in the deep Weddell Sea (Southern Ocean, Antarctica) and adjacent ...

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


1.2.2.2 Glyceridae GRUBE 1850 15

1.2.2.3 Goniadidae KINBERG 1866 15


1.2.2.5 Nephtyidae GRUBE 1850 16

1.2.2.6 Nereididae JOHNSTON 1865 16


1.2.2.8 Sabellariidae JOHNSTON 1865 17



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


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






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


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.


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


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).


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).


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.


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


133-3 07.03.2002 65°20.15'S- 65°20.09'S


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


80-9 23.02.2005 70°38.45'S-70°39.18'S


81-8 24.02.2005 70°31.08'S-70°32.23'S


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


102-13 06.03.2005 65°33.18'S-65°34.32'S

36°33.24'W-36°31.05'W


110-8 10.03.2005 64°59.20'S-64°00.91'S

43°02.05'W-43°02.10'W


121-11 14.03.2005 63°38.27'S-63°37.31'S


133-2 16.03.2005 62°46.73'S-62°46.33'S


142-5 18.03.2005 62°11.36'S-62°11.36'S


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.


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


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


# 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


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


Ampharetidae sp. 4


Ampharetidae sp. 5



Ampharetidae sp. 6

Stations (No of specimens): 80-9 (1), 81-1 (1)

Ampharetidae sp. 7


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


Amphicteis sp. 1


150-6 (8), 153-7 (3)

Amphicteis sp. 2


Amphicteis sp. 3

Stations (No of specimens): 74-6 (5), 80-9 (1), 142-5 (1), 150-6 (2), 154-9 (1)

cf. Amphisamytha



Amythas membranifera BENHAM 1921



Records: Benham, 1921; Hartman, 1966; Monro, 1939

Anobothrella antarctica (MONRO 1939)



Records: Hartman, 1967; 1978; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Monro,

1939

Anobothrella sp. 1


Anobothrus gracilis (MALMGREN 1866)


Distribution: cosmopolitan, shelf down to 4817 m

Records: Hartmann-Schröder, 1996; Hessle, 1917; Hilbig & Blake, 2006

Anobothrus pseudoampharete sp.n.


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


Distribution: Drake Passage to Weddell Sea, 298 – 4678 m

Records: Hartman, 1978


Grubianella antarctica MCINTOSH 1885


Distribution: subantarctic to Southern Atlantic, 412 –4720 m

Grubianella sp. 1


Melinna cristata (SARS 1851)



Records: Ehlers, 1908; Hartman, 1966; 1967; Hartmann-Schröder, 1996; Hartmann-

Schröder & Rosenfeldt, 1989; Hessle, 1917; Holthe, 1986; Monro, 1930

Mugga sp. 1BH


Muggoides cf. cinctus HARTMAN 1965


Distribution: cosmopolitan, down to 4419 m

Records: Ebbe (pers. comment)

Muggoides sp. 1BH


Neosabellides elongatus (EHLERS 1912)


81-8 (1), 133-2 (1), 153-7 (2)


Records: Benham, 1927; Ehlers, 1912; 1913; Hartman, 1953; 1966; 1967; 1978;

Hartmann-Schröder & Rosenfeldt, 1989; 1991; Monro, 1930; 1936; 1939


cf. Neosabellides


Phyllocomus crocea GRUBE 1877



Records: Augener, 1932; Benham, 1921; Grube, 1877; Hartman, 1966; 1967;

Hartmann-Schröder & Rosenfeldt, 1989; Hessle, 1917; Monro, 1930; 1936;

1939

Samytha sp. 1


Sosanopsis kerguelensis MONRO 1939


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


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


143-1 (31), 16-10 (4), 21-7 (2), 59-5 (14), 74-6 (43), 78-9 (47), 80-9 (45),


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)


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


Bathyglycinde sp. 1BH


Bathyglycinde sp. 2


Goniada maculata ÖRSTEDT 1843



Records: Hartman, 1950; Hilbig & Blake, 2006


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.


Notochaetae not serrated………………………………………………………………….A. sp. 1

# Median antenna attached frontally (App.-Fig. 3J). Eversible pharynx with 40

distal papillae……………………………………………………………….Gyptis

only G. incompta


Amphiduros serratus sp.n.


Amphiduros sp. 1


Gyptis incompta EHLERS 1912


Distribution: subantarctic and Southern Pacific, 18 – 1582 m

Records: Ehlers, 1912; 1913; Wesenberg-Lund, 1961

Hesionidae sp. 1


aff. Hesionides


Kefersteinia fauveli AVERNICEV 1972


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


Micropodarke cylindripalpata sp.n.


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.


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)


150-6 (3), 151-7 (17), 153-7 (1)


Records: Ehlers, 1912; 1913; Hartman, 1964

Parasyllidea delicata sp.n.


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



Aglaophamus paramalmgreni HARTMANN-SCHRÖDER & ROSENFELDT 1992


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)


110-8 (1), 150-6 (2), 154-9 (4)

Distribution: Weddell Sea, Antarctic Peninsula, 400 – 4698 m

Records: Hartman, 1978

Micronephtys sp. 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


# Eversible pharynx with conical paragnaths. Ventral cirri simple throughout.

Falcigers and spinigers in posterior notopodia…………………………….Nereis

only N. eugeniae


Ceratocephale sp. 1BH


Nereis eugeniae (KINBERG 1866)


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


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


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.


Ammotrypanella arctica MCINTOSH 1879


102-13 (2), 110-8 (14), 121-11 (22), 153-7 (2), 154-9 (3)


Records: Ebbe (pers. comment); Hartman & Fauchald, 1971; McIntosh, 1879


Ammotrypanella cirrosa sp.n.


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.


121-11 (1), 153-7 (1)

Ammotrypanella mcintoshi sp.n.


Ophelina breviata (EHLERS 1913)


78-9 (1), 102-13 (1), 110-8 (1), 121-11 (6), 133-2 (19), 150-6 (1), 153-7 (1),

154-9 (1)


Records: Augener, 1932; Hartman, 1953; 1966; 1978; Hartmann-Schröder &

Rosenfeldt, 1989; 1991; Monro, 1930; 1936; 1939

Ophelina gymnopyge (EHLERS 1908)


110-8 (3), 121-11 (2), 133-2 (14), 154-9 (4)


Records: Ehlers, 1908; Hartman, 1952; 1953; 1966; Hartmann-Schröder & Rosenfeldt,

1989; 1991

Ophelina nematoides (EHLERS 1913)


16-10 (2), 74-6 (18), 80-9 (1), 153-7 (1)


Records: Ehlers, 1913; Hartman, 1966; 1967; 1978; Hartmann-Schröder & Rosenfeldt,

1989; 1991


Ophelina scaphigera (EHLERS 1901)



Records: Ehlers, 1900; 1901; Hartman, 1953; 1966; Monro, 1936

Ophelina setigera HARTMAN 1978



Records: Ebbe (pers. comment); Hartman, 1978

Ophelina ammotrypanella sp.n.


121-11 (7), 154-9 (8)

Ophelina robusta sp.n.


151-7 (2), 153-7 (7)

Ophelina sp. 3


Ophelina sp. 4


Ophelina sp. 5



3.2.8 Sabellariidae JOHNSTON 1865

Only two specimens have been sampled for the Sabellariidae, belonging to one species.

Phalacrostemma elegans FAUVEL 1911



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


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



Asclerocheilus ashworthi BLAKE 1981

Station (No of specimens): 46-7 (1)


Records: Blake, 1981

Axiokebuita millsi POCKLINGTON & FOURNIER 1987



Records: Pocklington & Fournier, 1987

Axiokebuita minuta (HARTMAN 1967)


74-6 (8), 80-9 (7), 81-8 (1), 133-2 (15), 150-6 (2)


Records: Blake, 1981; Hartman, 1967; 1978; Persson & Pleijel, 2005

Hyboscolex equatorialis BLAKE 1981


Distribution: South Sandwich Trench, west coast of South America, 0 – 2313 m


Kesun abyssorum MONRO 1930


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)


Records: Augener, 1932; Hartman, 1966; 1978; Monro, 1930; 1939


Oligobregma blakei SCHÜLLER & HILBIG 2007


Distribution: Weddell and Scotia Sea, 1582 – 2889 m

Records: Schüller & Hilbig, 2007

Oligobregma collare (LEVENSTEIN 1978)


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)


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


Oligobregma notiale 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)




Oligobregma quadrispinosa SCHÜLLER & HILBIG 2007


59-5 (6), 88-8 (1), 94-14 (2), 110-8 (1), 121-11 (1), 133-2 (1), 151-7 (1),

153-7 (10)



Oligobregma sp. 1


Pseudoscalibregma bransfieldium (HARTMAN 1967)


88-8 (1), 133-2 (2), 150-6 (1), 151-7 (1), 153-7 (1)


Records: Blake, 1981; Hartman, 1967; 1978

Pseudoscalibregma papilia n. sp.


150-6 (5), 153-7 (1)

Pseudoscalibregma ursapium BLAKE 1981


Distribution: subantarctic, 1178 – 3690m


Scalibregma inflatum RATHKE 1843



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


Sclerocheilus cf. antarcticus ASHWORTH 1915


Distribution: Antarcic Peninsula and Weddell Sea, 45 – 3138 m


Travisia kerguelensis MCINTOSH 1885


150-6 (3), 153-7 (1), 154-9 (9)


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



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


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.


Clavodorum antarcticum HARTMANN-SCHRÖDER & ROSENFELDT 1990


Distribution: Antarctic Peninsula and Weddell Sea, 142 – 3404 m

Records: Hartmann-Schröder & Rosenfeldt, 1990; 1992

Ephesiella antarctica (MCINTOSH 1885)



Records: Ehlers, 1912; 1913; Hartman,1964; 1978; Hartmann-Schröder & Rosenfeldt,

1992; McIntosh, 1885; Monro, 1930; 1936; 1939

Ephesiella hartmanae sp.n.


Sphaerodoropsis distincta sp.n.



Sphaerodoropsis maculata sp.n.


Sphaerodoropsis parva (EHLERS 1913)


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




Sphaerodoropsis simplex sp.n.


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


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


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


Autolytus gibber EHLERS 1897



Records: Ehlers, 1897; 1901; Fauvel, 1936; 1951; Gravier, 1906; 1907; Hartman, 1954;

1964; Hartmann-Schröder & Rosenfeldt, 1992; Monro, 1930; 1936


Brania sp. 1BH


Braniella palpata HARTMAN 1967


121-11 (2), 133-2 (32)


Records: Ebbe (pers. comment); Hartman, 1967; 1978

Exogone heterosetosa MCINTOSH 1885



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


Distribution: Antarctic Peninsula and Weddell Sea, 10 – 2014 m

Records: Blankenstey & Lana, 1986; Hartman, 1953; 1964; 1967; 1978

Exogone sp. 2BH


Exogone sp. 5


Exogone sp. 6



Exogone sp. 7


Pionosyllis comosa GRAVIER 1906



Records: Benham, 1921; 1927; Ehlers, 1912; 1913; Gravier, 1906; 1907; 1911; Harman,

1954; 1964; 1967; Monro, 1930

Pionosyllis epipharynx HARTMAN 1953


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


Distribution: Weddell Sea, 238 – 1047 m

Records: Hartman, 1964; Hartmann-Schröder & Rosenfeldt, 1988; Monro, 1930

Pionosyllis sp. 1


Proceraea cf. mclearanus (MCINTOSH 1885)



Records: Benham, 1927; Ehlers, 1912; 1913; Hartman, 1964; Hartmann-Schröder &

Rosenfeldt, 1990; 1992; McIntosh, 1885


Sphaerosyllis antarcticus GRAVIER 1907



Records: Hartmann-Schröder & Rosenfeldt, 1988; 1990; 1992

Sphaerosyllis joinvillensis HARTMANN-SCHRÖDER & ROSENFELDT 1988



Records: Hartmann-Schröder & Rsoenfeldt, 1988; 1992; San Martín & Parapar, 1996

Sphaerosyllis lateropapillata uteae HARTMANN-SCHRÖDER & ROSENFELDT 1988


151-7 (4), 153-7 (38)



Syllides articulosus EHLERS 1897



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)



Records: Ehlers, 1879; 1901; Fauvel, 1936; Gravier, 1911; Hartman, 1964; Hartmann-

Schröder, 1996; Hartmann-Schröder & Rosenfeldt, 1992; Licher, 1999; Willey,

1902


Typosyllis variegata (GRUBE 1860)

Stations ( no specimens): 143-1 (4), 133-2 (60)


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


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


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


Eupistella grubei (MCINTOSH 1885)




Records: Hartman, 1966; 1978; Levenstein, 1964; McIntosh, 1885

Eupistella sp. 1


Hauchiella tribullata (MCINTOSH 1869)



Records: Hartman, 1966; 1978; Hartmann-Schröder, 1996; Hartmann-Schröder &

Rosenfeldt, 1989; 1991; Hessle, 1917; Holthe, 1986; Monro, 1930; 1936

Laphania cf. boecki MALMGREN 1866



Records: Hessle, 1917

Leaena antarctica MCINTOSH 1885



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



Records: Benham, 1921; Ehlers, 1913; Hartman, 1966; 1978; Hartmann-Schröder &

Rosenfeldt, 1991; Hessle, 1917

Leaena collaris HESSLE 1917




Records: Hartman, 1966; Hartmann-Schröder & Rosenfeldt, 1989; 1991; Hessle, 1917;

Monro, 1930; 1936

Leaena cf. collaris HESSLE 1917


Leaena pseudobranchiata LEVENSTEIN 1964



Records: Hartman, 1966; Levenstein, 1964

Leaena wandelensis GRAVIER 1907



Records: Benham, 1927; Gravier, 1907; 1911; Hartman, 1952; 1966; Levenstein, 1964

Leaena sp. 4


Lysilla sp. 1BH


Phisidia rubrolineata HARTMANN-SCHRÖDER & ROSENFELDT 1989




Phisidia sp. 1BH


Pista corrientis MCINTOSH 1885




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)



Records: Augener, 1932; Ehlers, 1900; 1901; Gravier, 1907; 1911; Hartman, 1966;

1967; Hartmann-Schröder, 1996; Hessle, 1917; Levenstein, 1964

Pista spinifera (EHLERS 1908)



Records: Augener, 1932; Ehlers, 1908; 1913; Gravier, 1911; Hartman, 1966; 1967

Polycirrus cf. antarcticus (WILLEY 1902)


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)


Records: Fauvel, 1951; Gravier, 1907; Hartmann, 1966; Hartmann-Schröder &

Rosenfeldt, 1989; Hessle, 1917

Polycirrus sp. 1



Polycirrus sp. 2


Streblosoma variouncinatum HARTMANN-SCHRÖDER & ROSENFELDT 1991


Distribution: Weddell Sea and Drake Passage, 134 – 2313 m

Records: Hartmann-Schröder, 1991

Thelepides koehleri GRAVIER 1911



Records: Gravier, 1911; Hartman, 1966; 1967; 1978; Hartmann-Schröder & Rosenfeldt,

1989; 1991

Thelepides venustus LEVENSTEIN 1964



Records: Hartman, 1966; Levenstein, 1964

cf. Thelepides venustus LEVENSTEIN 1964


cf. Thelepus cincinnatus (FABRICIUS 1880)



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



Terebellidae sp. 1


Terebellidae sp. 2


Terebellidae sp. 3


Terebellidae sp. 4


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



Octobranchus antarcticus MONRO 1936


153-7 (1)

Distribution: Weddell Sea and Antarctic Peninsula, 267 – 3138 m

Records: Hartman, 1966; 1967; Monro, 1936

Terebellides stroemi SARS 1835


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)


Records: Augener, 1932; Ehlers, 1897; 1900; 1908; Fauchald, 1972; Hartman, 1966;

Hartmann-Schröder, 1996; Hessle, 1917; Holthe, 1986

cf. Terebellides sp. 1BH


Trichobranchidae sp. 1


Trichobranchidae sp. 2



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).


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.


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


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


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.


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.


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.


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.


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).


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


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


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)


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,


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.


2002, 65°19.83’S, 51°31.62’W, 3049-3050 m, EBS, ZMH P-24751


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


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).


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.


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.


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.


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


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.


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


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


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


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).


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.


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.


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


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


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


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


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.


Fig. 9: Pseudoscalibregma papilia sp.n. A- entire animal, B- anterior segments, C- posterior parapodium,

D- furcate chaetae, E- pygidium


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


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


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


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)


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


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)


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


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)


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)


110

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


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)


112

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


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


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


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


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.


117

Fig. 25: AvTD of ANDEEP I-III-stations (95 % significance range)

Fig. 26: VarTD of ANDEEP I-III-stations (95 % significance range)


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.


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


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


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


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


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.


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.


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).


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.


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.


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


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


3683-3690

3049-4069

2258-2313

Pseudosaclibregma bransfieldium (HARTMAN 1967)

Antarctic Peninsula

Bransfield Strait

Weddell Sea

off Elephant Island

Ona Basin


Ross Sea

335

769

412-2086

2889-2892

3683-3690

773-752

916

Pseudoscalibregma papilia n.sp.

Ona Basin

off Elephant Island


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.

DISCUSSION

142

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

DISCUSSION

143

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

DISCUSSION

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

DISCUSSION

148

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

150

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,

DISCUSSION

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.


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.

DISCUSSION

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,

DISCUSSION

156

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.

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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




APPENDIX- FIGURES

203


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

















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

















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

















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

















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
























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




















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
























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





























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



























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





























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



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


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


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


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


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





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





















APPENDIX- TABLES

226

species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total





















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




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



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


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




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



APPENDIX- TABLES

228

species 16-10 21-7 59-5 74-6 78-9 80-9 81-8 88-8 94-14 102-13























APPENDIX- TABLES

229

species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total























APPENDIX- TABLES

230

species 41-3 42-2 46-7 131-3 132-2 133-3 134-4 135-4 141-10 143-1




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





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 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



















APPENDIX- TABLES

232

species 110-8 121-11 133-2 142-5 150-6 151-7 152-6 153-7 154-9 total



















total 6669

APPENDIX- TABLES

233

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


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


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

236

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

237

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


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


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 %


EOS 20°N 30°W 4600

Cirratulidae: 31 % Spionidae: 16 %

Paraoinidae: 11 % Sabellidae: 7 %

Flabelligeridae: 9 % others: 26 %


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


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


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 %



APPENDIX- TABLES

243

App.-Tab. 14: Composition of benthic deep-sea communities from world-wide sites

station latitude longitude depth (m) community structure


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



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



2-5 replicates

0.2 m2 Smith-

McIntyre grab

Alongi, 1992

WB 5 ~10°20'S ~151°20'E 2395



2-5 replicates

0.2 m2 Smith-

McIntyre grab

Alongi, 1992

WB 6 ~10°10'S ~151°70'E 2775



2-5 replicates

0.2 m2 Smith-

McIntyre grab

Alongi, 1992

WB 7 ~10°S ~152°E 2800



2-5 replicates

0.2 m2 Smith-

McIntyre grab

Alongi, 1992

QT 5 ~14°50'S ~146°E 2256



2-5 replicates

0.2 m2 Smith-

McIntyre grab

Alongi, 1992

CSP9 ~14°S ~146°50'E 1454



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



0.5 m2 giant box corer Brandt & Schnack, 1999

B 78°59'N 5°42'W 748-803




C 78°56'N 5°11'W 1082-1101




I 78°58'N 3°59'W 1952-1965

Polychaeta: 24 %Crustacea: 61.33 %

Bivalvia: 7.34 % others: 7.33 %


APPENDIX- TABLES

244

station latitude longitude depth (m) community structure


H 3-18 28°N 155°W 5497-5825

Polychaeta: 54% Tanaidacea: 18 %


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 %


64 38°46'N 70°06'W 2886




70 36°23'N 67°58'W 4680

Polychaeta: 8 % Mollusca: 50 %

Crustacea: 32 % others: 10 %


72 38°16'N 71°47'W 2864




Upper Spencer Gulf, S. AUS 1-10 total: 12396

Polychaeta: 39 % (4834)

Hutchings, 1998, Hutchings et al., 1993

Hong Kong Harbor total: 11608


Hutchings, 1998, Mackie and Oliver, 1993

North Carolina 0-200 total: 1500


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


others: 22 %

1.25 m2 0.25 m2 USNEL


4 18°16'S 4°44'E 5437


others: 22 %

0.25 m2 0.25 m2 USNEL


5 17°07'S 4°41'E 5465


others: 24 %

1 m2 0.25 m2 USNEL


6 16°17'S 5°27'E 5433


others: 19 %

1.25 m2 0.25 m2 USNEL


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


14 40°24.3'N 63°09.6'W 4626

Polychaeta: 53 % Isopoda: 17 %

Tanaidacea: 24 % others: 6 %



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

246

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“)

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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

in the deep Weddell Sea (Southern Ocean, Antarctica) and adjacent ...

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