DOCSLIB.ORG

Evolutionary Origin and Maintenance of Sociality in the Small Carpenter Bees

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Evolutionary Origin and Maintenance of Sociality in the Small Carpenter Bees Evolutionary Origin and Maintenance of Sociality in the Small Carpenter Bees by Sandra M. Rehan, B. Sc. A Thesis submitted to the Department of Biological Sciences in partial fulfilment of the requirements for the degree of Doctor of Philosophy Brock University St. Catharines, Ontario © Sandra M. Rehan, 2011 TABLE OF CONTENTS TABLE OF CONTENTS ........................................................................................................... .i LIST OF TABLES ................................................................................................................... vii LIST OF FIGURES .................................................................................................................. .ix THESIS SUMMARY ............................................................................................................... xi ACKNOWLEDGEMENTS .................................................................................................... xiii OVERVIEW OF THESIS CONTENT................................................................................... xiv CHAPTER 1 General Introduction ................................................................. ,/............................................... 1 Part 1: Phylogenetic Contrasts and the Origins of Sociality...................................................... 1 Age and Phylogenetic Relationships in the Xylocopinae ............................................. .2 Social Behaviour of the Xylocopinae ........................................................................... ..4 Social Evolution of the Ceratinini .................................................................................. 6 Part 2: Evolutionary Explanations of Altruism.......................................................................... 7 Kin Selection and the Evolutionary Origin of Eusociality............................................ 8 Maternal Manipulation as an Amendment to Kin Selection........................................ 10 Ecological Constraints and Selection for Group Living .............................................. 12 Model Systems to Study Social Origins ....................................................................... 12 The Small Carpenter Bees ............................................................................................ 13 Research Aims .......................................................................................................................... 14 CHAPTER 2 Social polymorphism in the Australian small carpenter bee, Ceratina (Neoceratina) australensis ............................................................................................................................... 15 Introduction ............................................................................................................................. 16 Methods ................................................................................................................................... 18 ii Statistical Analyses ..................................................................................................... 20 Results ..................................................................................................................................... 20 Frequency of social nesting .......................................................................................... 20 Colony cycle .......................................................... ....................................................... 21 Maternal care and longevity......................................... ................................................ 22 Reproductive hierarchies in social colonies ................................................................ .23 Nest architecture and brood productivity ..................................................................... 26 Discussion ................................................................................................................................ 27 Maternal behaviour and social preadaptations ..............{ ............................................ 28 Female dispersal and social nesting ............................................................................. 29 Reproductive differentiation in social colonies ............................................................ 30 Brood productivity and social benefits ......................................................................... 32 Acknowledgments .................................................................................................................... 34 References ............................................................................................................................... 35 Tables 1 - 2 .............................................................................................................................. 39 Figures 1 - 4 .............................................................................................................................. 41 CHAPTER 3 Fitness consequences of ecological constraints and implications for the evolution of sociality in an incipiently social bee ....................................................................................................... 46 Introduction ............................................................................................................................. 47 Methods ................................................................................................................................... 50 Statistical Analyses ............................................................................. 52 Results ..................................................................................................................................... 53 Weather variation among brood rearing periods .......................................................... 53 Nest site availability.......................................... ........................................................... 53 iii Variation in brood development. .................................................................................. 54 Variation in reproductive success in solitary nests ...................................................... 55 Reproductive success in solitary versus social nests .................................... :............... 56 Discussion................................................................................................................................ 57 Environmental constraints on development rates and brood production ..................... 58 The effect of brood parasitism on reproductive success .............................................. 59 Nest substrate limitation and social nesting ................................................................. 60 Acknowledgments .................................................................................................................... 61 References ................................................................................ ,/............................................ 62 Table 1...................................................................................................................................... 68 Figures 1 - 4 .............................................................................................................................. 69 CHAPTER 4 The costs and benefits of sociality in a facultatively social bee ............................................... 74 Introduction ............................................................................................................................. 7 5 Methods ................................................................................................................................... 79 Life history ................................................................................................................... 79 Nest collections and brood production ......................................................................... 79 Relatedness estimates ........................................................................ ;.......................... 80 Results ..................................................................................................................................... 81 Brood productivity....................................................................................................... 81 Relatedness estimates ................................................................................................... 82 Direct fitness of solitary and social females ................................................................. 83 Inclusive fitness of solitary and social females ............................................................ 84 Discussion ................................................................................................................................ 85 iv Implications of dispersal for social potential... ............................................................ 86 Reproductive success and direct fitness ....................................................................... 87 Cooperative behaviour and indirect fitness .................................................................. 88 Conclusions .............................................................................................................................. 88 Acknowledgments ...................................................................................................................
Load more

Recommended publications
  • Development of Native Bees As Pollinators of Vegetable Seed Crops
    Development of native bees as pollinators of vegetable seed crops Dr Katja Hogendoorn The University of Adelaide Project Number: VG08179 VG08179 This report is published by Horticulture Australia Ltd to pass on information concerning horticultural research and development undertaken for the vegetables industry. The research contained in this report was funded by Horticulture Australia Ltd with the financial support of Rijk Zwaan Australia Pty Ltd. All expressions of opinion are not to be regarded as expressing the opinion of Horticulture Australia Ltd or any authority of the Australian Government. The Company and the Australian Government accept no responsibility for any of the opinions or the accuracy of the information contained in this report and readers should rely upon their own enquiries in making decisions concerning their own interests. ISBN 0 7341 2699 9 Published and distributed by: Horticulture Australia Ltd Level 7 179 Elizabeth Street Sydney NSW 2000 Telephone: (02) 8295 2300 Fax: (02) 8295 2399 © Copyright 2011 Horticulture Australia Limited Final Report: VG08179 Development of native bees as pollinators of vegetable seed crops 1 July 2009 – 9 September 2011 Katja Hogendoorn Mike Keller The University of Adelaide HAL Project VG08179 Development of native bees as pollinators of vegetable seed crops 1 July 2009 – 9 September 2011 Project leader: Katja Hogendoorn The University of Adelaide Waite Campus Adelaide SA 5005 e-mail: [email protected] Phone: 08 – 8303 6555 Fax: 08 – 8303 7109 Other key collaborators: Assoc. Prof. Mike Keller, The University of Adelaide Mr Arie Baelde, Rijk Zwaan Australia Ms Lea Hannah, Rijk Zwaan Australia Ir. Ronald Driessen< Rijk Zwaan This report details the research and extension delivery undertaken in the above project aimed at identification of the native bees that contribute to the pollination of hybrid carrot and leek, and at the development of methods to enhance the presence of these bees on the crops and at enabling their use inside greenhouses.
    [Show full text]
  • Changing Paradigms in Insect Social Evolution: Insights from Halictine and Allodapine Bees
    ANRV297-EN52-07 ARI 18 July 2006 2:13 V I E E W R S I E N C N A D V A Changing Paradigms in Insect Social Evolution: Insights from Halictine and Allodapine Bees Michael P. Schwarz,1 Miriam H. Richards,2 and Bryan N. Danforth3 1School of Biological Sciences, Flinders University, Adelaide S.A. 5001, Australia; email: Michael.Schwarz@flinders.edu.au 2Department of Biological Sciences, Brock University, St. Catharines, ON L2S 3A1, Canada; email: [email protected] 3Department of Entomology, Cornell University, Ithaca, New York 14853; email: [email protected] Annu. Rev. Entomol. 2007. 52:127–50 Key Words The Annual Review of Entomology is online at sex allocation, caste determination, phylogenetics, kin selection ento.annualreviews.org This article’s doi: Abstract 10.1146/annurev.ento.51.110104.150950 Until the 1980s theories of social insect evolution drew strongly Copyright c 2007 by Annual Reviews. on halictine and allodapine bees. However, that early work suffered All rights reserved from a lack of sound phylogenetic inference and detailed informa- 0066-4170/07/0107-0127$20.00 tion on social behavior in many critical taxa. Recent studies have changed our understanding of these bee groups in profound ways. It has become apparent that forms of social organization, caste de- termination, and sex allocation are more labile and complex than previously thought, although the terminologies for describing them are still inadequate. Furthermore, the unexpected complexity means that many key parameters in kin selection and reproductive skew models remain unquantified, and addressing this lack of informa- tion will be formidable.
    [Show full text]
  • Nestmate Recognition in the Large Carpenter Bee, Xylocopa Virginica
    Nestmate Recognition in the Large Carpenter Bee, Xylocopa virginica by Marianne Peso, B.A.Sc. A Thesis submitted to the Department of Biological Sciences in partial fulfillment of the requirements for the degree of Master of Science Brock University St. Catharines, Ontario Marianne Peso, 2008 lAMESAGTOSONLIBRAFY BROCK UNIVERSITY ST. CATHARINES m Abstract Many species of social insects have the ability to recognize their nestmates. In bees, sociality is maintained by bees that recognize which individuals should be helped and which should be harmed in order to maximize fitness (either inclusive or individual) (Hamilton 1964; Lin and Michener 1972). Since female bees generally lay eggs in a single nest, it is highly likely that bees found cohabitating in the same nest are siblings. According to the kin selection hyjKjthesis, individuals should cooperate and avoid aggression with same sex nestmates (Hamilton 1964). However, in opposite sex pairs that are likely kin, aggression should increase among nestmates as an expression of inbreeding avoidance (Lihoreau et al. 2007). Female bees often guard nest entrances, recognizing and excluding foreign conspecific females that threaten to steal nest resources (Breed and Page 1991). Conversely, males that aggressively guard territories should avoid aggression towards other males that are likely kin (Shellman-Reeve and Gamboa 1984). In order to test whether Xylocopa virginica can distinguish nestmates from non-nestmates, circle tube testing arenas were used. Measures of aggression, cooperation and tolerance were evaluated to determine the presence of nestmate recognition in this species. The results of this study indicate that male and female X. virginica have the ability to distinguish nestmates from non-nestmates.
    [Show full text]
  • Genes Involved in Convergent Evolution of Eusociality in Bees
    Genes involved in convergent evolution of eusociality in bees S. Hollis Woodarda,1, Brielle J. Fischmana,1, Aarti Venkatb, Matt E. Hudsonb, Kranthi Varalab, Sydney A. Cameronc, Andrew G. Clarkd, and Gene E. Robinsona,c,e,f,2 aProgram in Ecology, Evolution, and Conservation Biology, Departments of bCrop Sciences and cEntomology, eInstitute for Genomic Biology, and fNeuroscience Program, University of Illinois, Urbana, IL 61801; and dDepartment of Molecular Biology and Genetics, Cornell University, Ithaca, NY 14853 Contributed by Gene E. Robinson, March 12, 2011 (sent for review February 17, 2011) Eusociality has arisen independently at least 11 times in insects. different social lifestyles among relatively closely related species. Despite this convergence, there are striking differences among Furthermore, the extensive knowledge of bee natural history (8, eusocial lifestyles, ranging from species living in small colonies 13, 14) provides a valuable framework for developing hypotheses with overt conflict over reproduction to species in which colonies about the adaptive significance of genetic changes detected in contain hundreds of thousands of highly specialized sterile work- eusocial bee lineages. ers produced by one or a few queens. Although the evolution of To study patterns of molecular evolution associated with eusociality has been intensively studied, the genetic changes eusociality in bees, we generated ~1 Gbp of expressed sequence involved in the evolution of eusociality are relatively unknown. tags (ESTs) from a set of nine bee species (Table S1). This set of We examined patterns of molecular evolution across three in- species reflects the remarkable social diversity in bees by in- dependent origins of eusociality by sequencing transcriptomes of cluding eusocial and non-eusocial species; three origins of euso- nine socially diverse bee species and combining these data with ciality (9, 10); and two different forms of eusocial lifestyle, “highly genome sequence from the honey bee Apis mellifera to generate eusocial” and “primitively eusocial” (ref.
    [Show full text]
  • Molecular Phylogenetics of Allodapine Bees, with Implications for the Evolution of Sociality and Progressive Rearing
    Syst. Biol. 52(1):1–14, 2003 DOI: 10.1080/10635150390132632 Molecular Phylogenetics of Allodapine Bees, with Implications for the Evolution of Sociality and Progressive Rearing MICHAEL P. S CHWARZ,1 NICHOLAS J. BULL,1 AND STEVEN J. B. COOPER2 1School of Biological Sciences, Flinders University, G.P.O. Box 2100, Adelaide, South Australia 5001, Australia 2Evolutionary Biology Unit, South Australian Museum, Adelaide, South Australia 5000, Australia Abstract.—Allodapine bees have long been regarded as providing useful material for examining the origins of social behavior. Previous researchers have assumed that sociality arose within the Allodapini and have linked the evolution of sociality to a transition from mass provisioning to progressive provisioning of brood. Early phylogenetic studies of allodapines were based on morphological and life-history data, but critical aspects of these studies relied on small character sets, where the polarity and coding of characters is problematic. We used nucleotide sequence data from one nuclear and two mitochondrial gene fragments to examine phylogenetic structure among nine allodapine genera. Our data set comprised Downloaded from https://academic.oup.com/sysbio/article/52/1/1/1656919 by guest on 03 October 2021 1,506 nucleotide positions, of which 402 were parsimony informative. Maximum parsimony,log determinant, and maximum likelihood analyses produced highly similar phylogenetic topologies, and all analyses indicated that the tropical African genus Macrogalea was the sister group to all other allodapines. This finding conflicts with that of previous studies, in which Compsomelissa + Halterapis formed the most basal group. Changing the basal node of the Allodapini has major consequences for understanding evolution in this tribe.
    [Show full text]
  • Woodward Et Al. 2011. Genes Involved in Convergent Evolution of Eusociality in Bees
    Genes involved in convergent evolution of eusociality in bees S. Hollis Woodarda,1, Brielle J. Fischmana,1, Aarti Venkatb, Matt E. Hudsonb, Kranthi Varalab, Sydney A. Cameronc, Andrew G. Clarkd, and Gene E. Robinsona,c,e,f,2 aProgram in Ecology, Evolution, and Conservation Biology, Departments of bCrop Sciences and cEntomology, eInstitute for Genomic Biology, and fNeuroscience Program, University of Illinois, Urbana, IL 61801; and dDepartment of Molecular Biology and Genetics, Cornell University, Ithaca, NY 14853 Contributed by Gene E. Robinson, March 12, 2011 (sent for review February 17, 2011) Eusociality has arisen independently at least 11 times in insects. different social lifestyles among relatively closely related species. Despite this convergence, there are striking differences among Furthermore, the extensive knowledge of bee natural history (8, eusocial lifestyles, ranging from species living in small colonies 13, 14) provides a valuable framework for developing hypotheses with overt conflict over reproduction to species in which colonies about the adaptive significance of genetic changes detected in contain hundreds of thousands of highly specialized sterile work- eusocial bee lineages. ers produced by one or a few queens. Although the evolution of To study patterns of molecular evolution associated with eusociality has been intensively studied, the genetic changes eusociality in bees, we generated ~1 Gbp of expressed sequence involved in the evolution of eusociality are relatively unknown. tags (ESTs) from a set of nine bee species (Table S1). This set of We examined patterns of molecular evolution across three in- species reflects the remarkable social diversity in bees by in- dependent origins of eusociality by sequencing transcriptomes of cluding eusocial and non-eusocial species; three origins of euso- nine socially diverse bee species and combining these data with ciality (9, 10); and two different forms of eusocial lifestyle, “highly genome sequence from the honey bee Apis mellifera to generate eusocial” and “primitively eusocial” (ref.
    [Show full text]
  • Kin Selection, Social Polymorphism, and Reproductive Allocation in Ants
    Katja Bargum Kin selection, social polymorphism, and reproductive allocation in ants Helsinki 2007 Kin selection, social polymorphism, and reproductive allocation in ants Katja Bargum Department of Biological and Environmental Sciences University of Helsinki Finland Academic dissertation To be presented, with the permission of the Faculty of Biosciences of the University of Helsinki, for public criticism in Auditorium 1041 of Biocenter 2, Viikinkaari 5, on March 30th, 2007, at 12 o´clock noon. Helsinki 2007 © Katja Bargum (chapters 0, II, IV) © Springer Publishing (chapter I) © Blackwell Publishing (chapters III, V) Cover illustration © Patrik Karell (2007) Layout © Katja Bargum Author´s address: Department of Biological and Environmental Sciences P.O. Box 65 (Viikinkaari 1) FI-00014 University of Helsinki Finland e-mail: [email protected] ISBN 978-952-92-1758-8 (paperback) ISBN 978-952-10-3795-5 (PDF) http://ethesis.helsinki.fi Helsinki University Printing House Helsinki 2007 _______________ 2 Kin selection, social polymorphism, and reproductive allocation in ants Katja Bargum This thesis is based on the following articles, which are referred to in the text by their Roman numerals: I. Bargum, K., Boomsma, J.J. & Sundström, L. 2004. A genetic component to size in queens of the ant Formica truncorum. – Behavioral Ecology and Sociobiology 57:9–16. II. Bargum, K. & Sundström, L. Colony-level life history trade-offs and reproductive skew in the ant Formica fusca. – Manuscript. III. Bargum, K. Helanterä, H. & Sundström, L. Genetic population structure, queen supersedure and social polymorphism in a social Hymenoptera. – Journal of Evolutionary Biology, in press. IV. Bargum, K. & Sundström, L. Multiple breeders, breeder shifts and inclusive fitness returns in an ant.
    [Show full text]
  • 299887753-Oa
    J Insect Behav (2017) 30:130–137 DOI 10.1007/s10905-017-9603-7 Sex Ratios in a Socially Parasitic Bee and Implications for Host-Parasite Interactions Nahid Shokri Bousjein1 & Morgan Staines1 & Cathy Vo1 & Natalie Puiu 1 & Carmen R. B. da Silva1,2 & Jack Harrington1 & Sara Wilkinson1 & Kelly Pratt1 & Michael P. Schwarz1 Revised: 27 January 2017 /Accepted: 30 January 2017 / Published online: 8 February 2017 # Springer Science+Business Media New York 2017 Abstract Obligate social parasites of Hymenoptera, known as inquilines, have re- ceived enormous attention due to the elaborate adaptations they exhibit for exploiting their hosts, and because they have frequently been used to infer sympatric speciation. Their population biology can be difficult to infer as they are both rare and difficult to extract from host nests. Sex allocation has been studied for very few inquilines of social Hymenoptera. Here we report sex ratio patterns in the allodapine bee Inquilina schwarzi, which is an obligate social parasite of another allodapine, Exoneura robusta. We show that the sex ratio of this inquiline varies with its brood number, it is female-biased in the smallest broods, but becomes more even in larger broods, where the population-wide sex ratio is close to parity. We argue that this pattern of bias is consistent with local resource competition, where inquiline females compete to inherit their natal colony. We also argue that extremely female-biased sex ratios of the host species, combined with overall sex ratio parity in the parasite, may help ameliorate disparity in effective population sizes between these two species which are locked in an evolutionary arms race.
    [Show full text]
  • Phylogenetics of Allodapine Bees: a Review of Social Evolution, Parasitism and Biogeography Simon M
    Phylogenetics of allodapine bees: a review of social evolution, parasitism and biogeography Simon M. Tierney, Jaclyn A. Smith, Luke Chenoweth, Michael P. Schwarz To cite this version: Simon M. Tierney, Jaclyn A. Smith, Luke Chenoweth, Michael P. Schwarz. Phylogenetics of allodapine bees: a review of social evolution, parasitism and biogeography. Apidologie, Springer Verlag, 2008, 39 (1), pp.3-15. hal-00891934 HAL Id: hal-00891934 https://hal.archives-ouvertes.fr/hal-00891934 Submitted on 1 Jan 2008 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie 39 (2008) 3–15 Available online at: c INRA/DIB-AGIB/ EDP Sciences, 2008 www.apidologie.org DOI: 10.1051/apido:2007045 Review article Phylogenetics of allodapine bees: a review of social evolution, parasitism and biogeography* Simon M. Tierney1,2, Jaclyn A. Smith1,LukeChenoweth1,MichaelP.Schwarz1 1 School of Biological Sciences, Flinders University, GPO Box 2100, Adelaide, S.A. 5001, Australia 2 Smithsonian Tropical Research Institute, Apartado Postal 0843-03092, Balboa, Republic of Panama Received 15 November 2006 – Revised 3 October 2007 – Accepted 3 October 2007 Abstract – It has been assumed that allodapine bees represent early stages in the evolution of social be- haviour.
    [Show full text]
  • The Ontogenetic Drivers of Behavioral Variation Across Groups
    REVIEW ARTICLE published: 10 December 2014 ECOLOGY AND EVOLUTION doi: 10.3389/fevo.2014.00081 The development of collective personality: the ontogenetic drivers of behavioral variation across groups Sarah E. Bengston 1* and Jennifer M. Jandt 2 1 Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA 2 Department of Ecology, Evolution and Organismal Biology, Iowa State University, Ames, Iowa, USA Edited by: For the past decade, the study of personality has become a topic on the frontier of Ann Valerie Hedrick, University of behavioral ecology. However, most studies have focused on exploring inter-individual California Davis, USA behavioral variation in solitary animals, and few account for the role that social interactions Reviewed by: may have on the development of an individual’s personality. Moreover, a social group Elva J. H. Robinson, University of York, UK may exhibit collective personality: an emergent behavioral phenotype displayed at the Marian Yi-ling Wong, University of group-level, which is not necessarily the sum or average of individual personalities within Wollongong, Australia that group. The social environment, in many cases, can determine group success, which *Correspondence: then influences the relative success of all the individuals in that group. In addition, Sarah E. Bengston, Department of group-level personality may itself evolve, subject to the same selection pressures as Ecology and Evolutionary Biology, University of Arizona, PO Box individual-level behavioral variation, when the group is a unit under selection. Therefore, 210088 Tucson, AZ 85721-0088, we reason that understanding how collective personalities emerge and change over USA time will be imperative to understanding individual- and group-level behavioral evolution.
    [Show full text]
  • Goulburn Broken, Victoria
    Biodiversity Summary for NRM Regions Species List What is the summary for and where does it come from? This list has been produced by the Department of Sustainability, Environment, Water, Population and Communities (SEWPC) for the Natural Resource Management Spatial Information System. The list was produced using the AustralianAustralian Natural Natural Heritage Heritage Assessment Assessment Tool Tool (ANHAT), which analyses data from a range of plant and animal surveys and collections from across Australia to automatically generate a report for each NRM region. Data sources (Appendix 2) include national and state herbaria, museums, state governments, CSIRO, Birds Australia and a range of surveys conducted by or for DEWHA. For each family of plant and animal covered by ANHAT (Appendix 1), this document gives the number of species in the country and how many of them are found in the region. It also identifies species listed as Vulnerable, Critically Endangered, Endangered or Conservation Dependent under the EPBC Act. A biodiversity summary for this region is also available. For more information please see: www.environment.gov.au/heritage/anhat/index.html Limitations • ANHAT currently contains information on the distribution of over 30,000 Australian taxa. This includes all mammals, birds, reptiles, frogs and fish, 137 families of vascular plants (over 15,000 species) and a range of invertebrate groups. Groups notnot yet yet covered covered in inANHAT ANHAT are notnot included included in in the the list. list. • The data used come from authoritative sources, but they are not perfect. All species names have been confirmed as valid species names, but it is not possible to confirm all species locations.
    [Show full text]
  • Canada Archives Canada Published Heritage Direction Du Branch Patrimoine De I'edition
    1 Nestmate Recognition in the Large Carpenter Bee, Xylocopa virginica by Marianne Peso, B.A.Sc. A Thesis submitted to the Department of Biological Sciences in partial fulfillment of the requirements for the degree of Master of Science Brock University St. Catharines, Ontario © Marianne Peso, 2008 Library and Bibliotheque et 1*1 Archives Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-46605-6 Our file Notre reference ISBN: 978-0-494-46605-6 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library permettant a la Bibliotheque et Archives and Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Plntemet, prefer, telecommunication or on the Internet, distribuer et vendre des theses partout dans loan, distribute and sell theses le monde, a des fins commerciales ou autres, worldwide, for commercial or non­ sur support microforme, papier, electronique commercial purposes, in microform, et/ou autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in et des droits moraux qui protege cette these. this thesis. Neither the thesis Ni la these ni des extraits substantiels de nor substantial extracts from it celle-ci ne doivent etre imprimes ou autrement may be printed or otherwise reproduits sans son autorisation.
    [Show full text]