GEOMETRICMORPHOMETRICSONSNAILSHELLS WarpedIdeas:GeometricMorphometricsasaComplementary TechniqueforStudyingGastropodShellMorphology ByMariamYousif,B.Sc. AThesisSubmittedtotheSchoolofGraduateStudiesinPartialFulfilmentofthe Requirementsfor theDegreeMasterofScience McMasterUniversity, ©CopyrightbyMariamYousif,July2012 McMasterUniversityMASTEROFSCIENCE(2012)Hamilton,Ontario(Biology) TITLE:WarpedIdeas:GeometricMorphometricsasaComplementaryTechniqueforStudying GastropodShellMorphologyAUTHOR:MariamYousif,B.Sc.(McMasterUniversity) SUPERVISOR:ProfessorJon StoneNUMBEROFPAGES:X,74 ii ABSTRACT Geometric morphometrics (GM) provides a complementary method for studying morphology.Snailshavebeenanalyzedinthefieldofmorphometricssincethe1960sbecause theirshellsservetorecordinformationabouttheirlifehistoriesandenvironmentalhabitats.In thisthesis,wepresentanannotatedbibliographyforadvancementsinGM,usingapplicationsto snail shells as a representative case study. We categorize 30 publications into four fields, morphology,ecology,taxonomyandevolution,andshowthatdevelopmentshavebeenunequal among them. We conclude by predicting that GM applications on snail shells will increase, especiallyinhybridfields,suchas,ecotoxicology,whichcurrentlyareunderrepresented.Asa demonstration, we describe an experiment wherein we applied GM as a complementary morphologicalmethodtostudythegardensnailspecies Cepeanemoralis inanorganophosphate- pesticide, ecotoxicological setting. We conclude by showing that GM reveals subtle morphologicaldifferencesamongtreatmentgroupswithnorelationtopesticidedosage. iii ACKNOWLEDGEMENTS First,IwouldliketothankmysupervisorDr.Jon Stone for his endless support and encouragement.Hiswillingnesstoguidemethroughthisprojectandhisgenerosityhasenabled metoenjoyandsuccessfullycompletethisthesis.Iwouldliketoalsothankmyothercommittee members:Dr.ChrisWoodandDr.MichaelO’Donnellfortheirsuggestivefeedback. Ialsothankthestudentvolunteerswhomparticipatedinrearingthespecimensduringthe study. Further thanks go to Cheminova for their generous donation of the chemical, Cygon (dimethoate).Finally,Iwanttothankmyfamilyfortheirongoingencouragementandsupport. iv TABLEOFCONTENTS TitlePage-------------------------------------------------------------------------------------------------------i DescriptiveNote-----------------------------------------------------------------------------------------------ii ABSTRACT---------------------------------------------------------------------------------------------------iii ACKNOWLEDGEMENTS---------------------------------------------------------------------------------iv TABLEOFCONTENTS----------------------------------------------------------------------------------vii LISTOFFIGURES&TABLES------------------------------------------------------------------------viii LISTOFABBREVIATIONS------------------------------------------------------------------------------ix DECLARATIONOFACADEMICACHIEVE----------------------------------------------------------x 1.0CHAPTERONE-----------------------------------------------------------------------------------------1 1.1INTRODUCTION--------------------------------------------------------------------------------2 2.0ADVANCEMENTSINGEOMETRICMORPHOMETRICSUSINGSNAILSHELLS ASACASESTUDY--------------------------------------------------------------------------------------4 2.1ABSTRACT-----------------------------------------------------------------------------------------------5 2.2INTRODUCTION----------------------------------------------------------------------------------------6 2.2.1Abriefhistoryofconchology-------------------------------------------------- ------- ---- ---7 2.2.2Traditionalmultivariatemorphometrics---------------------------------------------------8 2.2.3Geometricmorphometricsrevolution----------------------------------------- --------- ----8 2.2.4Outlineapproach------------------------------------------------------------ ----- ----- --- -----9 2.2.5Landmarkapproach-------------------------------------------------------- ----- ------ --- -----9 2.2.6Advancesingeometricmorphometrics----------------------------------- ----- ---- --- -----10 2.2.7Shellmorphologyadvantages----------------------------------------------- ------ --- --- ---11 2.2.8Shellstructure&production-------------------------------------------------- --------- ----12 2.3GEOMETRICMORPHOMETRICSONSNAILS-------------------------------------------------13 2.3.1Morphology-----------------------------------------------------------------------------------13 2.3.2Ecology----------------------------------------------------------------------------------------21 2.3.3Taxonomy-------------------------------------------------------------------------------------26 2.3.4Evolution--------------------------------------------------------------------------------------28 2.4CONCLUSION-----------------------------------------------------------------------------------------32 2.5REFERENCES------------------------------------------------------------------------------------------34 v 2.6APPENDIXI:TERMSANDCONCEPTS---------------------------------------------------------39 2.7APPENDIXII:FIGURES-----------------------------------------------------------------------------40 3.0GEOMETRICMORPHOMETRICSAPPLIEDTOGARDENSNAILCEPEA NEMORALIS ASAMORPHOLOGICALMETHODIMPLEMENTEDIN ANECOTOXICOLOGICALEXPERIMENT---------------------------------------------------41 3.1ABSTRACT---------------------------------------------------------------------------------------------42 3.2INTRODUCTION--------------------------------------------------------------------------------------43 3.2.1Dimethoate------------------------------------------------------------------------------------44 3.2.2Geometricmorphometrics------------------------------------------------------------------45 3.3MATERIALSANDMETHODS---------------------------------------------------------------------49 3.3.1Chemicals-------------------------------------------------------------------------------------49 3.3.2Exposureconditions-------------------------------------------------------------------------49 3.3.3Foodpreparation-----------------------------------------------------------------------------50 3.3.4Chronictoxicity-------------------------------------------------------------------------------50 3.3.5Reproductiveassessment--------------------------------------------------------------------51 3.3.6Foodintake-----------------------------------------------------------------------------------51 3.3.7MeasurementofAChEactivity------------------------------------------------------------51 3.3.8Growthassessment--------------------------------------------------------------------------52 3.3.9Bivariatestatisticalanalysis---------------------------------------------------------------53 3.3.10Principalcomponentsanalysis-----------------------------------------------------------53 3.3.11Geometricmorphometrics-----------------------------------------------------------------53 3.4RESULTS-----------------------------------------------------------------------------------------------54 3.4.1Survivorship----------------------------------------------------------------------------------54 3.4.2Foodintake-----------------------------------------------------------------------------------54 3.4.3Chronictoxicityandclinicalsigns---------------------------------------------------------56 3.4.4VariationinAChEActivity-----------------------------------------------------------------55 3.4.5Growthassessment--------------------------------------------------------------------------55 3.4.6Principalcomponentsanalysis------------------------------------------------------------56 3.4.7Relativewarpanalysis----------------------------------------------------------------------56 3.5DISCUSSION-------------------------------------------------------------------------------------------57 3.5.1Survivorship----------------------------------------------------------------------------------57 vi 3.5.2Foodintake-----------------------------------------------------------------------------------57 3.5.3AChEactivitymeasurement---------------------------------------------------------------58 3.5.4Growthinhibition---------------------------------------------------------------------------58 3.5.5Geometricmorphometricsasacomplimentarymorphometrictool-----------------61 3.5.6Impactofmorphologicalchange---------------------------------------------------------63 3.5.7Cepeanemoralisvs.Helixaspersa-------------------------------------------------------64 3.6REFERENCES----------------------------------------------------------------------------------------65 3.7APPENDIXI:Tables-----------------------------------------------------------------------------------69 3.8APPENDIXII:Figures---------------------------------------------------------------------------------70 4.0CHAPTERFOUR------------------------------------------------------------------------------------73 4.1CONCLUSION---------------------------------------------------------------------------------74 vii LISTOFFIGURES 2.2.7Geometricmorphoemtricspapersfrom1980-2010----------------------------------------------40 2.2.9Thin-platesplinerepresentingresearchdistributionfor30publications---------------------40 3.3.10ImageofaCepeanemoralissnailshellwithlabeledtraditionalvariablesandlandmark points-------------------------------------------------------------------------------------------------70