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In South Australia WAITE INSTIÏUTE LIBRARY TT{E ECOLOGY AND LIFF] HISTORY OF Ttr INTRODUCED MILLIPEDE, ArnnatoiuLus moveLetii (LUCAS, 1860) ' IN SOUTH AUSTRAI.IA. By GEOFFREY H. BAIGR B.Sc. Hons; (Adelaide) A thesis subrnitted for the Degree of Doctor of Philosophy in the Iraculty of Agricultural Science at the Llniversity of Adelaide. Departrnent of Frrtonology, Waite Agricultural Research fnstitute, university, of Adelaide . J.ure 1-976. ]. I'ABLE OF CONTENTS Pqge TABLE OF CONIENIS i SI.Nß{ARY i.ii DECLARATION vi ACKNOWLEDGUVIENTS vii 1 1. INTRODUCTION 2. DISTRIBUTION AND DISPERSAL J 2.r Ilistribution 3 2.2 Dispersal 6 9 3. POST-E\,{BRYONTC DEVELOPMENT 3.1 General 9 3.2 I 3.3 11 3.4 v, 3.5 I4 3.6 16 5./ Is 0. moreletii semelparous or iteroparous? 18 z4 3.8 Fec-un 3.9 Post- of A. castaneum 26 3.1 0 Post-Ernbryonic Develo of l_lqqlggg""u sp. 27 30 4 LIFE HISTORY, ABUMANCE A¡{D ACTN¡]TY 4.L Introduction 30 4.2 Literature Review 30 4.21 ]r{aturation in Iuliform N{illipedes 30 4.22 Seasonality of lr{ature and Intercalary tr4a1es in 33 Periodonorphic Species 4,23 Activity 34 I 36 4.3 Habitat and Weather ,\ 4.4 Methocls of Sampling for Life History, Abundance and Activity 39 4.4I Sampling of Soil and Litter j¡ the Grassland 39 4.42 Sampling of Litter by Itself in the Grassland and Woodland 42 4.43 Pitfall trapping in the grassland and woodland 43 4,44 Visual Assessment of Activity and l\4ating 4s 4.45 Fland-Collections and N'iiscellaneous Samples 46 4.s Results of Sampling for Life History, Abundance and Activity 46 4.51 Mating 46 4.52 Age Distributions 46 4.53 ltlaturation in Fenales 48 ii. Page 4.54 Maturation in N{ales 49 4.55 Sex Ratios 5Z 4. 56 Changes in Abr-rrdance Throughout Surrner s4 4,57 N{aturation and Subsequent Survival 57 4.58 Production of New Generations 62 4. 59 Actir¡ity 6.5 4.510 Life Flistories of the Native Species, Australiosoma castaneun and llfunerogonus sp. 6B 4.6 tJiscussîñ-- 70 5. WATER RET,ATIONSTilPS 77 5.1 Introduction 77 5.2 Literature Review 77 5.3 Resistance to Desiccation at Moderate Temperature (20'C) 79 5.4 Resistance to Desiccation at High Temperature (35'C) by O. noreletii 81 5.5 füniõ'íFEferences of 0. moreletii 85 5.6 Desiccation in Field Popülatiõñ3-õT-0. noreletii 87 5.7 Discussion 93 5.8 Sumnary 96 6. TEUPERATURE REI."ATIONSHIPS 97 6.1 fntroduction 97 6.2 Literature Review 97 6.3 Resistance to high Temperature 98 6.4 Temperature Preference s of 0. noreletiI 100 6.5 Surnnary 102 7. DISCUSSION 103 8. APPEMICES 9. BIBLIOGRAP}IY 1lr SLT\O/ÁRY The inil l-ipecle, Ommatoir"rl.us noreletii. (lulidae), is indigenous to South-Western Europe. The spec.ies has been intr:ocluced to Australia where it is in large nurnbers and constitutes a SeVeTe nuisance. The clistribution of O. moreletii in Australia is increasirig. Man is a rnajor factor in the establishnent of new outtrreaks of the species. Individual outbreaks spread slowly but steadily (an expansion of up to 200 metres in radius Per Year). The abr¡rdance of O. moreletii in Australia relative to its apparent rar:ity in South-l{estern Europe caffIot be e>çlained by differences in general clinate. The variety of habitats colonizedbT'cl. 1lgryþlii in Australia is already wide an<1 there is no ïeason to doubt t.hat the species wi.ll continue to spread nmch further. The life cycle of o. noreletii consists of an egg, pupoid and t¡en a seri.es cf up to sixteen stadj-a. lvfales may rrature at a:ry moult fron the eighth to the twelfth stadia, but nost are mature by the tenth or eleventh. Maturation is more difficult to determine in felnales but seens sinilar to the males with respect to stadial age. 9. ryrylçlt1 therefore has rnore than one adult stadir¡n. O. moreletii j.s periodomorphic. That is, there are nature and "intercalary" (inrnature) f-orms of the adult male. 9. rnoreletii is probably capable of iteroparity (producing moie than one brood). Iteroparity is however consicleted a rare occurrence in the field. Nests laid in the laboratory contain an average of 250 eggs' Field stuclies were made of O. noreletii in an open grasslancl and a sclerophyllous woodland. Surpfirg of soil and litter collectively, litter by itself, pitfall trapping and visual assessment of activity gave data on the life history, abrndance and activity of O. rnoigletii. IV. Females of O. nroreletii rnatured their eggs in late sunrner - auturn' They rnated ancl oviposi'Eed during autum - winter. Aft-er 1year, inrlividuals were i¡ the seventh, eighth or ninth staclirmr" After 2 yeats, the tenth or elevcnth stadia v¡ere reached. In subsequent year:s' 0. lgt"l"tli probably moulted twice T,er year (in spring and srlrnner) . In mal-es, the sunner noult rvas to the mature form (either fron the juvenile or intercalary\; the spring moult rvas to the intercalary fonn (from tTre mature) " Thus intercalary males were present in spring - sunmer. For the remainder oi the year:, the adults were mature. Maturation of adults was higher after â tr¡êt sulmer than after ü.vo dry sunrners. lvfaturation was ntore successful in the grassland than in the woodland. Production of young was nuch greater in the ggassland. Sgrvival of females from the autr¡rm breeding season to the subsequent spring was correlated (negatively) with their naturity in autunn. The advantage to O. moreletii of extended adult life in fenales after a poor breeding season is discussed with reference to Den Boerrs (1968) t"o concept of "spreacling of risk". The advantage of periodomorphosis this species is also discussed. Active 0. moreletii were rnost conrnonly seen and trapped in auturn. Day to day variations in activity were correlated with changes in tenperatutre and moisture. Older stadia were more active than yor.nger stadia. D-rring sutnner, O. morele-li:L aggregated in cool, moist sites (e.g. beneath tussocks cf Lornandra _tlÞæ. (Liliaceae)) or burrowed urrder- gror-rrd. Behaviour of the species in gradients of temperature and hunidity e4plained the aggregation j-n surnner beneath the tussocks. In a particularly hot, dry sunner, mortality was demonstrated in the grassland but not i¡ the cooler, moister woodland. A survey of the percentage water contents ir 9. no.reletti during this particular sunmer gave no evidence of desiccation of the species in either habitat. This suggesteC that high temperature \,\ras the funportant mortality factor. Flowever, using a tr:chnio,ue inr¡oh'ing the appearance of the mid-gut, desiccation of 9. moreletii was clemonstrated in a later, milder sunrner and cloubt was placed upon the earl.ier result. O. moreletii developed resistance to high tenperattrre and to desiccation at high tenrperature throughout sunrner. lvlature males were less resistant to high temperatures and desiccat.ion than females and juvenile ma1es. Indirect evidence suggested that mature males were less resistant to desiccation at high temperature than intercalary ma1es. T\,vo native speci.es of millipede, Dimerogongr sp. (Carnbal.idae) and Australiosoma castaneun (Paradoxosomatidae), were for¡:d in the r,rtoodla¡d but not in the grassland. The native species were rare relative to O. rnoreletii. They probably bred in late winter - spring. Juveniles of both native species h¡ere more susceptible to desiccation and high temperature than juveniles of O. Iglgle-lii. This is suggested as one reasolr for the differencers -'r-n distribution and abr¡rdance of the three species. Declaration The work presented in this thesis is nry ourt unless otherwise acknowledged, and has not previously been published or submitted to any university for the arvard of any degree. June 1976. a i vll. i ACK}.IOhII,EDGI$MNTS professor Ton y,]hi'Ee (now at the Uni-versity of the South Pacific) was first to recognize "the nillipede" as a profitable avenue of ecological research. Tom stimulated me as an undergraduate student to undertake thís present stucly. I thank him for many long discussions and his continued ínterest in rny rvork. I thank Dr. Roger Laughlin ancl Professor T.0. Brov¡ning for their advice throughout the studY. I{r. Bill King kindly allorved me the use of his property at Bridgewater. The National Trust of South Australia allowed me the use of Ergelbrook Reserve. Ron Vincent, Bob and "Tobe" Armstrong of Bridgewater information and help in the field-work, as well as were invaluable for t R.E. Oswald of the Stirling li "refreshmentil afterwards. Colin Wirth and Mr. ilr T. District Cotncil, Bill Anderson of the Keilor District Council, members of the Victorian Department of Agriculture and Nleville l{anklyn of the "Port Lincoln Times" províded welcome assistance in my surveys of the d.istríbution of O. noreletii in Australia. Dr. C.A.W. Jeekel (Amsterdam) identified O. rnoreletii, Djmerogonus- sp. and {. casta¡eum for me. Dr. David Symon (Waite Institute) iclentified the plants at Bridgewater. Drs. J.M. Demange and J.P. l4auriès (Paris) provided the available information on g. rnoreletii in Europe. Dr. David Lee (South Australian lrfuser:n-r) iàentified FI. feroniartrn. Professor Carroll l¡li11iams (Flaruard) assayed O_. noreletii for jguenile hormone. Dr. Robin Bedding (C.S.I.R.0., Tasmania) assayed O. rnoreletii rvith Negaplecjlg4e. Carol Crawford (lVaite Institute) provided urpublished data on soil moistures. l4arjorie and Robert Baker helped with t-he thesist diagrarns. I sincerely thank all these people. Finally, I thank Nicole.
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