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Interactive Processes in a Lanice Conchilega (Annelida: Polychaeta) Dominated Intertidal Community

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Interactive Processes in a Lanice Conchilega (Annelida: Polychaeta) Dominated Intertidal Community 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 UNIVERSITY OF SOUTHAMPTON INTERACTIVE PROCESSES IN ALANICE CONCHILEGA (ANNELIDA: POLYCHAETA) DOMINATED INTERTEDAL COMMUNITY by GRANT A. ROWE Submitted for the Qualification of PhD SCHOOL OF OCEAN AND EARTH SCIENCE September 1999 UNIVERSITY OF SOUTHAMPTON ABSTRACT FACULTY OF SCIENCE SCHOOL OF OCEAN AND EARTH SCIENCE Doctor of Philosophy INTERACTIVE PROCESSES IN ALANICE CONCHILEGA (ANNELIDA: POLYCHAETA) DOMINATED INTERTIDAL COMMUNITY by Grant A. Rowe An investigation was conducted into the factors influencing a soft-sediment intertidal benthic community dominated by the large tube building polychaete Lanice conchilega (Pallas, 1766). The environmental and biological characteristics of a mid-shore sampling site on a sheltered relatively sandflat were monitored over a two period. A total of 115 year taxa were recorded. Temporal variability in the community is discussed in relation to the sporadic occurrence of macroalgae, harsh winter conditions and physical disturbance to the sedimentary environment. Lanice conchilega had a significantly contagious distribution on the shore. The population was analysed using width measurements of both worms and tubes. Recruiting Lanice conchilega juveniles observed the first during year of the study were initially randomly distributed, but as these individuals developed the overall distribution of the population returned to being contagious. The presence of excess tubes towards the end of the sampling period indicated that individual Lanice conchilega were dying, being removed by predation or emigrating. Further sampling was designed to investigate the macrofauna and meiofauna in areas of sediment containing different densities of Lanice conchilega tubes. A consistently significant positive correlation existed between tube density and macrofaunal abundance, which was attributed to sediment-mediated interactions. At higher tube densities, an increase in the depth of sediment above the underlying clay equated to an increase in the volume of available infaunal space. In addition, Lanice conchilega-derived mucus and its associated microbes potentially enhanced levels of food resources available to feeders. deposit Species living at or near the sediment-water interface would also have been positively influenced by any sediment stabilisation in high tube density areas. Possible causes of a positive correlation observed between tube and density Exogone hebes abundance are discussed. No significant relationship existed between tube density and the abundance of macrofaunal juveniles, indicating that any inhibitory or facilitatory Lanice conchilega-juvenile interactions either were insignificant or acting in opposition to to produce an overall neutral effect. Details of the developmental morphology of several polychaete species are described. Meiofaunal abundance (in particular nematode and harpacticoid abundance) was consistently in greater samples taken from areas of high tube density. Potential mechanisms responsible for this relationship include the provision of habitat and heterogeneity areas of refuge by tubes, and the enhancement of sediment stability and food resources by mucus. Particle accumulation in high tube density areas also potentially influenced meiofaunal settlement and resettlement patterns. CONTENTS Abstract ............ i Contents ............ ii List of Tables v List of Figures ........... vi Acknowledgements .......... xiv CHAPTER 1 Introduction 1 1.1 Background . ',; . .1 . 1.2 The Site Study 6 1.3 Lanice conchilega (Pallas, 1766) ....... 8 1.4 Aims 10 CHAPTER 2 The Physico-Chemical Environment at Solent Breezes 12 2.1 Introduction 12 2.2 Meteorological and Hydrological Data . .12 . 2.2.1 Meteorological data 12 2.2.2 Seawater temperature and salinity measurements 13 2.3 Sediment Analysis . .18 . 2.3.1 Methods 19 2.3.2 Results 23 2.4 Discussion ........... 37 2.4.1 Meteorological and Hydrological Parameters 37 2.4.2 Sediment Parameters 38 CHAPTER 3 The Macrofaunal Sampling Transect 44 3.1 Introduction 44 3.2 Field Work and Laboratory Methods ....... 44 3.3 Data Analysis. .......... 46 3.3.1 Univariate analyses 46 3.3.2 Multivariate analyses 47 3.3.3 The Infaunal Trophic Index 50 3.4 Results 52 3.4.1 Univariate analyses 5 2 3.4.2 Multivariate analyses 5 5 3.4.3 The Infaunal Trophic Index 61 3.4.4 Summary 63 3.5 Discussion . ......... 64 CHAPTER 4 The Macrofaunal Monitoring Programme. .73 . 4.1 Introduction ........... 73 4.2 Field Work and Laboratory Methods ....... 73 4.3 Data Analysis........... 74 4.3.1 Univariate analyses 74 4.3.2 Multivariate analyses 75 4.3.3 The Infaunal Trophic Index 75 4.4 Results 76 4.4.1 Univariate analyses 76 4.4.2 The dominant species 81 4.4.3 Multivariate analyses 100 4.4.4 The Infaunal Trophic Index 112 4.4.5 Summary 115 4.5 Discussion . .117 . CHAPTER 5 The Lattice conchilega Population . .138 . 5.1 Introduction ........... 138 5.2 Methods 138 5.3 Results 140 5.3.1 Size-frequency analysis ofLanice conchilega and its tubes .141 . 5.3.2 Growth curves 146 5.3.3 Temporal variability in abundance . .148 . 5.4 Discussion ........... 152 ni CHAPTER 6 Lattice conchilega Tube Density Experiments .161 . 6.1 Introduction ........... 161 6.2 Field Work and Laboratory Methods . .161 . 6.2.1 Lanice and its conchilega tubes 161 6.2.2 Lanice tubes conchilega and the other infauna 162 6.3 Results 163 6.3.1 Lanice and its conchilega tubes 163 6.3.2 Lanice tubes conchilega and the other infauna 165 6.3.3 Lanice tubes conchilega and Exogone hebes 171 6.4 Discussion ........... 173 CHAPTER 7 The Meiofauna . 186 7.1 Introduction 186 7.2 Field Work and Laboratory Methods 186 7.2.1 The meiofauna permanent 187 7.2.2 The meiofauna temporary (juvenile macrofauna) 188 7.3 Results .190 . 7.3.1 The meiofauna permanent 190 7.3.2 The meiofauna temporary (juvenile macrofauna) 198 7.4 Discussion 227 CHAPTER 8 Summary and Conclusions 251 APPENDIX 1 261 APPENDIX 2 . 266 REFERENCES 267 IV LIST OF TABLES Page Table 3.1 The taxonomic literature used to identify the macrobenthic fauna. 45 Table 3.2 Values of the F-statistic calculated for Transect Sites 1 to 5. 54 Table 3.3 Results of the BIO-ENV procedure for the sampling transect. Combinations of the five sediment parameters giving the best matches between the faunal and abiotic matrices. Correlations were measured using weighted Spearman rank correlation (pw). 60 Table 3.4 Values of the Infaunal Trophic Index for Transect Sites 1-5. 61 Table 4.1 The dominant species at Solent Breezes. 81 Table 4.2 Combinations of the nine environmental parameters giving the best matches between the faunal and abiotic similarity matrices. Correlations were measured using weighted Spearman rank correlation (pw). 106 Table 6.1 Correlation coefficients (r) and p values calculated using linear regression analysis for the number of tube heads per sample (T) against the number of individuals (N), the number of species (s) and the diversity (//*). 166 Table 6.2 Correlation coefficients and p values calculated using Spearman rank order correlation for tube head density against the first two principal components. 170 Table 6.3 The correlation coefficients (r) and p values calculated for the relationship between Lanice conchilega tube heads and Exogone hebes. Ill LIST OF FIGURES Page Figure 1.1 The geographical position of the sampling site. 7 Figure 2.1 Meteorological data from the nearby Solent MRSC station (November 1990 - October 1992). Values shown for each sampling date refer to mean values calculated for the "between-sampling period" leading up to that date. 14 Figure 2.2 Meteorological data from the nearby Solent MRSC station (February 1993 - March 1994). Values shown for each sampling date refer to mean values calculated for the "between-sampling period" leading up to that date. 15 Figure 2.3 The temperature and salinity of the seawater adjacent to the site sampling at Solent Breezes - measurements made during each of the macrofaunal visits monthly sampling (November 1990 - October 1992). 16 Figure 2.4 The temperature and salinity of the seawater adjacent to the sampling site at Solent Breezes - measurements made during each of the meiofaunal visits monthly sampling (February 1993 - March 1994). 17 Figure 2.5 The total organic content of surface sediment samples taken during monthly macrofaunal sampling visits (upper graph) and monthly meiofaunal sampling visits (lower graph). Mean values ( S.D.) shown. 24 Figure 2.6 The total organic content of surface sediment samples from different densities of Lanice conchilega tubes (upper graph) and from five transect sites extending from high tide (Site 1) to low tide (Site 5) (lower graph). Mean values ( S.D.) shown. 25 Figure 2.7 Fluctuations in the five
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