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The Food of the Terrestrial Snail Helicella Virgata (Da Costa) In

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The Food of the Terrestrial Snail Helicella Virgata (Da Costa) In rHE FOOD U' TNE TENNH¡TN,IAT SI{AII, Eelicella virsetg (ila costa) ff SOTITE AIFTN¿LT¡ by .AIÁN JOEN BINIER B.Sc. (nons. ) DEP¡BS{ENT OF UOOI&Ir TFE T'NIVEASITY ü' AÐEIJAII)E À tbesis subnítteil to the Universíty of Adehide in part fulfilænt of the requireænte for the degree of Doctor of philosophy uaY 1972 TABI,E OF CONTENTS Page stttÍMAllr i DMIjNATIOIü iii ACI0IIOWIEDGEMM\TTS iv I. I}ITRODTETION I 2. DIET OF ffiLICELT,A VIRGATA IN SOTNH .ATISTRALIA 5 2.I INIRODIroIIG{ 6 2.2 FIETD STIR\IEY OF GTJT4OI{TENTS AT NORTIIT'IEI,D & BiTKTAND PAAI( I 3. DIETÂNY lã 3.I INTBODTICTION tõ 3.2 Îffi EFFECî OF 'CI]LTTIB,INGI THE MICBO-ORGANISII,S IN LITIts,R ON TI{E FOOÈVAH]E OF' fiIE LITTER 20 3.2. I First I culturing I erperinent 20 9.2.2 Second rculturingr experiment 3l 3.2.3 DÍscussion of tbe tculturingr experinents 36 3.3 TUE GRüTTE OF' S}TATI^S ON A VAAtrEIY OF POTMUIAL FOODS 88 3.4 TfiE GBtrTH OF S}IA.IIS FED ON DRIED PASTT]RE PTANTS HARVESTED IN DIEFEAÐ.¡Î STÂGES G' ìúATURITY 4L 3.4.I Pilot erperiment on dried lettuce 4t cs{ [BnF (co{'rrNrßp) Page 9.4,2 Fíret d¡ied-for¿ge e{p€risent 4l 3.4.3 Second dried-forage erperinent 49 8.4.4 Third dried-f orage experiment ã3 3.5 DISCUSSIOT¡ OF DIEIARY mQUIREMm.ITS õ9 4. FACTOR^S DBT&MINING WEETIIER SIüAII,S WILL EÀÎ SSATå,IN TUTNGS 6l 4.I INTRODÌrcITON 6l 4.2 1TIE EYPCIIHESIS TIIAT S.IAII,S FMD INDISCRIMIMTELY T]NI,ESS INETBITED 66 4.2.L The effect of variations in the texture of foorl 06 4.2.2. The effect of varying nutritiousnees rithout varying texture 68 4.2.3 Discusgion 7l 4.3 WHAT CEMTICAI^S STIMTTI,ATE FEDING? 72 4.3.1 Erperiments using the tspeed-of-leavingr tecbnique 72 4.3,2 Ân experinent using the ttwo-stripst technique T6 4.3.3 The effect of glutathione on the palatability of filter paper 8¿ 4,3.4 Erperiurents oa extracts of pasture plants 84 4.3.õ The effect of oil, peptides and carbohydrates 9I o0È{TENIS (Cü¡ÎINUEp) Page 4.3.6 Diecussion of the etinuli for feeiling 96 4.4 WEAT INHIBITS TEÐING? 98 4.4¡I Palatabílity of en êcid-€rtract of early-harveeted forage oats 98 4.4.2 &perinents with a plant alkaloitl 100 4.4.9 The toughness of grasses 108 4.4.4 Discr¡ssion of the inbibition of feeiling It6 5. MOrr,EtlENT OF SI.IAII.S WITH RtsSPBCf T0 F00D 119 ã.I INIAODæTION 119 5.2 ATTAACTION AT A DISTANCE 12s 6.2.L Technique for studying the behaviour of Helicella in Y-tubes t2g 6.2.2 Attraction to crushed sn¿ils - I pheromone? I27 5.2.3 Attraction to foods t80 6.3 DISCUSSION 148 6. CONDITIONS UNDm' WHICH SI{AII.S CANI TEßD 150 6.I THE STIMÜI,I WHICII AAOUSE SNAIIS TH,OM D.IACTTVIIY 150 6.2 IS DORIIANCY OBLIGATOfiT D(AING SIJ}Sftr¿? 154 c6[rnNrs -(cmrrnnmo) hge 7. IEE AUÎìÍDANCE @ F00I) 160 ?.1 DlrnounTro{ r00 7.2 CRffüDING UImIilBn - NORIEFIEIÐ 108 7.3 CRWDING ÐTPERIUEbN - HDKLAND PÁNK IE? 7.4 DISCÌISSION 208 !!BPI@ 1. The choice and D€asr¡rement of depentlent variables in feeding-treils anaL crording erperimenta 2tL 2. Statietical sethods and oonvention¡ for prerenttog re¡ults 221 8. Study-oreae 2¿1 4 Keepí4g 6nÀi1¡ in the laboratory q27 230 SilMARY i. the terreetrial snêil Belicella vireatå (da costa) *u" introduoed fron Errrope into South .Australia vhere ít is now vide- spreêd., ft feedg nainly on decayí",g plqnt naterialr altbough gut- contents indicate that green tissues of vaecular plante do fom a ¡nrt of its rliet. Ex¡rcrinents were conducüed to deter¡rine rhether there i¡ a specific dietary reguireænt which cen only be gatisfied by eating rlitterr. It is concluded. that this is unlikely, Certaín observations sr¡ggested that H. virgeta may feed indiscrininately r¡nless inbibited by physiaal or chenical properties of the food - thet ie, that it doeg not respond to positive chenical etimrli to feeding. Thie h¡rotheaie is rejected; so too is the h¡lotheeis that it responds to one specific stisulue. No atterrpt waa ¡rade to teet a conpreheneive list of chenicalg, but subetanceg shovn to sti¡ltr1ate feeding inclutle glutathíone, glucoae anil a vegetable oil, EVítlence wae obtained that H, vírgata is inhibited from feeding on certain plants not only by the toughneaa of their outer rayere but alao by chenical conponente of the plants. A p¡rrrolitidiae alkaloid was ehoûn to be inhibitory. However, toughness nay also be an inportant barrier to feeding. Evideuce is presented, thet thig ís ço rith at least two s¡rccies of grasães. The posaible role of gestropoile in the evolution of such protective devlces in plants is briefly discussed. ii, U_eirg y-tubea in the laboratory it rae eonfirned that f,. virgata ie.¿ttracted by the odour of.a crushed nrcmber of lte species. Haying fotrnd a crusheil suail, the live ones feed on it. there ie nq evidence of the presence of a phero.Eone and it ia tentatively accepted. that the snails are attracted by an od.our aesociated with food. rn agreement wíth this, there is evidence of attraetÍon to several other foods. The stinuli rhich arouse snôÍls frou inactivity are briefly iliseussed. Evidence is presented which confirms the earlier findÍng that donnaney is not obligatory Ín su¡mer and tbat an¡lrg briefly awakened from suruner dormancy êre able to feed. nro field experiments were eonducted to ínvestigate the likelihood of snails erperiencing e shortage of food. fn both experinente and in one by a previous worker, various !ûeaarrree of grorth, reproduotion, activity or survivel rere found to ilecreaee rith increasing density of sn¿ils. rt is suggested th¿t theee results app€ar ineonsistent with the h¡ryotheeis of an absolute shorüage of completely-accessible food. Tbe resurts could be explained by a nodificetion of that hlryothesis or by the h¡aothesis that sneíls at high tleneity interfere yith each other by neans other than competition for food. The second experinrent garze results suggeeting that, on the area etudied, the food naturalry an¿il¿ble Ís not of high quality¡ the sneils DÊy suffer a relative shortage of food, of the kind in which fooil is abundant but nuch of lt ie low in nutrients. iii ;\ DECIANATIO{ This tbesis contains no naterial rhich has been accepted for tbe award of ony other degree or ttÍplooa in any r¡niversity and, to the best of nryr knorledge anrl helief, contains no material previously published or çritten by enother person, except rrhen due reference ie nÞtle in the text of the thesis. rv ÂCIû\IOWTEDGEMENTS f should like to €xpress rry gratitude to ¡¡y supervisore, hof. H. G. Andrewartha end Dr. S. J. Ednonds, for tbeir advice and. encouragernent throughout the course of this study. I e,m grateful to ltr. N. Brooks of Buckland Park for perrnissíon to çork on hís property and to the Director of the Departuent of Âgriculture in South tu'etralia for perníesion to use the study-area at Northfield, f should like to thank Dr" S. Niveu for statietical advice and Messrs. P. D. and P. G. Kenpster for valuable technical assistance. f ras very fortunate to have the help of ì*lrs. D. J. l'{acpherson who t¡ryed. the thesis. Finally, I an nost grateful to uy wife, Jan, who is long-suffering and helped in laany ways. (:- I ai 1. The helicÍd gnail Helicella virgeta (da Coeta) ras introduced ínto south Auetralia from Europe early in this century and is nfr wideepread throughout agricultu¡al areas of the state. Pomeroy and Lews (196?) considereil that the activities of man heve prayed a great part in the spread of the snail antl that its spreail ie probably stitl continuing. Althoqgh II. v ta is r&re over much of its range it can reaoh deneities of over 200 per square metre in some placee (pomeroy, 1966). ït is especially numerous in agricultural areas, along roadsides EDd the borders of paddocks; relatively few are found arnongst native vegetetion. During the hot, dry South Australian surmer tbe enails spend long periode dornÊnt, many of then attaehed high on plants, fsnss-posts end other upright objects. Ifost of then êre young aninars that hatched from eggs during the prececling winter, but some ere aestivating for their second or, rarely, third sumerr Because of this behaviour II' virgata sometimes beçomeg a pest in the croplands; lerge numbers of snaile dornent on the cereal-starks rnay be incruded in the grain when it is harr€sted. Most of south Australiars rain can be erpected to farr between March and October or November. Temperatures during the winter are mild and at this tine H. virggta wilt be active on most nighte and many of the days. Adult snails which heve overeì¡nmered will feed., grow relatively little and shortly begin to lay eggs. Reproduction can take plece over a long period, some young hatching as late as 2. early 0ctober, but nost in April4{ey. The young grow quickly and nost of then weigh 250-400 llg by October-Novernber (Poneroy, 1969). Poneroy (tooo, 196?, 1968, 1969) undertook a study of the ecology of H" vírgata in South Australia. He exanined a number of aspecte of the generel biology of the aninal and concentrated particularly on dorr'rancy.
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