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University of Southampton Research Repository Eprints Soton University of Southampton Research Repository ePrints Soton Copyright © and Moral Rights for this thesis are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the copyright holder/s. The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the copyright holders. When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given e.g. AUTHOR (year of submission) "Full thesis title", University of Southampton, name of the University School or Department, PhD Thesis, pagination http://eprints.soton.ac.uk THE UNIVERSITY OF SOUTHAMPTON The Ecology of Deep-Sea Holothurians by David Stewart Martin Billett A thesis submitted to the University of Southampton for the degree of Doctor of Philosophy. Department of Oceanography, The University, April 1988 Southampton, S09 5NH. Table of Contents Abstract V 1 Preface V1 i 1. Introduction. 1 Feeding. 2 Locomotion and bioturbation 3 Swimming. 11 4 Reproduction. 13 5 Population structure. 16 6 Growth and longevity. 18 7 Metabolism. 19 8 Chemical composition. 19 9 Abundance and biomass. 21 10 Spatial distributions. 25 11 Bathymetric zonation. 26 1.12 Geographic distributions. 30 1.13 Summary. 32 2. Materials and Methods, 35 2. 1 Study area. 35 2. 2 Sampling details 52 3. Taxonomic details. 67 3. 1 The family Ypsilothuriidae Heding, 1942 : 68 taxonomy and ontogenetic changes in body wall ossicles. Echinocucumis hispida 69 Ypsilothuria talismani 82 Ypsilothuria bitentaculata attenuata 93 Comparison of the species. 105 11 3. 2 Ossicles of juvenile Bathyplotes natans. 108 3. 3 A new species of apodid holothurian. Siniotrochus myriodontus 112 4. Species synopsis. 121 4. 1 Species list. 121 4. 2 Description of the species. 123 Dendrochirotida 123 Psolus squamatus 123 Staurocucumis abyssorum 125 Thyone gadeana 128 Dactylochirotida 130 Echinocucumis hispida 130 Ypsilothuria talismani 138 Aspidochirotida 140 Stichopus tremulus 140 Bathyplotes natans 148 Benthothuria funebris 154 Paelopatides grisea 165 Mesothuria candelabri 172 Mesothuria lactea 173 Mesothuria bifurcata 179 Mesothuria cathedral is 179 Mesothuria maroccana 180 Mesothuria intestinal is 181 Mesothuria verrilli 183 Paroriza pal lens 186 Paroriza prouhol 190 Pseudostlchopus villosus 193 Pseudostichopus sp. 196 111 Elasipodida 198 Deima validum 198 Oneirophanta mutabilis 205 Laetmogone violacea 215 Benthogone rosea 224 Benthodytes typica 232 Benthodytes sordida 233 Psychropotes depressa 238 Psychropotes longicauda 242 Psychropotes semperiana 250 Enypniastes diaphana 251 Peniagone diaphana 255 Peniagone azorica 260 Amperima rosea 261 Kolga hyalina 261 Elpidia sp. 274 Molpadiida 280 Molpadia blakei 280 Cherbonniera utriculus 283 Hedingia albicans 287 Apodida 288 Labidoplax southwardorum 288 Protankyra brychia 288 Myriotrochus bathybius 292 Myriotrochus giganteus 294 Siniotrochus myriodontus 294 Prototrochus zenkevitchi rockallensis 295 Parvotrochus belyaevi 296 IV 5. Ecological Discussion. 297 5. 1 Holothurian biomass and abundance in 297 relation to depth. 5. 2 Holothurian bathymetric zonation, scale 307 and relationship to environmental characters. 5. 3 Holothurian feeding and selectivity. 319 5.3.1 Distribution of chloropigments and 322 organic carbon within the sediment. 5.3.2 Qualitative comparison of chloro- 325 pigments in the sediment and holothurian gut contents. 5.3.3 Quantitative comparison of sediment 333 and holothurian gut contents. 5.3.4 Discussion. 335 5. 4 Small scale spatial distributions of 339 holothurians. 5. 5 Holothurian reproduction and life- 355 history strategies. 6. Bibliography. 365 Appendix 1. Station data. 390 Appendix 2. Publication list. 396 UNIVERSITY OF SOUTHAMPTON ABSTRACT FACULTY OF SCIENCE OCEANOGRAPHY Doctor of Philosophy THE ECOLOGY OF DEEP-SEA HOLOTHURIANS. by David Stewart Martin Billett The biology of 45 holothurian species, collected between 150 and 4830m in the Porcupine Seabight and the Porcupine Abyssal Plain in the northeast Atlantic, is described. Epibenthic sledge and otter trawl hauls have been used to calculate the abundance and biomass of each species, and spatial distributions have been examined using transect photography. The population size distributions and egg sizes of many species are related to their reproductive biology and life-history strategies. No relationship was found between holothurian biomass and depth. Holothurians may account for up to 90% of the total invertebrate megafaunal ash-free dry weight biomass, and the greatest biomass was recorded at a depth of 2000m. Bathymetric zonation is considered in detail using data on the abundance of each species in 100m depth bands. Some species on the upper slope occur in abundance only within a narrow depth range of 100 to 200m. Narrow zonation on the slope appears to be related to hydrographic features. Many species have an upper bathymetric limit of 1400 to 1500m which coincides with the bottom of the perm^n^nt thermocline and a change in water mass. This may act as a physiological barrier or a limit for larval dispersal. Abyssal species are more common on the abyssal plain than close to the continental slope. Studies using chlorophyll pigments as markers of detr- ital food show that holothurians feed indiscriminately on freshly deposited detritus, but that selectivity occurs once the detritus is incorporated into the sediment. One species, Kolga hyalina, forms dense aggregations that move as a herd across the seabed. The spatial pattern changes temporally and appears to be related to the seas- onal deposition of detritus. The taxonomic features of several species are discussed. Ontogenetic changes in characteristics of ypsilothuriid holothurians are examined in detail and a new species of apodid holothurian is described. VI Preface The approach of this thesis differs from that of many other dissertations since it was not initiated with a specific problem in mind. Rather, it was started when it was unclear how future investigations would progress and hence what the content of the final thesis would be. This uncertainty was unavoidable since the work was carried out as part of an on-going deep-sea benthic biology programme which changed its sampling priorities with time, and which was subject to the vagaries of the weather, cruise planning and gear reliability. It is necessary, therefore, to describe the aims of the deep-sea benthic biology programme at the Institute of Oceanographic Sciences (lOS), and to recount how that programme has progressed so that this thesis may be set in context. The data presented have been amassed between November 1977 and December 1986 from samples taken by lOS in the Porcupine Seabight (PSB) and Porcupine Abyssal Plain (PAP) off the southwest coast of Ireland (Fig. 1). More than 200 samples have been taken over a wide bathymetric range (400 to 4800m) and from 11 months of the calendar year. This has provided a unique collection of deep-sea fauna from which seasonal and bathymetric changes in deep-sea populations can be examined. The PSB was chosen as a study area because 1) the site was close to the British Isles, 2) the area provided soft- bottom habitats over a wide bathymetric range within a small geographic area, 3) the area had a congenial terrain with gentle slopes suitable for sampling with towed gear. VI1 4) the area was situated in temperate latitudes where any seasonal fluctuations in surface water productivity might be manifested at abyssal depths, and 5) the soft-bottom communities could be compared with those sampled previously by lOS off northwest Africa. A principal aim of the lOS benthic biology programme has been to quantify the abundance and biomass of benthic invertebrate megafauna using transect photography and an acoustically monitored epibenthic sledge. Good biomass data are essential if the energetics of the deep-sea ecosystem are to be understood but few relevant data are available. Little is also known about the structure of deep-sea communities or of the life-histories and behav- iour of deep-sea organisms. These subjects have now been investigated by lOS using a combination of techniques including transect and time-lapse photography, mid-water trawls, benthic trawls, epibenthic sledges, box corers, multiple corers, remote traps, and, more recently, submersibles. The other aims of the benthic biology progr- amme,therefore, have been to describe the bathymetric zonation, spatial distribution, behaviour, population structure, reproduction and feeding of many organisms ranging from meiofaunal protozoans, to macrofaunal crust- aceans, to megafaunal echinoderms and fish. Ultimately an understanding of the structure of deep-sea communities, trophic relationships, carbon flux etc., will be achieved. The many facets of the work carried out by lOS has provided a stimulating and varied approach to deep-sea biology. However, it has also had the disadvantage of providing many competing demands on sampling time, which together with poor weather and gear failures, have conspired to make some data sets less than perfect. Despite these problems the sampling intensity in the Porcupine Seabight has allowed one of the dominant deep- vin sea groups, the holothurians, to be studied in consid- erable detail. This thesis, therefore, sets
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