Biodiversita' Ed Evoluzione
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Alma Mater Studiorum – Università di Bologna DOTTORATO DI RICERCA IN BIODIVERSITA’ ED EVOLUZIONE Ciclo XXIII Settore scientifico-disciplinare di afferenza: BIO/05 ZOOLOGIA MOLLUSCS OF THE MARINE PROTECTED AREA “SECCHE DI TOR PATERNO” Presentata da: Dott. Paolo Giulio Albano Coordinatore Dottorato Relatore Prof.ssa Barbara Mantovani Prof. Francesco Zaccanti Co-relatore Prof. Bruno Sabelli Esame finale anno 2011 to Ilaria and Chiara, my daughters This PhD thesis is the completion of a long path from childhood amateur conchology to scientific research. Many people were involved in this journey, but key characters are three. Luca Marini, director of “Secche di Tor Paterno” Marine Protected Area, shared the project idea of field research on molluscs and trusted me in accomplishing the task. Without his active support in finding funds for the field activities this project would have not started. It is no exaggeration saying I would not have even thought of entering the PhD without him. Bruno Sabelli, my PhD advisor, is another person who trusted me above reasonable expectations. Witness of my childhood love for shells, he has become witness of my metamorphosis to a researcher. Last, but not least, Manuela, my wife, shared my objectives and supported me every single day despite the family challenges we had to face. Many more people helped profusely. I sincerely hope not to forget anyone. Marco Oliverio, Sabrina Macchioni, Letizia Argenti and Roberto Maltini were great SCUBA diving buddies during field activities. Betulla Morello, former researcher at ISMAR-CNR in Ancona, was my guide through the previously unexplored land of non-parametric multivariate statistics. Philippe Bouchet and Philippe Maestrati, Muséum National d'Histoire Naturelle (Paris), were welcoming and helpful during my work with them. Navigating through 23.000 specimens of Indo-Pacific triphorids from their research campaigns has been an experience I will never forget. It is rare to have so much material of excellent quality to study. I have been able to touch biodiversity first hand. Paolo Vestrucci and Roberto Colzani manage NIER Ingegneria Spa. Working with this company allowed the first contacts and activities with the Marine Protected Area which then developed into the survey that gave most of the data for this thesis. Paolo Giuntarelli, former director of the public body RomaNatura which manages the Marine Protected Area, kept on the work done by Luca Marini in safe-guarding funds for the field activities during Luca’s absence. Paolo & Marisa, Gabriella & Vincenzo were great in helping me combining work and family. Moreover, Gabriella was supportive and was one of the people who shared my objectives most. Francesca Evangelisti, Eimi Ailen Font, Michela Kuan, Wanda Moshfegh, Linda Tragni, Francesca Orrù, Maria Teresa Greco helped sorting samples out in the field or in laboratory. Anna Maria Candela, Department of Matematics, University of Bari, helped reviewing the English of some mathematical methodological parts. Mirco Travaglini and Stefania Brunetti, Library of the Department of Experimental Evolutionary Biology, helped in locating and obtaining literature. Further help during these three years has been given by Francesco Criscione, Carlo Froglia, Maria Cristina Gambi, Giuliana Gillone, Virginie Héros, Susan M. Kidwell, Constantine Mifsud, Stefano Palazzi, Paul Sammut, Cristiano Solustri, Helmut Zibrowius. Key-words: Mollusca, ecology, biodiversity, conservation, Posidonia oceanica, coralligenous, detritic pools, Secche di Tor Paterno 1 Index 1 Index .......................................................................................................................................................... 1 2 Executive summary ................................................................................................................................... 5 2.1 The molluscan diversity..................................................................................................................... 5 2.2 Biocoenoses characterization ............................................................................................................ 5 2.3 Analysis of the Posidonia leaves species assemblage ....................................................................... 5 2.4 Analysis of the Posidonia rhizomes species assemblage ................................................................... 6 2.5 Analysis of the coralligenous species assemblage ............................................................................. 6 2.6 Analysis of the detritic species assemblage ....................................................................................... 7 2.7 Agreement between death and living assemblages ........................................................................... 7 3 Introduction ............................................................................................................................................... 9 3.1 The studied area ................................................................................................................................. 9 3.2 Why sampling in a marine protected area?...................................................................................... 11 3.3 The molluscan fauna ........................................................................................................................ 11 3.4 Abbreviations .................................................................................................................................. 12 4 Materials and methods ............................................................................................................................. 13 4.1 Field activity .................................................................................................................................... 13 4.2 Laboratory sorting ........................................................................................................................... 15 4.3 Data analysis techniques .................................................................................................................. 15 4.3.1 Standardisation ........................................................................................................................ 16 4.3.2 Transformation ........................................................................................................................ 16 4.3.3 Similarity matrix ...................................................................................................................... 16 4.3.4 One-way ANalysis Of VARiance (ANOVA) .......................................................................... 17 4.3.5 Mann–Whitney U test .............................................................................................................. 18 4.3.6 Cluster analysis ........................................................................................................................ 18 4.3.7 Non-metric Multi-Dimensional Scaling (MDS) ...................................................................... 19 4.3.8 Multivariate ANalysis Of SIMilarities (ANOSIM) ................................................................. 19 4.3.9 SIMilarity PERcentage breakdown (SIMPER) ....................................................................... 20 4.3.10 PERmutational Multivariate ANalysis Of VAriance (PERMANOVA) .................................. 21 4.4 Biodiversity indices ......................................................................................................................... 21 4.4.1 Number of species (S) ............................................................................................................. 21 4.4.2 Number of specimens (N) ........................................................................................................ 22 4.4.3 Margalef’s species richness (d) ............................................................................................... 22 4.4.4 Shannon index (H’) ................................................................................................................. 22 4.4.5 Equitability (J’) ........................................................................................................................ 22 4.4.6 Simpson index (λ) .................................................................................................................... 22 4.5 Biodiversity estimators .................................................................................................................... 23 4.6 Trophic groups and feeding guilds .................................................................................................. 23 4.7 The role of species in describing biocoenoses ................................................................................. 24 5 Sampling results and efficacy .................................................................................................................. 25 5.1 Results ............................................................................................................................................. 25 5.2 Discussion ........................................................................................................................................ 29 6 The molluscan diversity .......................................................................................................................... 30 1 6.1 Results ............................................................................................................................................