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The Ecology of Lepsiella Vinosa (Mollusca: Gastropoda) With

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The Ecology of Lepsiella Vinosa (Mollusca: Gastropoda) With THE ECOLOGY OF LEPSIELLA VINOSA (MOLLUSCA, GASTROPODA) VIITH PARTICTTLAR REGARD TO ITS FEEDING BEHAVIOUR by DAVID E. BAYLISS B.SC. M.SC. (UNSW) DEPARTMENT OF ZOOLOGY THE IJNI\TERSTTY OF ADELAIDE A thesis submitted to the University of Adelaide in fulfilment of the requirements for the degree of Doctor of Philosophy. January 1979 Awo,red ìç t'Jov. lq,þ' TAT]LE OF CONTNNTS Page SUMMARY ¡ ACKNOWLEDGEMENTS lv LIST OF FIGURES vi CHAPTER 1. 1" 1 Introduction I L.2 Lítexatuxe 3 L.2.L Switching 4 Á L.2"2 Experimental studies I 1"2.3 Switching and stability 7 I.2.4.1 Searching image 8 L.2.4.2 Profitability of hunting L2 L.2.5 Searching image formation and the concept of L4 snitching L.2.6 Apostacy 15 L.2.7.1 Hollingrs type 3 functional response 1ó L.2.7.2 More recent studies on type 3 functional response L7 CIIAPTER 2 2.L Location of study area I9 2.2 General description of the intertidaL zone 19 CHAPTER 3 Some aspects of the prey species 3.L Introduction 23 3"2 Líterature and description of prey species 23 3"2.i Balanus amphitrite amphitrite Darwin 23 3.2.2 Elminius modestus Darwin 24 3"3 Development of ovaries 25 3.4 Settlement seasons 26 3.5 Size 27 3.6 Refuges 27 3"7 Competition 29 3.7.I Interspecific 29 3.7 .2 Intraspecific 29 Page 3.8 Discussion 30 CHAPTER 4 Some aspects of the feeding behaviour of Lepsiella vinosa 4.L fntroduction 35 4"2 Prey evaluation 35 4.3 Growth rate 36 4.4 Feeding rate of L. vinosa in barnacle patches 37 4.4.L Introduction 37 4"4.2 fnfluence of season 38 4.4.3 Influence of patch 39 4.4.4 Influence of prey abundance 40 CH.APTER 5 Dynamics of barnacle populations in pneumatophore Patches 5.1 Inland patches 4L 5.1.1 fntroduction 4L 5.L.2 Sampling procedur 42 5.1"3 Experimenter influence 43 5.L.4 Pneunatophore dynamics 46 - 5 .L.4.1 Methods 46 5.L.4.2 Results 46 5"1"5 Barnacle density 48 5"1.5.1 Introduction 48 5.1.5 "2 B. amphitrite den,sity 48 5.1"5.3 X" modestus density 49 5.1"5.4 Ratio dead to living B. amphitrite 49 5.1.5"5 Effect of f" vinosa predation onå. jgpåæ 50 survival 5.1.5.6 PotentiaL L. vinosa predation rate on B . arnphitrite 52 5"1.5.7 Effect of L. vinosa predation on E. modestus 53 5"1"5.8 Barnacr" f rom L6/3/75 to 6/2/76 53 "*ar"r** 5.1.5.9 Survival of B. amphitrite which settlecl prior to 54 16/3/7 s . Page 5.1.6.0 Relative importance of factors causing mortality 55 in the B. amPhitrite PoPulation" 5"2 Dynarnics of barnacle populations on a smal1 island, 58 subiect to L" vinosa and B- paivae predation 5.2.L Introduction 58 5.2.2 Characteristics unique to I patch 58 5.2.3 GastroPod Predation 59 5.2.4 PneumatoPhore dYnamics 60 5.2.5 B. amphitrite density and mortaLity 60 5.2.6 E. modestus density and mortality 62 5.3 Discussion 64 GIAPTER 6 Movenrent of L. vinosa in respect to barnacle patches 6.1 Introduction 66 6.2 Movement of marked individuals 67 6.2.L Methods 67 6.2.2 Re sults 67 ó.3 Movement of L. vinosa into and out of patches 69 6.3 .L Methods 69 6.3.2 Results 69 CHAPTER 7 Prey selection by L. vinosa undel natural conditions in the field. 7.t Introduction 74 7.2 lviethods 74 7.3 Predator response to changes in relative prey density 76 7"4 Influence of absolute dens-í ty of preferred prey species 78 on the percent age of L" vinosa eating that species. 78 7 "4.L High densities 7 .4.2 Moderate densities 80 7"5 Examination of possible mechanisrns by which the L. 81 vinosa population could adjust to changes in the-relative iñ-uun¿".t.e of barnacle species- 7"5"L Change in prey species selected by individual- 8r L" vinosa" Page 7.5.2 Ernigration as a possible factor influencing, 82 predation on the two PreY sPecies. 7"5.3 Immigration of new L. Y¿9.9-3-into the patches' 84 7.5"4 Possibility of different feeding rates causing 85 discrepancies in apparent prey selection 7.6 Discussion 86 CIIAPTER 8 Preliminary laboratory experimeirts on switching 8" 1 fntroduction 93 8.2 Experiments on switching using barnacles as prey 93 8.2.L Methods 93 8.2.2 Results 95 8.2.2.1 Traiirin g effect 95 8.2.2.2 Switching exPeriments 98 8.3 Training to herbivorous gastropods 100 8"3.1 Methods 100 8.3 .2 Results r01 8.4 Discussion 103 CHAPTER 9 Fíeld cage experíments on switching 9"1 Introduction 104 105 9 ^2 Methods 9.3 Results 110 9.3.L Switching exPeriments 110 9.3.1.1 L. vi-nosa trained to B. amphitrite 110 9.3.1.2 L. vinosa trained to E. mo,lestus 111 9.3"1.2.I. Prey not clumPed 111 9.3 .L.2.2 Prey clunPed 113 9.3.2 Reverse Switching experiments LL4 9.3.2"L L. vinosa trained to E. modestus LL4 9"3.2"2 L" vinosa trained to B" amPhitrite 115 9.3.3 Determination of "C" value 118 9.3.4 Controls L20 9 .3.5 Statistical analysis of evidence for switching L23 by L" vinosa in field cages Page 9.3.ó Feeding rate L25 9"3.7 Functional response and switching L27 9"3.8 Influence of absolute density on predation 130 9.4 Discussion L32 CFIAPTER 10 General discussion on the stability of the inter- L39 action between L. vínosa and barnacles REFERXNCES L4L Irretr x!¡9. 27 lln¡ ? for 5.F r¡rd 5.l0o 29 llnc eO for oo¡nrll (1961) r¡¡d Co¡¡oll (1961b) n li¡c , for Conncll (rg6r) ro¡d Oonnol.l (196rr) t I1¡c 17 for (Eurlcy 19?6) rcad (unrlry 1n5, ,tt ll¡c t for Co¡¡cLI (1961) ¡r¡û Go¡¡cL[ (fg61b) 66 rl¡c 1, for La¡drnbcrgas 6%?) rcad lr¡dcabcrgcr (1968) n llnc , for Ecl1r ¡onotle rcrd lcfl¡Î ro¡atla ?4 lfnc 11 lor con¡dl (rg6r) rerü GonncLI (1961r) 11? llno 26 for P ¡. O.24 rc¡d P - O.?8 îîa ltno eg for Pt0.255 rcad PeO.74 11t llnc 18 lon Pt¡0.085 ¡c¡ô P¡0.1O 119 ll¡c ¿? for ?-Oo79 r¡ad P*Oo7'l 118 Il¡c 7 for P=OoJ.1 rcrd P.0156 118 llne I for P=O.11 rr¡d PzOo29 1r1 llnc ö lor ¡¿-- o.oa8 rcad lt-- o.O28 Oheptcr t thcrr 1¡ ao Teblc 45 l_ SUMMARY Lepsielta vinosa is a muricid gastropod which is found in South Austratian mangroves where it eats two .species of barnacles, Balanus amphitrite and Elminius modestus, which are attached to the pneumatophores and branches of the mangrove Avicennia marina. The distribution of pneumatophores is heterogeneous and distinct prey patches are formed. A series of these patches lrlere surveyed to deÈermine the impact of L. vinosa predation on the barnacle populations a;rd the influence of barnacle density on prey selection by the predator. B. amphitrite settles between Spring and mid-sufltmer. It is Subject to fluctuations in reproductive success and t'failures" were recorded in 1975 and 1976. L. vinosa eliminated 9-33 to 30-8? of the heavy 1974 Spring settlement in one year. After three years only those B. amphitrite which settled in refuges survived. L. vinosa couÌd potentially elininate all B. amphitrite in the patches between successive breed.ing seasons. E. modestus can settle conÈinuously and unlike B. amphitrite its numbers recover quickly after a "catastrophe". In crowded situations it is crushed by B. amphitrite. Frequent restocking of the patches prevent its elimination by L. vinosa predation. prey selection by L. vinosa is influenced by both relative and absolute densities of the barnacle species. At moderate densities Iess than 25 barnacles/metre pneumatophore, the predator fed proportionally; L. vinosa feeding on the less abundant species changed to the more abundant species as its relative density increased. AÈ high densities, 50 to 70 barnacles,/metre pneumatophore, E. modestqq r¡ras progressivety dropped from the dietcsr the absolute density increased. I]- Laboratory experiments indicate that L. vinosa can be trained to either barnacle species and shows switching in terms of Murdochrs (1969) model. Field cage experimenÈs indicate that L. vinosa shows switching provided that the absolute densiÈy of B . amphitrite is not too high. The switch tends to become stronger over an eight week period. In both the taboratory and field experiments L. vinosa had a weak preference for B. amphitrite, c = L.26, when equal numbers of each species were present. The preference appears to be consistent between individual L. vinosa. In the field L. vínosa switches to B. amphitrite when it is more abundant irrespective of prior training. L. vinosa trained to E. modestus or unfed switched to E. mod.estus when it was more abundant. Those trained to B. amphitrite however only slowly switched to E. modestus over a period of eight. weeks. A clumped prey distribution accelerated the switch. L. vinosa also showed aslzmmetry in feeding rate.
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