Corella.1991, 15\2t: 45-52

Au.enldsiun Rird Reviews - Number 3

THE SHORT-TAILEDSHEARWATER: A REVIEWOF ITS BIOLOGY

IRYNEJ SKIRA

Dcpann1entof Parks,Wildlife and Heriragc, 13,{ Macquarie St.. Hobarr-Tas 7000

ReceivedI May 1990

The lile history of the Shorl-tailedSheaMaler tenuirostrishas been well documenled since it firsl came to lhe attentionof naturalists.Approximately 23 million breed in about 250 colonies in soLrtheasternAustralia trom Septemberto April. The Short-tailedShearwatef commences to breedwhen 4 to 15 years.During their completed tiletimes 27 per centoI all individualsproduce no youngand 19 per cent only one chick.Mortality is age-relatedwith lhe mediansurvival time for breedingbeing 9.3 years after first breeding.Many areas remainopen for study,with a particularneed for interdisciplinaryresearch that includesoceanography.

INTRODUCTION The Short-tailedShearwater was one of the lirst Australian birds to be banded in large numbers The Short-tailed Shcarwater P uffinut tenuirostris, (Serventy 1957, 1961) and to be subjected commonly known as the Tasmanian muttonbird, to a long-term scientiRcstudy (Guiler e/.r/. 1958). is one of about 100 in the Procellarii- This studv was commenccd on Fisher lsland in formes. The diagnosticfeature of the order is the the Futncaux Group of Tasmania in March 1947 by cxternal nostrils produccd into tubes extending Dominic Serventy lormcrly of the CSIRO onto the bill. Other distinctive features are the (Serventy 1977) and continuesto thc present, 42 hooked and plated bill, and the glandular part of years later. Due to the long-term nature of the the stomach which is greatly extended and pro- study together with the banding of some 92 00t) duces the well known'oil', actuallywax esters birds in Australia, the life history of the (Warham 1977).Most membersin this order Short- have tailed :t distinctivc musky odour. is one of the bcst documented in the world 'I'he of any (Bradley eral. 1989,1990: familv is the most diversc Seryenty 1974;Serventy and Curry 1984;Wooller group in the order and contains 61 spccics of et at. 1988. 1989. 1990). pctrels and shcarwaters (Serventy et al. 1971). The majority are nocturnal and nest in holes, EVOLUTION burrows ot crevices.which serve to protecl them and their young from predators. The gcnus The Procellariiformesare an ancientgroup of Puffnas consistsof 15 medium-sizedspecics that birdsthat probablyoriginated from aquaticbirds are among thc world's most numerous . presentat the end of the Cretaccous,some 64 Thcir high ncsting densitiesand their fidelity to a million years ago. The phylogenetichistory of particular site has meant that thcy are highly shearwatersis little known (Kuroda 1954). vulnerable to exploitation (Seryenty et al. 197l). Accordingto Olson (1985p.211), 'mosr of the The harvcsting of Short-tailed Shearwalers, or modernspecies-gror.rps, or subgencra, of Puffinus muttonbirding as it is known in Tasmania, is onc were in existenccby the Middle Miocenc. and of the bcst documented instanccs of there has bcen very little morphologicalchange harvesting(Skira 1987, 1990). within theselineages in 15 million yearsor so'. I SklfarThe biologyofthe Short-lailedShearwater Corella15(2) P. tenuirostris has not been found in any and Norman 1981), South Australia 600 000 deposits apart from pre-historic archaeological burrows in 33 colonies (A. C. Robinson, pers. srtes(Bowdlcr 1984;Friedman 1934a and b, 1941; comm.), New South 25 700 breeding pairs Vanderwal and Horton 19{34).One ot the fossil in 13 colonics (Lane 1979)and Western Australia procellariids, P. c.onradi from the Middle 10 000 burrows in several colonies (Johnstoneet Mioccne of Maryland, USA. seemsto have lived al. 7990aand b; I-ane 1983). It is estimated that to thr: Pleistocene, one ntillion years ago. p. 23 rnillion birds breed in about 250 colonies. lenuirostrismay have evolvcd from a P. conradi- Recent rescarch by Japanesebiologists has con- typc ancestor through the P. inceptor line centratcd on mapping vegetationtypes in sevcral (Kuroda 1954). P. inceptor is known from the colonies in Tasmania (Kuroda 1986). Middle Miocene of California suggesting that P. tenuirostris and its congener thc Sooty Shcarwater P. griseus which is abundant in MIGRATION Ncw Zealancl. were diflerentiated in the North Atlantic-North America areas. Having settled in Thc Short-tailedShcarwater is a circum-Pacific the Southern Hemisphcrc. perhaps as late as migrant spending the boreal sunmer in the the Plcistocene. their postbreeding migration N()rthern Pacific region (Fig. l). Sexually nriry be an instinctivc responseto return to thc immature birds depart from Australia near thc Northern Hemispherc (Kuroda 1954; Marshall cnd of March, followed by the brecding birds and Scrventy 19-56). around mid-April, with the flcdged chicks leaving at the end of April to early May. Thc l'hc firssil record left by seabirdsin Australia migratc rapidly (Serventy 1956)and arrive in the rs meagrc compilrcd to that of the Northern Northern Hcmisphere on a broad front acrossthc Hcmisphere. It spans too short a period of time central Pacific Ocean (Shuntov 197,1;Maruyana and is too low in tliversity to test biogeographic et al. 1986). hypothcses. The oldest petrcl and shearwatcr fbssilsoccur in the latc Pleistoceneor Holocene Most arrivalsto thc northern part of thc Pacific from co.rstaldeposits around Ocean arc from the end of April to the end of 'l'asmaniir. mainland Australia and In these deposits the genera May or the beginning of June. The largestflocks I'terodroma and Puftinus can be observcd in the easternpart of the Bering 'l'cts predominate (Rich and van 1982). Sea in the Northern Hemisphcrc spring and the beginning of summcr. In the second half of Thc interval from 2-5000 to l0 000 yearsbefbrc summer many pass into the Chukchi Sca, and present was a period of great faunal and climatic while birds are still dispersinginto the Chukchi changc in Australia. Climatic disruptions would Sca migration southwards begins. Some birds have affected the location of Short-tailcd Shear- also migrate along the Nortll American coast water colonies through changesin sca levels. At and others across the Pacific bctwcen the times. thc coastline was up to 50 km away from Ilawaiian Islands and North Amcrica (Shuntov its currentposition (Blom l9lili; .Tcnnings1971). 1974).This results in vcry large flocks occasion- ally occurring off the west Canadiancoast in May under certain wind conditions (Guzman and DISTRIBUTION AND ABUNDANCE Myrcs 1983). Thc Short-tailed 'l'hcre Shearwatcr only breeds in In the Gult of Alaska. the Short-tailed and Australia. are known to be at least 167 Sooty Shearwatcr arc thc dominant birds in colonies around the coast of Tasmania and its spring and thev prefer the contincntal shelf, near oft.shorc islands. The total area of these which is between 100 and 150 km offshore colonies is 1 -522ha and the numbcr of burrows is (Harrison 1982). Numbers are greatest in May cstimatedat I1.4 rnillion(Skira a n1.1986). The and try Junc their estimateddensity has droppcd largestcolonies are in Tzrsmania,on Babel Islancl by half. In thc north-east of the Gulf. thc with 2.8(rmillion burrows and'l'rcfiril Island with Kodiak area, Short-tailed Shearwaltcrs out- 1.5.1million burrows ('I'owney and Skira l9U5a number Sooty Shcarwatcrs by about 1.2:1 with ancl b). Of other Australian States,Victoda has flock sizes numbering 32 (XX) (Gould et ril. 1.45 million burrows in about 30 colonies (Harris 19it2). Iray,1991 /. Sklra.'Thebiology ofthe Short-tailedShearwater

Chukchi

Gulf of Alaska ,,'4--- ..\.,. CANADA

lt rl Californra ftl Ill an tYr lslands B,i

.,1\,, . <-- {\r "€a-'

/t I AUSTRALIA

,,\v

/

Figurc I MaP conpuring new miSration (A) rcute oJ Shott-tailed Shaorwakrs based on Jupenese ship boa survcys to the figure- oJ-eight nrcvement (B) ol SerrcntJ (1953). BreedinS arcas are ipple(I.

The return journey commences at the begin 1981). The prcscnceof birds along the coastsof ning ol Septemher.Many Shorr-tailedShiar- Japan and North America led Serventy to pos- waters have been obscrved moving through the tulate a figure-of-eightmigration (Serventy 1953). we\tcrn \ectur trf lhe Pacific (Maruyama cl al. However. data from seabird survevsbv.Iaoanese lqiJb). Some flocks pass sourh antl *ell oltthore ornithologistsindicate thar migratiLnoccuir,,n a through the Gulf of Alaska to off California broad front acrossthc Pacific Ocean (Maruyama before heading acrossto Australia, but a lack of et al. l98b). RegartJlessot roule. it is appirent sightingsindicate that there is no migratorv move- that thc movement of shearwatersbetween the ment along lhe Crnadian c,rastafter August. two hemisphercsoccurs on a very broad front. It Sunrehirtls remain in rhe Norrhern Hemisohcre is possiblethat the route fbllowed varies with the during their tirstboreul \\ inter( l-i)rselland Gould age or soecificbehaviour of the bird. /. Sklra.'Thebiology of the Short-lailedShearwater Corella15(2)

FEEDING high phosphoruslcvels and, thus, a high produc- tivity. Factors Shcarwarerscommonl) form largeaggregarions that affect these warer masses .rafts aflect Shorftailcd on lhc sea \urlace edlled . Thc:\e tre Shearwaters because the abundanceand availability common in calm weather and birds may thus of their food isdepen- denl up('n the aIributes of assembleeither when feeding or resting. Short- the individurl r.rater masses. tailcd Shearwatersare one of the most niquaticof the shearwatersand have a long narrow 'knee-pelvis and compressed tarsum. well diveloped BREEDING BIOLOGY processj long sternum, Jolnt short thick com_ On arrival from their migration in September- presscdhumerus and short smooth body plumagc October, the birds clean out and refurbish their (Brooke 1990; Kuroda 1954). They liave beJn burrows. Breeding Short-tailcd Shearw.lterstend scenup ro l0 m helow rhe surfircepursuing prrl to occup) lhe sumc lrurrow as in pre\i()u\ yeJrs (Skira 1979). Their feeding mettroOs lattei or one in close proximity. During October the Ashmole 1971) rankcd in order of importance colonies arc a hive of noisy social activity. For are: pursuit plunging in which birds chasc their three weeks in November prior to egg-laying, prcy afterplunging into the water; surfaceseizing; however, thc colonies are deserted. This prc- pursrril ,.living:.cavenging; hl dr,rplaning: and laying absence cnables femalc shearwaters to hott',m lceding(Morgrn 1082:Morgan and Rirz build up body reservesto producc the egg and 1982; Ogi et al. 1980; Skira 1979). During the males the reservcs for incubation of thc egg brecding seasonthe Short-tailed Shearwatei is a (Fitzherbert 1985). neritic feeder obtaining its food close to the colony. Food. in ordcr of importance (,/o Upon their return, cggs are laicl from 19 frcquency of occurrence), are the Nlc- Novembcr to 2 December with 8-5pcr cent of thc tipllunes australis, the arrow squid Notodarus eggsbeing laid within three days on cach side of sloanigouldi anclother squid,lish and the mean laying date 24-26 November (Serventy (Montague et a1. 1986;Skira 1986).N- australisis 1963).There is no annualvariationin thispattern. abundant in large swarms particularly when Only one egg is laid and no re-lavingoccurs if the breeding betwecn October irnd December and is egg is lost. The inc[bation period varics bctween restrictcd to thc continental shelf (Blackburn 52 and 55 days and averages 53 days. Both 1980). The diet of the birds changcs from pre- partners incubate the egg in altcrnative shifts, the dominantly krill to a mixturc ol . squid and male usually taking thc lirst shift. The length of crustaccawhen cggs hatch in January. The trans- the shifts varies from l0 to l6 days and occasion- ition could be duc to reduced swarming of krill ally up to 20 days. Eggs can be left un ttcnded and irn increasein thc numbers of schoolingpost- for up to seven davs and still rentain viable as llrrrrrlhrh tMuntrguc , / o/. Iu86.l. with other shcarwaters.such as the Manx Shear- water P. pufjinus in Britain (Brooke 1990; Of sevenwater masscsmoving along the shorcs Matthews 195,1). Nearly all brccding failurcs ot northern and eiistern Australia (Rochford occur during the egg stagei only three per cent of 1957), four govern the food regime of the Short- successfullyhatchecl chicks on Fisher Island die tailcd Shearwater. Thcy are thc Subantarctic or disappearbefore banding (Serventyand Curry derived from the Southern Occan; the Southwest 1981). Tasman from the eastern approaches to Bass Strait; thc North Bass Srrait liom the South The majority of chicks hatch betwecn l0 and Austrirliiln gulfs: and the East Tasmanian-West 23 January (Oka 1989).They are broodcd by the Tasmanianfrom the centralTasman Sca and Sub- parents for the lirst few days then left unirttended antarctic.The mixing, during movement from the cluring the day. Thc chick is fed nightly for rhe sourceregions. and changingpatterns of distribu- lirst week then at longer intervals with up to 16 tion of these watcr massesdctcrmine the major days between nrealsand parentsalternate in leed- phvsicaland chcmicalcharacteristics of thc region ing. The final visit of the parents is from l-23 (Harris et al. 1987).These mixed bedies are sub- days (mcan l4) before the chicksdepart (Serventy jcct to local weather effccts such as heating and 1967). The timc between the linal feed and ctxrling which var),considerably with the seasons; departure is termed the starvation'or'desertion' thcir tenrpcraturerangc is l0'to 17".Thev have pcriod. The Short-tailcd Sheiuwater chick is in May,199'1 /. SklrarThe biologyof the Short{ailedShearwater the burrow fbr 88 to 108 days (mean 94 days). TABLE 1 They grow quickly forming large fat depositsand Brccding success.of attain a maximum mean weight of g, Short-tailed Shearwatcr rccorded in 800 nearly burrows on Fisher Island between 1947and 1984. twice that of their parcnts, in the secondweek of April (Lill and Baldwin 1983). Adult birds In DUrrows In tlrc second week of April, chicks begin to Adult birds Birds not ECC. which were cmerge from burrows at night and attempt to fly. recordcd in nssociated resulting fledged on They wander burrows with eggs in chicks Fishcr Island around and may enter any burrow "/. "/. "/. during the day, generallymoving closerto the sea total prior to dcparture.This'travel'phase is recog- 1947-1957 965 19 4l I nized by muttonbirders who may go over the 19571966 619 t'7 same area up to three times during the season. 1967-1976 161 2l 58 Chicks leave from the third week in April to the 19',77t984 009 L,{ 6li first week in May. They leave at night working their way down to the sea shore and swimming Fisher Island (Wooller et al. 1988).The breeding out to sea. Strong prevailing westerlywinds facili successof shearwatcrson Fisher Island from 1947 tate departure but also result in chicks that are to 1984 is shown in Table 1. Annual banding of not yet fully developed leaving too early and later all chicks and adults found in burrows com- perishing at sea. menced in March 1947,and it took a minimum of Chicks tend to return to their natal colony 20 years for the percentageof banded birds in (Sewenty et al. 1989)but there is probably much burrows which were fledged on Fisher Island to exploration by young birds of other areasbefore stabilizc bccauseof the longevity of Short-tailed they breed. For example. in any year only 40 per Shcarwaters. (rangc cent 16-61)of the brceding birds on Fisher The breedingsuccess ofyoung birds, 6-year-old lsland were hatched on Fisher Island (Serventy or younger, during their lirst attempt (38'/") was and Curry 1984). However, once they begin to mrrkedlylower than that r-rf hirds starring rr seven brccd at a specific colony, Short-tailed Shear- or more years(-58%) (Wooller et a/. 1988).There- watcrs have a very strong tendency to return to after, breeding successimproved with increasing breed in that colony until death (Serventy1967). experience with a particular partner. and the number of previous mates (Wooller Short-tailedShearwaters breed for the iirst time et al. 1989). The ratio of chicks fledged (termed when 4-15 yearsof agc. the mean for malesbeing to cggs laid breeding success)is about 60 per 7.J and female:7.ttyeilrs.Mate retentionappears cent annually throughout Tasmania (Naarding 1979, 1980. to be related to reproductive performance.Some 1981;Skira and Wapstra 1980). 33 per cent of all pairs which failed to produce an cgg in the preceding seasonchanged patrtncrsby divorce. However. the divorce rate was down to MORTALITY 23 per ccnt in pairs which produced an egg but Mortality is agc-rclated. Annual mortality which lailed to hatch. and 15 per cent in birds (+SE) is 7.e t 1.5% in maleand 10.61 1.8"/.in which fledgedyoung (Bradley era1. 1990; Wooller lemale shearwatersin the year of nrst recorded ct a/. 1981i).During the complered lifetimes of breedingdecreasing to 6.6 + 2.1'k and7.6 + 418 malc and temalc shearwaters,27 per cent of 2.3ok afternine yearsrising to 12.7 t 1.9% and all individualsproduced no young and 19 per ccnt 15.6 + 1.8'l. after l8 years.The median survival only onc young. Overall, 71 per cent of birds pro- time is 9.3 years after first breeding (Wooller et duced no offspring that rcturned to breed. In fact, a/. t988) although four birds on Fisher Island arc ti pcr cent of all birds that had completed their known to have been at least 36 years old (Skira, rcproductivc careers produced -53per cent of all unpubl. clata).More vigorous biids, as measured young that returned to FisherIsland to breed. and by their survival and reproductive success,may 26 per cent of all birds were responsiblefor all tend to have a greater reproductiveoutput earlier reproducing offspring. Shearwatersthat formcd in life u hcrerr'intlir iduals with lowcr vigourmry known pairs produced on averitge 5.3 eggs, 3.1 produce fewer offspring and die earlicr. How- fledglings.and 0.,13reproducing offspring eachon cvcr, among birds which have brcd for 15 years, /. Sk//arThe btology of rheShort-tailed Shearwatel Corella15(2) those which 'health' havc fledged fewer young have a in Short-tailed Shearwatersparticularly .lrghl bul significanlly higher sulivul iatc rhan w-hen in the Norrhcrn Hcmisphcre iDay er ai. lhosc whieh hare produced morc offsorins .1q85).At thc momenrrhis'impairmcnr'has nrrr (Bradfcy er a/. 1989;Woollcr et al. 1990). bcen mcaslrred. The grcatestmortality (52%) occursin thc first ycar.of life_(Scrventy1967). In some years,large FUTURE RESEARCH numbers of Shorftailed Shearwatersare washdd Much is known about the onto Japanesebeaches when casterlywinds blow breeding biology of short-tailed Shearwatcrs weakencd birds off thcir normal route. Autopsies while at the breeding colony. This is due to the have establishedthat death is due to starvation long-term study on Fisher Island which has also and lhirl Ihr mrjoriry arc fledglings(Nishigai er made very important contributions to the ecology a/. lqxl: Okr antl Maruyama tOSo.1.Oi tne -haveof veitebrate . Strayer et aL. (1986) rcturn tnp annual mortalities are invcrsely pro- reviewed the essentialqualities of long-term studics portional to fluctuations in plankton numbe;s in and most of them are exemplifiedin the thc Tasman Sea (Serventy ei al. 1971).Autopsies Shorftailed Shear- water researchon Fisher Island. on l4 shearwaters found dead along one Tas- manian beach in Deccmber 1983 showed that There are many questionsstill to answer. For death was due to starvation(Skira, unpubl. data). examplc, the roles of chancc and othcr factors in determiningwhich Naluftrl causes ol murtalit) are prcdation. birds havc long and productivc lifespans have yet to be determined. Jlseirs(.\tarvaliitn und floodingol low-lyingnest- Virtually nothing is known of the social ing areas. Ouite scvere mortalities are'cauied in behaviour and pelagic biology of shearwaters. somc years by a condition known as'limy-bird The reasons for their great abundance is disease'.which is associatedwith blockageof the unclear. Advances in tcchnological developmcnts lo-werpart of thc alimentary canal by concretions such as automatic burrow monitors of socliumurate (Mykytowycz 1963j. and radiotelemctry will make someof the work easier.There is a need for inter- Apart lrom muttonbirding which takesapproxi- disciplinary resezrrchthat includes oceanography t,,1ql1 l(100{lll chicks e:tch yelr. irn(,lherhuman becauseof the depcndenceof Short-tailed .Shear- rclrvitv hrs hrd Jisa:rrous atfects. Gillnel waters on the marine environment. fisherics in thc North Pacific operate using mothcrships and the Japaneseenterprise drown,s ACKNOWLEDGMENTS between 131000 and 281 000 Shorrtailed Shear- urlers unnualll(King lu84:Ogi lu84).Korerr and Hans Wapstra, David Pemberton, Darryl Jones I ir$itn rlso ,'perarcgillner llsheries in the North and Durno Murray made very useful suggestions Pacific.The effcctsof a salmon land-basedfishcry for this paper. I would also like to thank Ron and a squid fishcry are not known but they Wooller and Stuart Bradlcy for providing details opcrate in an area through which large numbcr.s for Table 1. ,'i slr,)n-ruiledShcrrrwaiers parr duiing migra- tion. The Uniled Slillc\ and C-anadulrre curre"ntly REFERENCES devcloping obscrver programmes with Japan. 'by-catcli' Ashmolc.N. P. (1971).Seabird ecolog! and rhe marine Korca and Taiwan to look at the of environmcnt.In Avian Biology.Vol. 1. (Eds D. S. squid gillnet lishcries in thc North Pacific(Gould, Farncrand J. R. King)pp 223-286.(Acrdcmic Pressr New Pcrs_comm.J. York.) Azzrrelb. M. Y- and van Vled. E. S. (1987). Small plastic particles are commonly found Marinebirds in andplastic pollution. Mar. Ercl. Prog.,tsr. stonachs of 37:295,303. scabirds (Azzarello and van Vleet Blackburn.M. (l9tj0). Observarionson the disrribulioDof l9ll7). A high proporrion of Short-tailed Shear- Ny&phanes dastralts Sars (Crustacea. Euphausiidae)in watcrs contilin plastic particles in their stomachs Australian walcrs. CSIRO Div. I.ish. Orcanogr. Ilep. tlgl on their rcturn to the Southern Hcmisphere but t i0. k)se thcm as thc seasonprogrcsscs (Skira 1986). Blon. W. M. (l9lt8). Late Quaternarvscclimenrs and sca levclsin BassB:rsin. southcasrefn The effccts of plastic ingesti6n are unknown Ausrralia A prclimin but ar!,rcport..Scdl.lr 19: 9+96. there rs some suggestionof a link between high Bowdler. S. (19134).llunter Hill. Hunrcr Island. 7elr..] amountsof plasticingested and decreasedphysical .4llrrr..r/tl8: l-14u. N4ay,1991 /. Skl/.arThe biologyofthe Short.iailedShearwater

Bradley. J. S., Wooller, R. D.. Skira. L J. aDd SeNcnty. D. Johnslonc,R. E.. Smith, L. A.:'nd Klo:rp. N. L (1991)b). t-. (l9lt9). Age-depcndcnt survival of bfccding shorl rail Scabird lslands. No. 204 Gulch lsland. Archipelago of ihc sheirrwaicrs Pufiinus !?nuirostri!. L Anim. Eol. 58: Rcchcrchc, Wcstcrn Austr:rlia. Corella 14:133 131. 175-luti. King, W. B. (19E4).Incidental morlrlily of seabirdsiD gillnets Bradley. J. S.. Wooller. R. D.. Skira. L J. irnd Scrvcnty. D. in tbc Norlh Pitcilic. In Slatus rnd ConseNati(nr of thc L. ( 1990) Thc infiucnccof mirc rctcnli()nand divorcc upon Worlds Scabirds. ([ds J. P. Crox.r]I. P. C. H. Evirns rnd fcproduclive successin Sho tailed ShearwatcrsPr,?inrs R. W. Schrcibcr) pp. 709 715. lnt. ( ounril Bird I'rcwva totuirosrri.\.L Anutl. E.ol.59: .187.196. tiui Ie(h. I'uh. Na. 2. Rrookc. M. (1991)) The Manx Shearwater'. (T. & A. D. Kuroda, N. (1954). On the classificationand phyl()gcnvof Poyser: London.) the ordcr Tubinarcs. particularlv the shearwrtcr\ (l'uJrtnus). D:'y. R. H.. wchle. D. H. S. and Colcman,F. C. (t9u5) \\irh special considerationson thcir oslcolog) Ingestionof plasticpollutants bymariocbirds. In Proceed- and habitat diffcrcntiation (Avcs).' ('fhc Author: Tok),o.) (19t6). ings of the Workshop on the Fatc and Impact of Marinc Kuroda. N. SynrhcticStudy of thc Mass-norrality of (Study Debris. (Eds R. S. Shomura ard H. O. Yoshida) pp.344, Shor.'tailcd Shcarwatct Pulfnus tenui.osttis. group of Shoft lailed Shearwrter: Abiko. (ln 3t16.(U.S. Dcpart. Commer: NOAA Tech.) Japan) Jitpanesc.) Lanc. S. C. (1979). Summary of lhe brocdirg scabirLlson Demcnliev.G. P. and Cladkova,N. A. (1951).Birds of thc Ne*. South Wales coastrl islands.Corcllu 3:1 1\) Sovicl ljnion. Vol. 2. pp. 370 -]7,1.(Moscow) (Translarcd Lanc. S. C. (1983).Short{ailed shcarwalcrin l.ieurc oi!ighl froln Russian.) lshnd Archipelago of the Recherchc. Wcsrcrn Auslralia. Filzhorbcrl, K. (1985). Thc rolc of energericfaclors in thc tr?rr 83: 37 38. cvolulioD of thc brccding biology of the sbort-tailedshcar Lill, A. and Baldwin. J. (1983).WciSht changcsand lhe modc wirlcr (/'rdrrrr /.r//iro.!r/i.r Temminck). Ph.D. Thesis, of dcpot fat iccumulation iD migrrrcrv short-lailcd shcar- Monash Uni\rersit!. Australir. walers. ,4r1.r/.J. Zool.3l.891 902. Forscll. D. J. an(l Could. P. J. (l9lr1). Disrribution and Marshall, A. J. irnd Serventy. D. I-. (1956). 1hc brccding abundanceof nlilrinc birds and mammals winteriDgin thc cycle of thc short tililcd shcarwater, Puf/1nus k\tuirostri! Kodiak arca of Alaski. Biok)gicalServiccs Program El/13. (lemminck). in rclation to lranscquationallni.qration aod (US Fish irnd Wildlife Servicc:Washingrcn.) its cnviroDmenl. Pto(. Zool. So.. L.rr1. 12?:489 510. Fricdnann. H. (1931a1.Ilird bones irom cskimo ruins on St Maruvamx. N.. Oka. N.. Watabc. Y . Kuroda. N. and Skira. Lawrcncc Island. BerinS Sca. J. WashingtonAcatl. Sci.24: L J. (l9116).Migrarion route of shor!-lailcd shcar*atcf in fiz 96 North Pacific.In Syrthclic Sludy ol thc Mass,mortalityof Friedm nn. H. (1934b). Uird bones frcm old cskimo ruins in Shorttailed Shcnrwaters /1/rrilr.\ /{,?Iriro.r/r.r.r'.(Ed. N. n laska.J. Wd.rrrr8l.rt Atatl. Sci.24:230-231. Kuroda) pp 3(F12. (Study Ciroup of Shorl lrilcd Shearwat- I:ricdmann. H. (1911). Bird bone\ from cskimo rui s at Capc crs: Abiko. Japan.) (ln Jap:',rcsc.) I'rincc of Walcs. Alirska. "L W.rr,trr?g1anA&d. Sd.4l: Matthews. Ci. V. I . (1951) Somc aspccrsof incubationin lhc ,10.]_-109 Mrnx shearw.rterI'r'o. ellariu pulJinur. wilh parricullrr refcf, (n)uld. I' J.. Forscll.D. J. rnd I-cnsink,C. J. (1982).Pcllgic c cc 1o chilling resistrncein thc embryo. /rir 96: ,1114.10 drstributiona d bundanceof seirbirdsin rhc aiulf of Monlnguc. T. L.. alllon. J. l\{. lnd Filzhcrbcrr. K. (l9lJ6) Altiskl 'fhc rnd eastcrn Ilcring Scl. LtS l.ish Wildl .lcr. 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