Corella, 1991 , 15(2): 45-52

A11srralasia11 Hird l?<'l'iews - N11111ber 3

THE SHORT-TAILED : A REVIEW OF ITS BIOLOGY

IRYNEJ SKIRA

Department of Parks. Wiltllifc and Heritage, 134 Macquarie St.. Hobart. Tas 7000

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The life history of the Short-tailed Shearwater tenuirostris has been well documented since it first came to the attention of naturalists. Approximately 23 million breed in about 250 colonies in southeastern from September to April. The Short-tailed Shearwater commences to breed when 4 to 15 years. During their completed lifetimes 27 per cent of all individuals produce no young and 19 per cent only one chick. Mortality is age-related with the median survival time for breeding being 9.3 years after first breeding. Many areas remain open for study, with a particular need for interdisciplinary research that includes oceanography.

INTRODUCTION The Short-tailed Shearwater was one of the first Australian birds to be banded in large numbers The Short-tailed Shearwater Puffinustenuirostris, (Serventy 1957, 1961) and to be subjected to a commonly known as the Tasmanian muttonbird, long-term scientific study (Guiler et al. 1958). This is one of about I 00 species in the Procellarii­ study was commenced on Fisher Island in the formes. The diagno�tic feature of the order is the of in March 1947 by external nostrils produced into tubes extending Dominic Serventy formerly of the CSIRO onto the bill. Other distinctive features are the (Scrventy 1977) and continues to the 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 92 000 duces the well known 'oil'. actually wax esters birds in Australia, the life history of the Short­ (Warham 1977). Most members in this order have tailed Shearwatcr is one of the best documented a distinctive musky odour. in the world of any ( Bradley et al. 1989. 1990: The family Proecllariidac is the most diverse Serventy 1974; Servcnty and Curry 1984: Wooller group in the order and contains 61 species of et al. 1988, 1989. 1990). petrels and (Serventy et al. 1971). The majority are nocturnal and nest in holes, EVOLUTION burrows or crevices, which serve to protect them and their young from predators. The genus The Procellariifonncs are an ancient group of Puffinus consists of 15 medium-sized species that birds that probably originated from aquatic birds are among the world's most numerous . present at the end of the Cretaceous. some 64 Their high nesting densities and their fidelity to a million years ago. The phylogenetic history of particular site has meant that they are highly shcarwaters is little known (Kuroda 1954). vulnerable to exploitation (Serventy et al. 1971). According to Olson (1985 p.211), ·most of the The harvesting of Short-tailed Shearwaters, or modern species-groups, or subgenera, of Puffinus muttonbirding as it is known in Tasmania, is one were in existence by the Middle Miocene. and of the best documented instances of there has been very little morphological change harvesting (Skira 1987, 1990). within these lineages in 15 million years or so'.

45 46 I. Skira: The biology of the Short-tailed Shearwater Corella 15(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. sites (Bowdler 1984: Friedman 1934a and b, 1941; comm.), New South Wales 25 700 breeding pairs Vanderwal and Horton 1984). One of the fossil in I 3 colonies (Lane 1979) and Western Australia procellariids. P. conrudi from the Middle 10 000 burrows in several colonies (Johnstone et Miocene of Maryland. USA. seems to have lived al. 1990a and b; Lane 1983). It is estimated that to the Pleistocene. one million years ago. P. 23 million birds breed in about 250 colonies. 1e1111iros1ris may have evolved from a P. conradi­ Recent research by Japanese biologists has con­ type ancestor through the P. inceplor line centrated on mapping vegetation types in several (Kuroda 195--l). /'. inceptor is known from the colonies in Tasmania (Kuroda 1986). Middle Miocene of California suggesting that P. 1e11uirostris and its congener the Sooty Shearwatcr P. griseus which is abundant in MIGRATION New Zealand. were differentiated in the North Atlantic-North America areas. Having settled in The Short-tailed Shcarwater is a circum-Pacific the Southern Hemisphere. perhaps as late as migrant spending the horcal summer in the the Pleistocene. their post breeding migration Northern Pacific region (Fig. l). Sexually may be an instinctive response to return to the immature birds depart from Australia near thc Northern Hemisphere (Kuroda 1954; Marshall end of March. followed by the breeding birds and Serventy 1956). around mid-April, with the Acdged chicks leaving at the end of April to early May. The shearwaters The fossil record left by seabirds in Australia migrate rapidly (Scrvcnty 1956) and arrive in the is meagre compared to that of the Northern Northern Hemisphere on a broad front across the Hemisphere. It spans too short a period of time central Pacific Ocean (Shuntov 1974; Maruyama and is too low in diversity to test biogeographic et al. 1986). hypotheses. The oldest petrel and shearwatcr fossils occur in the late Pleistocene or Holocene Most arrivals to the northern part of the Pacific from coastal deposits around mainland Australia Ocean arc from the end of April to the end of and Tasmania. In these deposits the genera May or the beginning of June. The largest flocks l'terodroma and Puffinus predominate (Rich and can he observed in the eastern part of the Bering van Tets 1982). Sea in the Northern Hemisphere spring and the beginning of summer. In the second half of The interval from 25 000 to 10 000 years before summer many pass into the Chukchi Sea. and present was a period of great fauna! and climatic while birds are still dispersing into the Chukchi change in Australia. Climatic disruptions would Sea migration southwards begins. Some birds have affected the location of Short-tailed Shear­ also migrate along the North American coast water colonies through changes in sea levels. At and others across the Pacific between the times. the coastline was up to 50 km away from I lawaiian Islands and North America (Shuntov its current position (Blom 1988; Jennings 1971). 1974). This results in very large flocks occasion­ ally occurring off the west Canadian coast in May under certain wind conditions (Guzman and DISTRIBUTION AND ABUNDANCE Myres 1983). The Short-tailed Shearwater only breeds in In the Gulf of Alaska. thc Short-tailed and Australia. There are known to be at least 167 Sooty Shcarwatcr arc the dominant birds in colonies around the coast of Tasmania and its spring and they prefer the continental shelf. near offshore islands. The total area of these which is between 100 and 150 km offshore colonies is I 522 ha and the number of burrows is (Harrison 1982). Numbers are greatest in May estimated at 11.--l million (Skira el al. 1986). The and by June their estimated density has dropped largest colonies are in Tasmania. on by half. In the north-east of the Gulf. the with 2.86 million burrows and Trefoil Island with Kodiak area. Short-tailed Shearwaters out­ 1.54 million burrows (Towney and Skira 1985a number Sooty Shcarwatcrs by about 1.2: I with and b). Of other Australian States. Victoria has flock �izes numbering 32 000 (Gould et al. I .--l5 million burrows in about 30 colonies (Harris 1982) May, 1991 /. Skira: The biology of the Short-tailed Shearwater 47

' CANADA ' ' '\ \ \ \ \ \ \ \ I I I California I I IIJ II I I I I I ,Is , I I I I I I I I I / / / ,,/ � ��/ � / ,,

AUSTRALIA

r-igur<: I. Map cn111pari11g 11ew 111igm1io11 ro111e (A) of Shorr-/(/i/ed Sheanvmers based 011 Japa11ese ship-board s1,r1•eys 10 rhe fig11re­ ofe1gh1 111ove111e111 ( B} of Serve111y ( 1953). Breedi11g areas are srippled.

The return journey commences at the hegin­ 1981). The presence of birds along the coasts of ning of September. Many Short-tailed Shear­ Japan and North America led Scrvcnty to pos­ waters have been observed moving through the tulate a figure-of-eight migration (Scrvcnty 1953). western sector of the Pacific (Maruyama et al. However. data from seabird surveys by Japanese 1986). Some flocks pass south and well offshore ornithologists indicate that migration occurs on a through the Gulf of Alaska to off California broad front across the Pacific Ocean (Maruyama before heading across to Australia. but a lack of et al. 1986). Regardless of route. it is apparent sightings indicate that there is no migratory move­ that the movement of shcarwaters between the ment along the Canadian coast after August. two hemispheres occurs on a very broad front. It Some birds remain in the Northern Hemisphere is possible that the route followed varies with the during their first boreal winter (Forsell and Gould age or specific behaviour of the bird. 48 I. Skira: The biology of the Short-tailed Shearwater Corella 1 5(2)

FEEDING high phosphorus levels and. thus. a high produc­ Shcarwatcrs commonly form large aggregations tivity. Factors tha t affect these water masses on the sea surfa ce called ·rafts'. These arc affect Short-tailed Shcarwaters be cause the common in calm we athe r and birds may thus abundance and availability of their food is depen ­ den t upon the a ttributes of the individual water assemble either when feeding or resting. Short­ tai led Shcarwatcrs are one of the most aqua tic of masses. the shearwaters and have a long narrow pelvis BREEDING BIOLOGY and compressed tarsum. well developed kn ee­ joint proce ss_ long sternum. short thi ck corn­ On arrival from th ei r migration in Scptcmbcr­ prcs-; cd humerus and short smooth body plumage October, th e birds clean out and refurbish their (Brooke 1990: Ku roda 1954 ). They have be en burrows. Breeding Short-tai led Shearwa tcrs ten d seen up to lO m be low the surface pursuing prey to occupy the same burrow as in previous years (S kira 1979). Their feeding metho ds (after or one in close proximity. During October the Ashmo le 1971) ranked in order of importance co lonies arc a hive of noisy so cial activity. For arc: pursuit plungin g in which birds chase their three weeks in No vember prior to egg-laying. prey after plunging into the water; surface seizing; ho wever, the co lonies arc deserted. This pre­ pursui t diving; scavenging; hy droplaning; and laying absence enables female shearwaters to bo ttom feeding (Morgan 1982; Mo rgan and Ritz bu i ld up bo dy reserves to produce th e egg and 1982: Ogi et al. 1980 ; Skira 1979). During the males the reserves for incubation of the egg br eeding season the Short-tai led Shcarwatcr is a (Fitzherbert 1985). neritic feeder obtaining its food close to the co lony. Foo d. in order of importance ('¾, Upon their return, eggs are laid from 19 frequency of occurrence). arc the krill Nyc­ November to 2 December with 85 per centof the tiplwnes australis, the arrow sq ui d Notodarus eggs being laid within th ree days on each si de of sluani gouldi and other squi d. fish and crusta ceans the mean laying date 24-26 November (S crventy (Montague et al. 1986: Skira 1986). N. australis is 1963). There is no annual variation in this pattern . abundant in large swarms particularly when Only on e egg is laid and no re-laying occurs if the breeding betwe en October and December and is egg is lo st. The incubation period varies between restri cted to the con tinen tal shelf (Blackburn 52 and 55 days and averages 53 clays. Both 1980 ). The diet of the bi rds changes from pre­ partners incubate the egg in alte rnativeshi fts, th e dominantly kri ll to a mixture of fish. squi d and male usua lly takin g th e first shift. The length of crusta cc a when eggs ha tch in January. The trans­ th e shifts varies from 10 to 1 6 days and occasion­ ition co uld be due to reduced swarming of kri ll ally up to 20 days. Eggs can be left unatten ded and an increase in th e numbers of schooling post­ for up to sev en days and sti ll remain viable as larval fish (Montague et al. 1986). wi th other sh carwaters. such as th e Manx Shear­ wa te r P. puffinus in Britain (Brooke 1990; Of seven water masses moving along the shores Matthews 195-t). Nearly a ll breeding fai lures of northern and easte rn Australia (Rochford occur during the egg stage; on ly three per cen t of 1 957). four go vern the food regime of th e Short­ succcssfullv hatched chicb on Fi sher Island die tai led Shearwater. Th ev are the Suban tarctic or disappc;n before ban ding (S ervcnty and Curry derived from the Southern Ocean: the Southwest 1 984). Tasman from the eastern approaches to : th e Nort h Bass Strait from th e South The majority uf chicks hatch betwe en 10 and Australian gulfs; and the East Ta sm anian-West 2J January (Oka 1989). They are brooded by th e Tasmanian from the central and Sub­ parents for the firstfew clays then left un atten de d an tarctic. The mixing. during movemen t from the during th e day. The chick is fed nightly for the so urce regions. and changing pattern s of distribu­ first week then at lo nger intervals with up to 1 6 tio n of th ese wa ter masses dete rmine th e major days between meals and parents a lternatein feed­ phy�ical and chemical characteristi cs of the region ing. The final visit of the parents is from 1-23 ( Ha rris et al. 1987). Th ese mixed bo dies are sub­ days (mean 14) before the chicks depart (Serven ty ject to loca l weather effects such as hea ting and 1 967). Th e time between the final feed and coo ling whi ch vary considerably with the sea sons; departure is term ed the ·starvation· or ·desertion· th eir temperature range is 10° to 17°. They have period. The Short-tailed Shcarwater chick is in May, 1991 I. Skira: The biology of the Short-tailedShea rwater 49 the burrow for 88 to 108 days (mea n 94 days). TABLE l They grow quickly forming large fa t deposi ts and Breeding success of Shon-tailed Shcarwatcr recorded in attain a maximum me an wei ght of 800 g, nearly burrows on Fisher Island between 1947 and 1984. twice tha t of their parents . in the second week of Apr il (Lill and Baldwi n 1983). Adult birds in burrows In the second week of April, chicks be gin to Adult birds 13irdsnot Eggs which were emerge from burrows at ni ght and attemp t to fl y. recorded in associated resulting fledged on They wander around and may enter any burrow hurrows with eggs in chicks Fisher Island total o during the day. generally mov ing closer to the sea Years %, (½) (Y prior to de parture. This 'travel' phase is recog­ 1947-1957 I 965 19 41 niz ed by muttonbirdc rs who may go over the 1957-1966 I 619 17 66 16 same area up to three times dur ing the season. 1967- 1976 I 163 �I 58 -n Chic ks leave from the thi rd week in April to the 1977-1984 I 009 14 68 45 fi rst week in May. They leave at night working their way down to the sea shore and swimming Fisher Is land (Wooller et al. 1988). The bre eding out to sea. Strong prevailing weste rly wi nds facili­ success of shearwatcrs on Fisher Is la nd from 1947 ta te departure but also resu lt in chicks tha t arc to 1984 is shown in Table I. Annua l banding of not yet fully develop ed leaving too early and la ter all chic ks and adults found in burrows com­ perishing at sea. me nced in March I 94 7, and it took a minimum of Chicks tend to return to their na tal colony 20 years fo r the percentage of banded bi rds in (Serve nty et al. 1989) but there is probably muc h burrows which we re fledged on Fisher Is land to explorati on by young birds of other areas before stabi liz e because of the longevity of Short-tailed they breed. For example. in any year only 40 per Shearwaters. cent (range 16-61) of the breeding birds on Fisher The breedi ng success of young birds, 6-year-old Island wc rl: hatched on Fisher Is land (Scrventy or younger, during their firs t atte mpt (38%) was and Curry 1984). However. once they begin to markedly lower tha n tha t of bi rds starting atseven breed at a specific colony, Short-tailed Shear­ or more years (5 8%) (Wooller et al. 1988). There­ waters ha ve a very strong tendency to return to after, breeding success improved wi th increasing breed in that colony until dea th (Serventy 1967). ex perience with a particular partner. and the m1 mber of previous ma tes (Wooller et al. 1989). Short-tailed Shcarwatc rs breed for the fi rst time The ra ti o of chicks fledged to eggs laid (termed when -l-- 15year � of age. the mea n for males being 7.3 and fe males 7.0 years. Ma te retention appears breeding success) is about 60 per cent annual ly to be rela te d to reproductive performance. Some throughou t Tasma nia (Naarding 1979. 1980, 33 per cent of all pairs which failed to produce an 1981; Skira and Wapstra 1980 ). egg in the prece di ng season changed pa rtners by MORTALITY divorce. However . the divorce rate was down to 23 per cent in pai rs which produced an egg bu t Mortali ty is age-related. Annu al mortality which fa ikd to hatch. and 15 per cent in bi rds (±SE) is 7. 8 ± l.5% in male and 10.6 ± 1.8% in which fledged young (Bradley et al. I 990: Woo lier fema le shearwaters in the year of fi rs t recorded e1 al. 1988). During the completed lifetimes of breeding decreasing to 6. 6 ± 2.1 % and 7. 6 ± -1 18 male and fema le shearwatcrs. 27 per cent of 2.3°/4-, after nine ye ars rising to 12.7 ± 1. 9% and all individua ls produced no young and 19per cent 15.6 ± 1.8% after 18 years. The median surv ival only one young. Overall. 71 per cent of birds pro­ time is 9.3 years after first breeding (Woollc r et duced no offspring tha t return ed to breed. In fact. al. 1988) although four birds on Fisher Island arc 8 per cent of all birds tha t had completed their known to have been at least 36 years old (Skira. repro ductive careers produced 53 pe r cent of all unpubl. data). More vigorous birds, as measured young tha t retu rned to Fisher Is land to breed. and by their survival and reproductive success, may 26 per cent of all birds were responsible for all tend to have a grea te r reproductive output earlier reproducing offspring. Shearwatc rs that forme d in life whereas individuals with lower vigour may known pairs produced on average 5.3 eggs . 3. l produce fewe r offspring and die earlier. How­ fledglings. and 0.-13 reproducing offspring each on ev er. among birds whic h have bred for 15 years. 50 I. Skira: The biology of the Short-tailedShearwater Corella 15(2) those which havc Hedged fewer young have a 'health' in Short-tailed Shcarwate rs part icula rly slight but significantly higher survival rate than when in the Northern Hemisphere (Day el al. tho se which have pro duced more offsprin g 1985). At the moment this 'impairment' ha s not (Bradley el al. 1989; Wo oller e1 al. 1990). been measured. The greatest mortality (52%) occurs in the first year of life (Serven ty 1967). In some years, large FUTURE RESEARCH numbers of Sho rt-tailed Shearwaters arc wa shed Much is known about the breeding biology of onto Japanese beaches whe n easterly wi nds blow short-tailed Shearwatcrs while at the breeding we akened birds off their normal route. Autopsies colony. This is due to the lo ng-t erm study on ha ve established that death is due to starvation Fishe r Isla nd whic h has also made very important and that the majority arc tledglings (Nishigai el contributions to the ecology of ve rt ebra te al. 1981; Oka and Maruyama 1986). On the animals. Strayer e1 al. ( 1986) have reviewed the return trip annual morta lities arc in versely pro­ essen ti al qualities of long-term studies and most po rtional to tl uc tuations in plankton numbers in al. of them arc exemplified in the Sho rt-tailed Shear­ the Tasman Sea (Serve nty el 1971). Autopsies wate r research on Fisher Island. on 14 shearwaters fo und dead along one Tas­ manian beach in December 1983 showed that There arc many questio ns still to answer. Fo r death was due to starvation (Skim. unpubl. da ta ). example. the ro les of cha nc e and other factors in determ ining whic h birds have lo ngand productive Nat ura l causes of mortality are predation. lifespans ha ve yet 10 be det erm ined. Virtually disease. sta rva tion and floodin g of lo w-lyin g ne st­ nothin g is known of the social behaviour and in g areas. Quite severe mortalities arc ca used in pelagic biology of shcarwatcrs. The re asons for some years by a condition known as 'limy-bird their great abundance is unclea r. Advances in disease·. whic h is asso ciated wi th blockage of the techno lo gical develo pm ents such as automatic lower part of the alim entary canal by concretio ns burro w monitors and radiotc lcmctry will make of sodium urate (Mykytowycz 1963). so me of the wo rk easier. There is a ne ed for inter­ Apart from mutton birdin g which takes appr oxi­ disciplinary re search tha t inc ludes oceanogra phy mately 300 000 chicks each year, anothe r human because of the dependence of Short-tailed Shear­ activitv has ha d disastrous affects. Gillnet waters on the marine environment. fisheries in the Nort h Pacific operate us ing mothc rships and the Japanese en terprise drowns ACKNOWLEDGMENTS between l3 I 000 and 281 000 Sho rt-tailed Shear­ wa ters annually (King 1984; Ogi 1984). Korea and Hans Wa pstra, David Pemberton. Darry l Jo ne s Taiwa n also operate gillne t fisherie s in the Nort h and Durno Murray made very useful suggest ions Pacific. The effects of a salmon la nd-base d fishery fo r this pa per. I wo uld also like lo thank Ron and a squid fo,hcry arc not kno wn but they Woolle r and Stuart Bradley for pro vidin g details opera te in an area through whic h large numbers for Table I. of Short-tailed Shcarwa tcrs pass durin g migra­ tion. 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