0 0
Eusamythella sp. 1 0 0 0 0 0 0 0 0 0 Exogone heterosetosa 0 0 0 0 0 0 0 0 0
Exogone minuscula 0 0 0,85911 0 0 13,0152 0 0,51177 0 Exogone sp. 5 0 0 0 0 0 0 0 0 0 Exogone sp. 6 0 0 13,7457 0 0 0 0 0 0 Exogone sp. 7 0 0 36,0825 0 0 0 0 2,04708 0
Glycera kerguelensis 9,98623 0 16,323 0 3,82897 0 0 2,04708 1,9802 Glyphanostomum scotiarum 0 0 0 0 0 0 0 0 0
Goniada maculata 0 0 0 0 0 0 0 0 0 Grubianella antarctica 0 0 0 0 0 0 0 0,51177 0
Grubianella sp. 1 0 0 0 0 0 0 0 0 0 Gyptis incompta 0 0 632,302 0 0 0 0 0 0
Hauchiella tribullata 0 0 0 0 0 0 0 0 0 Kefersteinia fauveli 0 0 174,399 0 0 7,95372 0 0 0 Kesun abyssorum 1,37741 7,19794 1,71821 1,77699 16,5922 0 0 27,6356 4,35644 Leaena antarctica 0 0 1,71821 0 0 0 0 3,5824 0 Leaena arenilega 0 0 0 0 0 0 0 0,51177 0 Leaena cf. collaris 0 0 0 0 0 0 0 0 0
Leaena collaris 0 0 54,9828 0 0 0 0 0 0 Leaena pseudobranchia 0 0 0 0 0 0 0 0 0
Leaena sp. 4 0 0 0 0 0 0 0 0 0 Leaena wandelensis 0 0 0 0 0 0 0 0 1,18812
APPENDIX- TABLES
235
species 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13Melinna cristata 0 0 0 0 0 2,24972 0 0 0 0
Micropodarke cylindripalpata sp.n. 0,3127 0,68423 1,73732 0 0 2,81215 1,36286 1,72018 0,28769 0 Mugga sp. 1BH 0,93809 0 0 0 0 0 0 0 0 0
Muggoides cf. cinctus 0 0 0 0 0 0 3,74787 0 0 0 Muggoides sp. 1BH 0,62539 0 0 0 0 0 0 0 0 1,2184
Neosabellides elongatus 0 0,34211 0 5,62588 0,84175 0,56243 0,34072 0 0 0 Octobranchus antarcticus 0 0 0 26,7229 0 2,24972 0 0 0 0
Oligobregma blakei 0 0 0 0 0 0 0 0 0 0 Oligobregma collare 0 0 0,34746 1,40647 2,94613 0,56243 0 0 0 0
Oligobregma hartmanae 0 0 1,04239 0 0 0 0 0 0 0 Oligobregma pseudocollare 0 0 0 0 0 1,12486 0 0 0 0 Oligobregma quadrispinosa 0 0,34211 2,08478 0 0 0 0 0,2867 0,57537 0
Oligobregma sp. 1 0 0 0 0 0 0 0 0 0 0 Ophelina ammotrypanella sp.n. 1,25078 0 0 0 4,20875 0 1,36286 0 0 0
Ophelina breviata 0,3127 0 0 22,5035 0,42088 0 0 0 0 0,3046 Ophelina gymnopyge 0,62539 0 0,34746 0 0 0 1,02215 0 0 0 Ophelina nematoides 0,62539 0 0 25,3165 0 0,56243 0 0 0 0
Ophelina robusta sp.n. 0 0 0 0 4,20875 1,12486 0 0 0 0 Ophelina scaphigera 1,25078 0 0,34746 0 0 0 0 0 0 0
Ophelina setigera 0 0 0 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 0 0 0 0 0 0 0 0
Ophiodromus calligocervix sp.n. 0,62539 1,02634 0,69493 0 5,89226 6,74916 0 2,29358 1,15075 0,3046 Ophiodromus comatus 0 0 0 35,1617 0 3,93701 1,02215 0 0 0
Parasyllidea delicata sp.n. 0,62539 1,02634 0 0 0 2,24972 0 0 0 0 Phalacrostemma elegans 0 0 0 0 0 0,56243 0 0 0 0
Phisidia rubrolineata 0 0 0 0 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0 0 0 0 0 0
Phyllocomus crocea 0 0 0,69493 2,81294 0 0 0,68143 0 0 0 Pionosyllis cf. maxima 0 0 0 1,40647 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 9,84529 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 0 19,6906 0 0 0 0 0 0
Pionosyllis sp. 1 0 0 0 0 0,42088 0 0 0 0 0 Pista corrientis 0 0 0 1,40647 0 0 0 0 0 0 Pista cristata 0 0 0 2,81294 0 0 0 0 0 0
Pista spinifera 0 0 0 7,03235 0 0 0 0 0 0 Polycirrus cf. antarcticus 0 0 0 1,40647 0 0 0 0 0 0
Polycirrus insignis 0 0 0 18,2841 1,26263 0 0 0 0 0 Polycirrus sp. 1 0 0 0,34746 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 1,40647 0 0 0 0 0 0
Proceaea cf. mclearanus 0 0 0 1,40647 0 0 0 0 0 0 Proclea cf. graffii 0 0 0 0 0 0,56243 0 0 0 0
Pseudoscalibregma bransfieldium 0 0 0 2,81294 0 0,56243 0 0,2867 0 0 Pseudoscalibregma papilia sp.n. 0 0 0 0 0 0 0 0 0 0
Pseudoscalibregma ursapium 0 0 0 0 0 1,68729 0 0 0 0 Samytha sp. 1 0 0 0 1,40647 0 0 0 0 0 0
Scalibregma inflatum 0 0 0 0 0 0 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 0,56243 0 0 0 0
Sosanopsis kerguelensis 1,25078 2,05269 1,38985 4,21941 0,42088 1,12486 0 0,2867 0,86306 0,3046 Sosanopsis sp. 1 0 0 0 0 0 0 0 0 0 0
Sphaerodoropsis maculata sp.n. 0 0 0 32,3488 0 0 0 0 0 0 Sphaerodoropsis parva 0,3127 0,34211 2,08478 122,363 1,26263 3,37458 9,54003 0 0 0
Sphaerodoropsis polypapillata 0 0 0 0 0 2,24972 0 0 0 0 Sphaerodoropsis simplex sp.n. 0 0 0 4,21941 0 0,56243 0 0 0 0
Sphaerosyllis antarctica 0 0 0 1,40647 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 0 16,8776 0 0 0 0 0 0
Sphaerosyllis lateropapillata uteae 0 0 0 219,409 0 0 0 0 0 0 Syllides articulosus 0 0 0 1,40647 0 0 0 0 0 0 Terebellidae sp. 3 0 0,34211 0 0 0 0 0 0 0 0
Terebellides stroemii 0,3127 0 0,34746 8,43882 5,05051 2,81215 0,68143 0 0,57537 0,3046 Thelepides koehleri 0 0 0 2,81294 0 0 0 0 0 0 Thelepides venustus 0 0 0 4,21941 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 0 1,40647 0 0 0 0 0 0 Travisia cf. lithophila 0 0 0 0 0 0 0 0 0 0 Travisia kerguelensis 0 0 0,34746 8,43882 1,6835 0 0 0 0 0 Typosyllis cf. hyalina 0 0 0 1,40647 0 0 0 0 0 0 Typosyllis variegata 0 0 0 0 0 0 0 0 0 0
APPENDIX- TABLES
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species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 Melinna cristata 0 0 0 0 2,55265 0 0 0 0
Micropodarke cylindripalpata sp.n. 0,68871 0,51414 0 0 1,91449 0 0 0 0,79208 Mugga sp. 1BH 0 0 0 0 0 0 0 0 0,39604
Muggoides cf. cinctus 0 0 0 0 0 0 0 0 0 Muggoides sp. 1BH 0 0 0 0 0 0 0 0 0
Neosabellides elongatus 0 0 0,85911 0 0 0 0 1,02354 0 Octobranchus antarcticus 0 0 102,234 0 0 0 0 0,51177 0
Oligobregma blakei 0 0,51414 1,71821 0 0 0 0 0 0 Oligobregma collare 0,34435 5,65553 2,57732 0,44425 0,63816 0,72307 0 5,11771 0,39604
Oligobregma hartmanae 0 0 1,71821 0 0,63816 0 0 0 0 Oligobregma pseudocollare 0 0,51414 6,87285 0 0,63816 0 0 1,02354 0 Oligobregma quadrispinosa 0,34435 0,51414 0,85911 0 0 0,72307 0 5,11771 0
Oligobregma sp. 1 0 0 0 0 0 0,72307 0 0 0 Ophelina ammotrypanella sp.n. 1,37741 3,59897 0 0 0 0 0 0 3,16832
Ophelina breviata 0,34435 3,08483 16,323 0 0,63816 0 0 0,51177 0,39604 Ophelina gymnopyge 1,03306 1,02828 12,0275 0 0 0 0 0 1,58416 Ophelina nematoides 0 0 0 0 0 0 0 0,51177 0
Ophelina robusta sp.n. 0 2,05656 0 0 3,19081 1,44613 0 3,5824 0 Ophelina scaphigera 0 0 1,71821 0 0 0 0 0 0
Ophelina setigera 0 0,51414 0 0 0 0 0 0 0 Ophelina sp. 5 0 0 8,59107 0 1,27632 0 0 0 1,9802
Ophiodromus calligocervix sp.n. 1,03306 0,51414 0 0 0 0 0 3,5824 0,79208 Ophiodromus comatus 0 2,05656 36,9416 0 1,91449 12,2921 0 0,51177 0
Parasyllidea delicata sp.n. 0,68871 0 0 1,33274 4,46713 0 0 0,51177 0 Phalacrostemma elegans 0 0 0 0 0 0 0 0 0
Phisidia rubrolineata 0 0 228,522 0 0 0 0 0 0 Phisidia sp. 1BH 0 0 0 0 0,63816 0 0 0 0
Phyllocomus crocea 0 0 0 0 0 1,44613 0 0 0 Pionosyllis cf. maxima 0 0 0 0 0 0 0 0 0
Pionosyllis comosa 0 0 0 0 0 0 0 0 0 Pionosyllis epipharynx 0 0 20,6186 0 0,63816 0 0 0,51177 0,79208
Pionosyllis sp. 1 0 0 0 0 0 0 0 0 0 Pista corrientis 0 0 0 0 0 0 0 0 0 Pista cristata 0 0 0 0 0 0 0 0 0
Pista spinifera 0 0 0 0 0 0 0 0 0 Polycirrus cf. antarcticus 0 0 0 0 0 0 0 0 0
Polycirrus insignis 0 0 207,045 0,88849 0,63816 0 0 0 0,39604 Polycirrus sp. 1 0 0 0 0 0 0 0 0 0 Polycirrus sp. 2 0 0 0 0 0 0 0 0 0
Proceaea cf. mclearanus 0 0 0 0 0 0 0 0 0 Proclea cf. graffii 0 0 0 0 0 0 0 0 0
Pseudoscalibregma bransfieldium 0 0 1,71821 0 0,63816 0,72307 0 0,51177 0 Pseudoscalibregma papilia sp.n. 0 0,51414 0 0,44425 3,19081 0 0 0,51177 0
Pseudoscalibregma ursapium 0 0 4,29553 0 0 0,72307 0 0 0 Samytha sp. 1 0 0 0 0 0 0 0 0 0
Scalibregma inflatum 0 0 0,85911 0 1,27632 0 0 0 0 Sclerocheilus cf. antarcticus 0 0 0 0 0 0 0 0 0
Sosanopsis kerguelensis 0,68871 1,54242 6,87285 0 1,27632 0 0 3,07062 0,39604 Sosanopsis sp. 1 0 0 0 0 3,19081 0 0 1,02354 0
Sphaerodoropsis maculata sp.n. 0 0 0 0 0 0 0 0 0 Sphaerodoropsis parva 0 0 162,371 0 1,27632 7,95372 0,47326 1,53531 0,79208
Sphaerodoropsis polypapillata 0 2,57069 0 0 0 0 0 1,02354 0 Sphaerodoropsis simplex sp.n. 0 0 0 0 0,63816 0 0 0 0
Sphaerosyllis antarctica 0 0 0 0 0 0 0 0 0 Sphaerosyllis joinvillensis 0 0 244,845 0 0 0 0 0 0
Sphaerosyllis lateropapillata uteae 0 0 186,426 0 3,19081 2,89226 0 19,4473 0 Syllides articulosus 0 0 272,337 0 0 0 0 0 0 Terebellidae sp. 3 0 0 0 0 0 0 0 0 0
Terebellides stroemii 1,03306 7,71208 0,85911 2,66548 2,55265 0 0 0 0 Thelepides koehleri 0 0 0 0 0 0 0 0 0 Thelepides venustus 0 0 0 0 0 0 0 0 0 Thelepodinae sp. 1 0 0 0 0 0 0 0 0 0 Travisia cf. lithophila 0 0 0 0 0 0 0 0 0,39604 Travisia kerguelensis 0 0 1,71821 0 1,91449 0 0 0,51177 3,56436 Typosyllis cf. hyalina 0 0 4,29553 0 0 0 0 0 0 Typosyllis variegata 0 0 51,5464 0 0 0 0 0 0
APPENDIX- TABLES
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App.-Tab. 8: Resemblance matrix for ANDEEP I-III (Bray-Curtis, presence/absence transformation)
41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1 16-10 21-7 59-5 74-6 41-3 42-2 15,38 46-7 16,67 38,96
131-3 6,67 40,68 32,35 132-2 0,00 15,38 8,33 13,33 133-3 0,00 5,41 8,70 7,14 0,00 134-4 0,00 25,53 14,29 42,11 22,22 12,50 135-4 22,22 21,05 4,26 6,90 22,22 0,00 23,53 141-10 20,00 30,51 35,29 32,00 13,33 14,29 26,32 13,79 143-1 20,00 24,49 24,14 10,00 20,00 11,11 7,14 21,05 20,00 16-10 25,00 45,90 20,00 42,31 6,25 0,00 25,00 19,35 30,77 19,05 21-7 18,18 27,45 16,67 28,57 9,09 10,00 40,00 19,05 33,33 25,00 36,36 59-5 21,43 38,60 27,27 41,67 7,14 0,00 38,89 14,81 37,50 21,05 56,00 50,00 74-6 8,82 28,87 30,19 13,64 2,94 9,09 10,53 5,97 25,00 25,64 22,22 20,00 23,26 78-9 24,24 38,71 36,62 37,74 6,06 0,00 43,90 25,00 22,64 23,26 40,00 31,11 47,06 24,18 80-9 15,79 35,82 47,37 44,83 10,53 5,56 26,09 10,81 34,48 25,00 30,00 40,00 32,14 37,50 81-8 27,59 27,59 23,88 24,49 0,00 14,81 21,62 14,29 24,49 20,51 31,37 34,15 38,30 25,29 88-8 15,38 23,81 11,76 24,24 30,77 0,00 28,57 33,33 24,24 26,09 28,57 48,00 38,71 8,45
94-14 16,67 24,39 16,00 37,50 16,67 0,00 40,00 36,36 31,25 18,18 29,41 50,00 46,67 8,57 102-13 12,50 31,11 14,81 38,89 12,50 0,00 41,67 26,67 22,22 15,38 42,11 28,57 41,18 10,81 110-8 8,70 46,15 26,23 60,47 8,70 0,00 45,16 18,18 27,91 12,12 53,33 45,71 63,41 17,28 121-11 6,25 45,90 28,57 61,54 6,25 6,67 45,00 12,90 26,92 14,29 40,74 45,45 48,00 20,00 133-2 15,38 29,63 40,00 27,78 3,85 12,00 20,00 3,92 30,56 35,48 29,73 31,25 42,86 49,09 142-5 0,00 21,74 25,45 21,62 0,00 13,33 24,00 0,00 10,81 22,22 10,26 27,59 17,14 13,33 150-6 15,38 35,29 49,35 33,90 5,13 16,22 17,02 5,26 30,51 40,82 26,23 27,45 35,09 39,18 151-7 10,00 24,49 17,24 20,00 10,00 0,00 14,29 0,00 20,00 20,00 9,52 12,50 31,58 20,51 152-6 25,00 10,81 8,70 0,00 25,00 0,00 12,50 0,00 7,14 11,11 6,67 10,00 15,38 3,03 153-7 20,00 46,38 38,46 43,33 10,00 5,26 41,67 15,38 36,67 32,00 35,48 34,62 37,93 30,61 154-9 13,33 37,29 29,41 40,00 6,67 14,29 31,58 13,79 32,00 30,00 42,31 38,10 45,83 25,00
78-9 80-9 81-8 88-8 94-14 102-13 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 41-3 42-2 46-7
131-3 132-2 133-3 134-4 135-4 141-10 143-1 16-10 21-7 59-5 74-6 78-9 80-9 36,07 81-8 34,62 35,09 88-8 22,22 24,39 12,50
94-14 28,57 30,00 25,81 66,67 102-13 41,03 22,73 28,57 31,58 55,56 110-8 56,52 35,29 28,57 46,15 56,00 55,17 121-11 50,91 43,33 39,22 22,86 35,29 36,84 57,78 133-2 32,00 42,50 39,44 18,18 18,52 24,14 24,62 37,84 142-5 25,00 22,22 16,67 0,00 21,05 17,39 26,67 30,77 20,34 150-6 41,94 53,73 34,48 19,05 24,39 22,22 38,46 39,34 51,85 39,13 151-7 18,60 25,00 15,38 17,39 9,09 0,00 18,18 19,05 29,03 7,41 28,57 152-6 12,90 5,56 7,41 0,00 0,00 0,00 9,52 0,00 4,00 0,00 10,81 22,22 153-7 53,97 44,12 27,12 27,91 23,81 34,78 45,28 51,61 46,34 25,53 52,17 36,00 10,53 154-9 52,83 31,03 32,65 30,30 31,25 44,44 60,47 42,31 36,11 27,03 47,46 10,00 7,14 40,00
APPENDIX- TABLES
238
App.-Tab. 9: Resemblance matrix for ANDEEP I/II (Bray-Curtis, presence/absence transformation)
41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 41-3 42-2 15,38 46-7 16,67 38,96
131-3 6,67 40,68 32,35 132-2 0,00 15,38 8,33 13,33 133-3 0,00 5,41 8,70 7,14 0,00 134-4 0,00 25,53 14,29 42,11 22,22 12,50 135-4 22,22 21,05 4,26 6,90 22,22 0,00 23,53 141-10 20,00 30,51 35,29 32,00 13,33 14,29 26,32 13,79 143-1 20,00 24,49 24,14 10,00 20,00 11,11 7,14 21,05 20,00
App.-Tab. 10: Resemblance matrix for ANDEEP III based on standardized species abundances
(Bray-Curtis, fourth root transformation)
16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13 110-8 121-11 133-2 142-5 150-6 151-7 153-716-10 21-7 35,13 59-5 50,54 44,52 74-6 13,53 11,82 15,36 78-9 34,59 24,54 41,29 19,11 80-9 25,01 35,00 32,00 28,89 37,37 81-8 29,61 30,01 34,92 16,54 29,99 30,56 88-8 25,44 44,23 37,58 5,67 22,03 24,87 13,11 94-14 27,71 47,08 43,77 6,08 25,59 26,94 22,11 65,81
102-13 39,58 25,65 38,50 6,13 33,31 19,18 24,17 31,31 51,33 110-8 50,91 42,50 58,76 11,52 48,99 33,55 27,67 45,71 53,61 50,05
121-11 35,16 35,02 39,88 15,54 47,94 38,72 32,56 15,85 25,71 28,17 48,91 133-2 17,21 16,56 24,82 48,52 21,19 30,29 22,79 9,28 10,07 11,70 14,67 23,77 142-5 9,04 25,16 15,79 8,16 23,21 21,39 14,69 0,00 17,81 16,44 26,02 26,28 11,00 150-6 21,25 23,13 33,43 28,78 41,77 51,42 29,54 17,34 20,77 18,50 33,51 38,63 33,52 34,03 151-7 8,03 9,23 29,78 18,53 15,44 23,46 17,62 11,30 6,99 0,00 13,62 16,16 20,33 5,55 25,83 153-7 30,92 28,59 38,33 21,55 49,62 39,79 21,54 22,68 20,77 28,81 37,75 46,68 30,91 20,76 48,05 31,19 154-9 38,82 33,69 44,00 16,12 48,48 29,04 31,32 28,32 28,28 40,11 56,59 37,20 21,65 24,34 42,91 8,18 35,07
App.-Tab. 11: Ecological parameters taken into account for BIO-ENV analysis of ANDEEP I/II stations
(data taken from Diaz, 2004; Howe et al., 2004)
41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1 min grainsize (µm) 10 10 10 10 10 10 7 8 40 35 max grainsize (µm) 27 31 27 23 31 27 17 18 235 150
depth (m) 2360 3690 2889 3050 2086 1123 4069 4678 2313 774 02 depth (cm) 2.6 2.3 2.7 2.1 1.3 2.1 2.0 3.4
APPENDIX- TABLES
239
App.-Tab. 12: Global distribution of 84 named species found during the expeditions ANDEEP I-III (C-
cosmopolitan, LR- locally restricted to sampling area, SA- South Atlantic, SH- southern
hemisphere, SP- South Pacific, SU- sub-antarctic) (records compare chapter 2.4.2)
species family distribution station A I-II station A III
Amage sculpta Ehlers, 1908 Ampharetidae SU 42 21, 74, 80, 121,133
Ampharete kerguelensis McIntosh, 1885 Ampharetidae SA/SU 41, 46 16, 59, 78, 81, 133
Amphicteis gunneri (Sars, 1835) Ampharetidae C 46 78, 80, 121
Amythas membranifera Benham, 1921 Ampharetidae SU 133, 142
Anobothrella antarctica (Monro, 1939) Ampharetidae SU 143 74, 81, 150
Anobothrus gracilis (Malmgren, 1866) Ampharetidae C 74, 81, 102, 133, 154
Glyphanostomum scotiarum Hartman, 1978 Ampharetidae LR 42, 135 59, 74, 78
Grubianella antarctica McIntosh, 1885 Ampharetidae SA/SU 42 16, 153
Melinna cristata (Sars, 1851) Ampharetidae C 80, 150
Muggoides cf. cinctus Hartman, 1965 Ampharetidae C 80, 81
Neosabellides elongatus (Ehlers, 1912) Ampharetidae SU 134 21, 74, 78, 80, 81, 133, 153
Phyllocomus crocea Grube, 1877 Ampharetidae SU 59, 74, 81, 151
Sosanopsis kerguelensis Monro, 1939 Ampharetidae SA/SU 131, 134 16, 21, 59, 74, 78, 80, 88, 94,
102, 110, 133, 150, 153, 154 Glycera kerguelensis
McIntosh, 1885 Glyceridae SA/SU 41, 42, 46, 135, 141, 143
16, 21, 59, 74, 78, 80, 81, 88, 94, 102, 110, 133, 150, 153, 154
Goniada maculata Örstedt, 1843 Goniadidae C 16
Gyptis incompta Ehlers, 1912 Hesionidae SU/SP 74, 133
Kefersteinia fauveli Averincev, 1972 Hesionidae SA 141 16, 59, 74, 133, 151
Ophiodromus comatus (Ehlers, 1912) Hesionidae SU 74, 80, 81, 121, 133, 150, 151,
153 Aglaophamus paramalmgreni
Hartmann-Schröder & Rosenfeldt, 1992 Nephtyidae LR 42, 46 59, 78, 88, 110, 150, 151, 152, 153
Aglaophamus trissophyllus (Grube, 1866) Nephtyidae LR 42, 46 59, 78, 80, 110, 150, 154
Nereis eugeniae (Kinberg, 1866) Nereididae SH 42, 46, 132, 141
Ammotrypanella arctica McIntosh, 1879 Opheliidae C 131, 134 16, 59, 78, 102, 110, 153, 154
Ophelina breviata (Ehlers, 1913) Opheliidae SU/SA 42, 46, 131 16, 74, 78, 88, 102, 110, 121,
133, 150, 153, 154 Ophelina gymnopyge
(Ehlers, 1908) Opheliidae SA 42, 131 16, 59, 81, 110, 121, 133, 154
Ophelina nematoides (Ehlers, 1913) Opheliidae SU/SA 41, 42, 46, 131, 141 16, 74, 80, 153
Ophelina scaphigera (Ehlers, 1901) Opheliidae SA 16, 59, 133
Ophelina setigera Hartman, 1978 Opheliidae SA 46, 131 121
Phalacrostemma elegans Fauvel, 1911 Sabellariidae C 46 80
APPENDIX- TABLES
240
species family distribution station A I-II station A III
Ascleirocheilus ashworthi Blake, 1981 Scalibregmatidae SU 46
Axiokebuita minuta (Hartman, 1967) Scalibregmatidae C 46, 131, 133, 134,
141 74, 80, 81, 133
Axiokebuita millsi Pocklington & Fournier, 1987 Scalibregmatidae C 143
Hyboscolex equatorialis Blake, 1981 Scalibregmatidae SP 141
Kesun abyssorum Monro, 1930 Scalibregmatidae SU/SA 46, 131, 134, 141 16, 21, 59, 78, 80, 81, 94, 102,
110, 121, 133, 142, 150, 153, 154Oligobregma blakei
Schüller & Hilbig, 2006 Scalibregmatidae LR 46 121, 133
Oligobregma collare (Levenstein, 1978) Scalibregmatidae SU 42, 46, 131, 134 59, 74, 78, 80, 110, 121, 133,
142, 150, 151, 153, 154 Oligobregma hartmanae
Blake, 1981 Scalibregmatidae LR 59, 133, 150
Oligobregma notiale Blake, 1981 Scalibregmatidae SU 42
Oligobregma pseudocollare Schüller & Hilbig, 2006 Scalibregmatidae LR 46, 131 80, 121, 133, 150, 153
Oligobregma quadrispinosa Schüller & Hilbig, 2006 Scalibregmatidae LR 42, 131, 134, 141 21, 59, 88, 94, 110, 133, 151,
153 Pseudoscalibregma bransfieldium
(Hartman, 1967) Scalibregmatidae SU 42, 46, 143 74, 80, 88, 133, 150, 151, 153
Pseudoscalibregma usarpium Blake, 1981 Scalibregmatidae SU 42, 46, 131 80, 133, 151
Scalibregma inflatum Rathke, 1843 Scalibregmatidae C 42, 46 133, 150
Sclerocheilus cf. antarcticus Ashworth, 1915 Scalibregmatidae LR 133
Travisia cf. lithophila Kinberg, 1866 Scalibregmatidae SU/SP 154
Travisia kerguelensis McIntosh, 1885 Scalibregmatidae SU 46, 143 59, 74, 78, 133, 150, 153, 154
Clavodorum antarcticum Hartmann-Schröder & Rosenfeldt, 1990 Sphaerodoridae LR 46 142, 150
Ephesiella antarctica (McIntosh, 1885) Sphaerodoridae SU 42, 46, 143 74, 142, 153
Sphaerodoropsis parva (Ehlers, 1913) Sphaerodoridae SH 41, 42, 46,
141, 143 16, 21, 59, 74, 78, 80, 81, 133,
150, 151, 152, 153, 154 Sphaerodoropsis polypapillata
Hartmann-Schröder & Rosenfeldt, 1988 Sphaerodoridae SA 42, 131 16, 80, 110, 121, 153
Autolytus gibber Ehlers, 1897 Syllidae SU/SP 74
Braniella palpata Hartman, 1967 Syllidae C 141 21, 59, 74, 80, 81, 121, 133
Exogone heterosetosa McIntosh, 1885 Syllidae C 42 74
Exogone minuscula Hartman, 1953 Syllidae LR 74, 133, 151, 153
Pionosyllis cf. maxima Monro, 1930 Syllidae LR 74
Pionosyllis comosa Gravier, 1906 Syllidae SU 74
Pionosyllis epipharynx Hartman, 1953 Syllidae LR 46, 141 74, 133, 150, 153, 154
Proceraea cf. mclearanus (McIntosh, 1885) Syllidae SU/SP 74
Sphaerosyllis antarcticus Gravier, 1907 Syllidae LR
Sphaerosyllis joinvillensis Hartmann-Schröder & Rosenfeldt, 1988 Syllidae LR 141 74, 133
Sphaerosyllis lateropapillata uteae Hartmann-Schröder & Rosenfeldt, 1988 Syllidae LR 141, 143 74, 133, 150, 151, 153
Syllides articulosus Ehlers, 1897 Syllidae SU 74, 133
Typosyllis hyalina (Grube, 1863) Syllidae C 143 74, 133
Typosyllis variegata (Grube, 1860) Syllidae C 143 133
APPENDIX- TABLES
241
species family distribution station A I-II station A III
Eupistella grubei (McIntosh, 1885) Terebellidae SU 74, 80, 133
Hauchiella tribullata (McIntosh, 1869) Terebellidae C 42 74
Laphania cf. boecki Malmgren, 1866 Terebellidae C 46
Leaena antarctica McIntosh, 1885 Terebellidae SU 811, 102, 133, 153
Leaena arenilega Ehlers, 1913 Terebellidae SU 74, 153
Leaena collaris Hessle, 1917 Terebellidae SU/SA 16, 74, 133
Leaena pseudobranchia Levenstein, 1964 Terebellidae SU 74
Leaena wandelensis Gravier, 1907 Terebellidae SU 21, 154
Phisidia rubrolineata Hartmann-Schröder & Rosenfeldt, 1989 Terebellidae LR 133
Pista corrientis McIntosh, 1885 Terebellidae SU/SA 74
Pista cristata (Müller, 1776) Terebellidae C 74
Pista spinifera (Ehlers, 1908) Terebellidae SU 74
Polycirrus cf. antarcticus (Willey, 1902) Terebellidae SU 74
Polycirrus insignis Gravier, 1907 Terebellidae SU 46, 143 74, 78, 133, 142, 150, 154
Proclea graffii (Langerhans, 1884) Terebellidae C 46 74, 80
Streblosoma variouncinatum Hartmann-Schröder & Rosenfeldt, 1991 Terebellidae LR 141
Thelepides koehleri Gravier, 1911 Terebellidae LR 74
Thelepides venustus Levenstein, 1964 Terebellidae SU 74
Thelepus cincinnatus (Fabricius, 1880) Terebellidae C 74
Octobranchus antarcticus Monro, 1936 Trichobranchidae LR 46, 141 74, 80, 133, 153
Terebellides stroemi Sars, 1835 Trichobranchidae C 42, 46, 131 16, 59, 74, 78, 80, 81, 94, 102,
110, 133, 142, 150,
APPENDIX- TABLES
242
App.-Tab. 13: Polychaete community composition from world-wide deep-sea basins
station latitude longitude depth (m)
family composition
sampled area gear references
PAP 48°N 16°W 4800
Spionidae: 27 % Cirratulidae 26 %
Paraoinudae: 11 % Sabellidae: 6 % Pilargidae: 4 %
others: 26%
1.25 m2 Spade Box Core Glover et al., 2001
TAP 38°N 11°W 5035
Cirratulidae: 17 % Opheliidae: 15 % Spionidae: 15 %
Paraoinidae: 13 % Pilargidae: 11 %
others: 29 %
0.54 m2 Vegematic SBC Glover et al., 2001
MAP 31°N 21°W 4900
Spionidae: 22 % Cirratulidae: 19 % Paraoinidae: 18 % Ampharetidae: 8 %
Pilargidae: 7 % others: 26 %
1.25 m2 Spade Box Core Glover et al., 2001
EOS 20°N 30°W 4600
Cirratulidae: 31 % Spionidae: 16 %
Paraoinidae: 11 % Sabellidae: 7 %
Flabelligeridae: 9 % others: 26 %
2.0 m2 Spade Box Core Glover et al., 2001
O 21°N 31°W 4600
Cirratulidae: 25 % Spionidae: 19.1 %
Scalibregmatidae: 8.2 %Syllidae: 6.9 %
Ampharetidae: 6.6 %
1.25 m2 0.25 m2 USNEL
box corer
Cosso-Sarradin et al., 1998
M 19°N 21°W 3100
Cirratulidae: 26.4 % Spionidae: 18.5 % Sigalionidae: 6.4 % Paraoinidae: 6.1 %
Syllidae: 6.0 %
1.25 m2 0.25 m2 USNEL
box corer
Cosso-Sarradin et al., 1998
E 21°N 18°W 1700
Spionidae: 17.7 % Paraoinidae: 14.4 %
Syllidae: 8.8 % Cirratulidae: 7.2 % Capitellidae: 5.9 %
1.00 m2 0.25 m2 USNEL
box corer
Cosso-Sarradin et al., 1998
133 65°20.18'S 54°13.71'W 1138
Ampharetidae: 50 % Sabellidae: 15 %
Syllidae: 11 % Polygordiidae: 5 %
Polynoidae: 5 % Trichobranchidae: 5 %
others: 9 %
0.1 m2 Sandia box corer Hilbig, 2004
7 40°24.0'N 63°07.4'W 4626
Ampharetidae: 58 % Paraonidae: 14 % Spionidae: 14 % Cirratulidae: 4 % Hesionidae: 4 %
others: 6 %
0.77 m2 0.25 m2 box corer
Thistle, Yingst & Fauchald,1985
14 40°24.3'N 63°09.6'W 4626
Ampharetidae: 64 % Paraonidae: 15 % Spionidae: 13 % Cirratulidae: 4 % Hesionidae: 0 %
others: 4 %
0.77 m2 0.25 m2 box corer
Thistle, Yingst & Fauchald,1985
APPENDIX- TABLES
243
App.-Tab. 14: Composition of benthic deep-sea communities from world-wide sites
station latitude longitude depth (m) community structure
sampled area gear references
PNGS 1 ~11°S ~151°E 693
Polychaeta: 84 %Crustacea: 13 % Bivalvia: 0.6 % others: 2.4 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
PNGS 2 ~11°10'S ~151°E 2426
Polychaeta: 57 %Crustacea: 38 %
Bivalvia: 3 % others: 2 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
PNGS 3 ~11°20'S ~151°E 3264
Polychaeta: 64 %Crustacea: 32 %
Bivalvia: 2 % others: 2 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
CSB 13 ~13°S ~148°E 4008
Polychaeta: 55.3 %Crustacea: 30.3 %
Bivalvia: 5.4 % others: 9 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
CSB 4 ~12°S ~150°50'E 4350
Polychaeta: 67 %Crustacea: 28 %
Bivalvia: 3 % others: 2 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
WB 5 ~10°20'S ~151°20'E 2395
Polychaeta: 72 %Crustacea: 13 %
Bivalvia: 5 % others: 10 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
WB 6 ~10°10'S ~151°70'E 2775
Polychaeta: 37 %Crustacea: 52 %
Bivalvia: 0 % others: 11 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
WB 7 ~10°S ~152°E 2800
Polychaeta: 69 %Crustacea: 25 %
Bivalvia: 2 % others: 4 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
QT 5 ~14°50'S ~146°E 2256
Polychaeta: 64 %Crustacea: 17 %
Bivalvia: 4 % others: 15 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
CSP9 ~14°S ~146°50'E 1454
Polychaeta: 67 %Crustacea: 15 %
Bivalvia: 4 % others: 14 %
2-5 replicates
0.2 m2 Smith-
McIntyre grab
Alongi, 1992
Discol 3 (undisturbed) 7°S 88°W 4122-4201
Polychaeta: ~56 %Tanaidacea: ~21%
Isopoda: ~15 % Bivalvia: ~8 %
0.75 m2 0.25 m2 box corer Borowski & Thiel, 1998
A 78°57'N 7°42'W 183
Polychaeta: 47 %Crustacea: 33 %
Bivalvia: 16 % others: 4 %
0.5 m2 giant box corer Brandt & Schnack, 1999
B 78°59'N 5°42'W 748-803
Polychaeta: 57 %Crustacea: 33 %
Bivalvia: 3 % others: 7 %
0.75 m2 giant box corer Brandt & Schnack, 1999
C 78°56'N 5°11'W 1082-1101
Polychaeta: 38 %Crustacea: 44 %
Bivalvia: 2 % others: 16 %
0.75 m2 giant box corer Brandt & Schnack, 1999
I 78°58'N 3°59'W 1952-1965
Polychaeta: 24 %Crustacea: 61.33 %
Bivalvia: 7.34 % others: 7.33 %
0.75 m2 giant box corer Brandt & Schnack, 1999
APPENDIX- TABLES
244
station latitude longitude depth (m) community structure
sampled area gear references
H 3-18 28°N 155°W 5497-5825
Polychaeta: 54% Tanaidacea: 18 %
Bivalvia: 7 % others: 21 %
3 m2 0.25 m2 USNEL
box corerHessler & Jumars, 1974
73 39°46.5'N 70°43.3'W 1330-1470
Polychaeta: 43 % Mollusca: 27 %
Crustacea: 17 % others: 13 %
EBS Hessler & Sanders, 1967
62 39°26'N 70°33'W 2496
Polychaeta: 4 % Crustacea: 7 %
Echinodermata: 82 %others: 7 %
EBS Hessler & Sanders, 1967
64 38°46'N 70°06'W 2886
Polychaeta: 6 % Crustacea: 36 %
Echinodermata: 51 %others: 7 %
EBS Hessler & Sanders, 1967
70 36°23'N 67°58'W 4680
Polychaeta: 8 % Mollusca: 50 %
Crustacea: 32 % others: 10 %
EBS Hessler & Sanders, 1967
72 38°16'N 71°47'W 2864
Polychaeta: 6 % Crustacea: 16 %
Echinodermata: 64 %others: 14 %
EBS Hessler & Sanders, 1967
Upper Spencer Gulf, S. AUS 1-10 total: 12396
Polychaeta: 39 % (4834)
Hutchings, 1998, Hutchings et al., 1993
Hong Kong Harbor total: 11608
Polychaeta: 73 % (8351)
Hutchings, 1998, Mackie and Oliver, 1993
North Carolina 0-200 total: 1500
Polychaeta: 40 % (6384)
Hutchings, 1998, Day et al, 1971
H-39 50°58'N 171°37.5'W 7298
Polychaeta: 49 % Bivalvia: 11.5 %
Aplacophora: 10.5 %others: 29 %
0.25 m2 box corer Jumars & Hessler, 1976
2 19°58'S 2°59'E 5494
Polychaeta: 65 % Nematoda: 9 % Peracarida: 6 %
others: 20 %
1 m2 0.25 m2 USNEL
box corerKröncke & Türkay, 2003
3 19°07'S 3°51'E 5468
Polychaeta: 67 % Nematoda: 6 % Peracarida: 5 %
others: 22 %
1.25 m2 0.25 m2 USNEL
box corerKröncke & Türkay, 2003
4 18°16'S 4°44'E 5437
Polychaeta: 41 % Nematoda: 14 % Peracarida: 23 %
others: 22 %
0.25 m2 0.25 m2 USNEL
box corerKröncke & Türkay, 2003
5 17°07'S 4°41'E 5465
Polychaeta: 46 % Nematoda: 19 % Peracarida: 11 %
others: 24 %
1 m2 0.25 m2 USNEL
box corerKröncke & Türkay, 2003
6 16°17'S 5°27'E 5433
Polychaeta: 54 % Nematoda: 17 % Peracarida: 10 %
others: 19 %
1.25 m2 0.25 m2 USNEL
box corerKröncke & Türkay, 2003
7 40°24.0'N 63°07.4'W 4626
Polychaeta: 67 % Bivalvia: 11 % Isopoda: 11 % others: 11 %
0.77 m2 0.25 m2 USNEL
box corer
Thistle, Yingst & Fauchald,1985
14 40°24.3'N 63°09.6'W 4626
Polychaeta: 53 % Isopoda: 17 %
Tanaidacea: 24 % others: 6 %
0.77 m2 0.25 m2 box corer
Thistle, Yingst & Fauchald,1985
ACKNOWLEDGES
245
8 Acknowledges
First of all I would like to thank Dr. Brigitte Ebbe for introducing me to the “wonderful
world of polychaetes” and making me recognize that there is more beauty and variety in
this group than seen on first view. Thanks to you a worm has become much more than a
worm to me.
I’m more than grateful to Prof. Dr. J.-W. Wägele for giving me the chance to work on
this project, and offering me a helpful hand whenever needed.
I’d also like to thank HD Dr. G. Dehnhardt for agreeing to be co-corrector for this
study.
I’m thankful to Prof. Dr. Angelika Brandt, who together with B. Ebbe made the
ANDEEP expeditions possible and successful, and always showed great interest into
my work and progress. Also, the crew of the RV Polarstern, the participating scientist,
and my colleagues of the DZMB and the University of Hamburg deserve a warm “thank
you” for working day and night on the successful sampling and subsequent sample
management.
Many thanks are due to all my present and former colleagues from the department of
Animal Evolution, Ecology and Biodiversity of the Ruhr-University of Bochum –
especially Dr. I. Burghardt, Dr. N. Brenke, S. Janett, M. Rudschewski, C. Brefeldt, I.
Manstedt, Dr. M. Raupach, and Dr. C. Held – who accompanied me with helpful hands,
good advise, constructive discussions, and many pleasant conversations during my
studies.
I’d like to thank Prof. Dr. Ralph Tollrian for making work at the department very
uncomplicated and pleasant, and offering help whenever needed. Also, I’d like to thank
Dr. S. Brix, S. Kaiser, and Dr. W. Brökeland for many creative and often funny
discussion, and their helpful criticism and corrections.
ACKNOWLEDGES
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For volunteering to correct my spelling in spite of lack of scientific background, my
thanks go to Nic, Jen and Lorraine.
A big “Thank you” belongs to all my friends (you know who you are) who stuck with
me during the last three years and put up with my changing moods and lack of time
during the final weeks before completion of this thesis.
Viv, Mikel & Keanu, Christiane & Daniel, Johanna & Jens, and Birgit- I owe you a lot
of time and patience. Thanks for your constant support.
Last but not least I thank my family: Mutti, Jürgen und Tina- vielen Dank für euer
Vertrauen in mich. Wann immer ich euch brauche, seid ihr für mich da, und ich hoffe,
euch die ganze Geduld und Unterstützung irgendwann, irgendwie zurückgeben zu
können. Ich liebe Euch.
I also thank my dad, who can’t be here with me anymore. Nevertheless, he gave me the
constant will to persue my aims, and showed me that you can achieve anything in life as
long as you are happy with what you are doing.
I’d like to thank the German Science Foundation (DFG) for financing this project WA
530/29-1/2.
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9 Lebenslauf
PERSÖNLICHE DATEN
Myriam Schüller
Gröpperstr.11
58454 Witten
Geb. am 03. März 1980 in Aachen
Ledig, deutsch
SCHULBILDUNG
1986 – 1999 Allgemeine Hochschulreife (Abschlussnote 1,3)
STUDIUM
Oktober 1999 – September 2001 Vordiplom in Biologie an der Ruhr-Universität
Bochum (Note: 1,8)
September 2001 – Mai 2004 Diplom für Biologie an der Ruhr-Universität
Bochum (Note: 1,3)
Hauptfach Zoologie, Vertiefung in Ökologie
Nebenfach Botanik
Außerbiologisches Nebenfach Paläontologie Diplomarbeit am Lehrstuhl für Spezielle Zoologie (Titel: „Morphologie und Evolution der
Sexualdimorphismen bei den Sphaeromatidae (Crustacea: Isopoda) unter Berücksichtigung bestehender
phylogenetischer und ökologischer Kenntnisse“)
LEBENSLAUF
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BERUFSERFAHRUNG
Oktober 2004 - September 2006 Wissenschaftliche Mitarbeiterin am Lehrstuhl für
Spezielle Zoologie (Ruhr-Universität, Bochum).
Seit Oktober 2006 Wissenschaftliche Mitarbeiterin am Museum Alexander König,
Rheinische Friedrich Wilhelms Universität Bonn.
SONSTIGES
Seit 1998 Übungsleiterin im Jugendsportbereich (Fußball und Baseball)
2001 – 2004 Mitglied des Jugendvorstands des TuS Witten-Stockum 1945 e.V.
seit 2004 Mitglied der Gesellschaft für Biologische Systematik
seit 2006 Mitglied des Jugendvorstands der Sport Union Annen e.V., Abteilung
Baseball
seit 2006 Beiratsmitglied des Stockumer Theatervereins
Witten, den 28.06.2007
Myriam Schüller