Common Murre •.• Thick-Billed Murre
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j. Field Ornithol., 54(3):266-274 THE FLEDGING OF COMMON AND THICK-BILLED MURRES ON MIDDLETON ISLAND, ALASKA BY SCOTT A. HATCH Three speciesof alcids,Common and Thick-billed murres (Uria aalge and U. lornvia)and the Razorbill (Alca torda),have post-hatchingdevel- opmental patterns intermediate to precocialand semi-precocialmodes (Sealy1973). The youngleave their cliff nestsites at aboutone quarter of adult weight and completetheir growth at sea.At departure, an event here looselyreferred to as "fledging," neither primary nor secondary flight feathersare grown, but well-developedwing covertsenable lim- ited, descendingflight. The adaptivesignificance of thispattern maybe that leadingthe young to distantfeeding areasis more efficientthan lengthy foraging flights by adultsonce the chicks'energetic requirements exceed some critical level (Sealy 1973, Birkhead 1977). The risk to predation is probably greater in exposednest sitesthan at sea, contributing further to the selectiveadvantage of earlyfledging (Cody 1971, Birkhead1977). Given these constraints,however, larger chickswould be expected to better survivethe rigorsof fledging,increased activity at sea,and the vagaries of weather. Hedgren (1981) analyzed278 recoveriesof CommonMurres banded as fledglingsand found no significantrelationship between fledging weight and subsequentsurvival. This result is paradoxicalin view of a strong expectationto the contrary and evidencethat such an effect occursin other speciesof birds (Perrins 1965, Perrinset al. 1973, O'Con- ner 1976). Hedgren suggestedthe nestlingperiod of murres,averaging about3 weeks,is determinednot by a thresholdin bodysize, but by the relativelyconstant time requiredto completefeather growth.Birkhead (1977) alsoemphasized the need for chicksto attain a critical weight: wing-arearatio prior to fledging. Clearly, however,body weight and feather developmentare not mutuallyexclusive factors affecting fledg- ling survival. Murres breeding on Middleton Island, Alaska use nest siteslocated sofar from the water'sedge that direct flightsto the seaby the fledglings are impossible.In 1978, I capturednearly 400 fledglingsas they made their wayoverland to the sea.In thispaper I analyzemensural data for thesebirds with a viewto testingthe abovehypotheses regarding optimal fledging condition. My data also document population changesand breedingphenology of murres in a region for which there is little pub- lished information. STUDY AREA AND METHODS Middleton Island is in the north-central Gulf of Alaska about 115 km from the mainland and 18 km from the edge of the continental shelf 266 Vol.54, •o. • FledgingofMurres in Alaska [267 ,, I 'L,..,• Alaska \ /I ,//••', '.,--.....'"'-'-'.... ,'// /// Present low water ' ' •-'•, reefs ?'- I km F•cuRE1. Map of MiddletonIsland showing changes in the shorelineand the distribution of murres in 1978. CM, Common Murres; TBM, Thick-billed Murres. (Fig. 1). The islandis generallyflat, and reachesa maximum elevation of about 40 m on its southwestside. The Alaskan earthquakeof 1964 raisedthe islandabout 4.5 m, exposinglarge areasof previouslysub- merged land. On the east, south, and southwestthe distancebetween the sea cliffs and the present high water mark averagesabout 400 m. The intertidal zone extends another 200 to 1200 m seaward. Most murres nested near the top of a steep cliff surrounding the southernthird of the island.Nesting ledges did not exceedabout 35 m in height. The largestgroup of Common Murres, designatedthe "main colony," nestedon a broad ledge only 10-15 m high but 500 m from the high water level. Flatsbelow the cliffsconsisted of marshyground with a scatteringof large bouldersand numerousshallow ponds of fresh or brackish water. A censusof murres was carried out on 26 July when all birds were counted with a spotting scopefrom the flats. The proportions of the two speciesin the populationwere estimatedfrom the ratio of Common to Thick-billed murres in the population of chicksleaving the cliffs in August. 268] s.A. Hatch j. FieldOrn,thol. Summer 1983 Chicksfrequently did not completethe journey to seaon the same eveningthey left the cliffs,but spentone or moreof the followingdays accompaniedby adultsin the ponds.Chicks were capturedby hand duringdaily searches of the pondsbetween 26July and 16 August.Wing chord wasmeasured to the nearestmm with a stoppedruler; weight wasdetermined to within 1 g with a 300 g Pesolascale. All chickswere marked with U.S. Fish and Wildlife Servicemetal leg bandsat first capture,which permitted me to separaterecaptured chicks from those newlyarrived in the pondseach day. Everychick present in the ponds was captured each day, and the measurementof weight was repeated on recaptured chicks. RESULTS AND DISCUSSION Censusand populationchange.--A total of 6803 murres was counted in the census.This figure agreeswith independentcounts by Gouldand Zabloudil,and Gouldand Nysewanderin 1981 and 1982, respectively (P. J. Gould, pers. comm.). Southern et al. (1965) showedthat the pro- portion of murrespresent on ledgesat any time during incubationand broodingis about 60% of the total population(including both breeders and nonbreedingnest-site holders). Extensivecounts at the Semidi Is- lands, Alaska (Hatch, unpubl• data) substantiatethis value as a conser- vativeadjustment for nest-siteattendance. Thus, the total population on MiddletonIsland in 1978 wasabout 11,000 birds(Fig. 1). Thick-billedMurres occurredonly north of the main colonyon the westside of the island.Here, theymade up about7.4% of the population, asestimated from the numberof Thick-billed Murres among311 chicks capturedin the area during fledging.I assumethe two specieshad similar nestingsuccess. The ratio agreeswith my general impressionfrom ob- servationsof birds on the ledges.Thick-billed Murres thus made up only 3% of the censustotal, or about 350 birds. Rausch(1958) visited Middleton Island in 1956 and found a popu- lation of only about 400 murres, mostlyThick-billed Murres. Thus, although the population of Thick-bills has changed little, Common Murres increasedgreatly between 1956 and 1978. The increaseis all the more remarkablein view of physicalchanges brought about by the 1964 earthquakewhich might havebeen expectedto discouragemurres from nestingon Middleton Island. The changein numbersof murres was paralleled by large increasesin the populationsof Black-legged Kittiwakes(Rissa tridactyla) and PelagicCormorants (Phalacrocorax pe- lagicus)(Hatch, unpubl. data). Timingoffledging.--The distribution of fledgingbetween 26 July and 16 Augustwas approximately normal, exceptfor a conspicuousdecline in the number of chicksleaving the cliffson 31 July and 1 August(Fig. 2). The lull occurredduring an unusualspell of foggyweather, although suchconditions might have been expectedto have the oppositeeffect. Peak numbersof birds left the cliffsas soonas the fog lifted. A few chicksprobably fledged before daily observationswere begun. vo•.54, No. • Fledgingof Murres in Alaska [269 50 Common Murres (exceptmain colony) 40 Thick-billed Murres Fog Common Murres (maincolony) '• 50 o '" 20 I0 ? I •5 50 5 July August FIGURE 2. Distributionof fledgingdates of murres on Middleton Island in 1978. On 27 May severalhundred murres were flushed from ledgesand only 3 eggswere present.This date probablymarked the commencementof egg-laying.If an incubationperiod of 33 daysand a nestlingperiod of 21 daysare assumed(Tuck 1960, Hedgren and Linnman 1979), these young shouldhave fledged about 20 July. Back-calculatingthe latest fledgingdates, I estimatethe lasteggs were laid about 23 June. There wasno discernibledifference between species in breedingphe- nology.The fledgingof CommonMurres in the main colony,however, seemedto be more synchronousthan elsewhereon the island. There was also evidencethat egg-layingmay have occurred later within the main colony(see below). Turr•overand survival in po•ds.--Lacking information on rates of breedingsuccess or the total productionof chicks,I cannotestimate the proportionof fledglingsthat remainedin the pondslong enoughto be banded.There islittle question,however, that the majorityof fledglings of both speciesreached the seawithin hoursof leavingthe cliff and were thereby excludedfrom this study. Thick-billedMurres showeda greatertendency to linger in the ponds than did Common Murres. More than 70% of Thick-billed Murres that remainedin the pondslong enoughto be bandedstayed at leastanother full day; 23c/cstill remainedafter 3 days(Fig. 3). By comparison,only 24% of CommonMurres were recapturedone or more times. 270] S.A.Hatch j. FieldOmithal. Summer 1983 Common(rl =352)Murre •.•Thick-billed (n = 22) Murre I 25 5O i i No. Days After First Capture FmURE 3. Length of residencyand mortalityof chicksin pondsafter fledging. Most chicksreceived no food while they stayedin the ponds.They lost weight at a mean rate of 17.7 g/day (samplesfor both species combined)and facedan increasedprobability of deatheach day. Of 374 chicksbanded on or before 12 August, 18% were later recovereddead. In contrast,Thick-billed Murres had cumulativemortality of nearly50% (Fig. 3). These data suggestThick-billed Murres are behaviorallyless capableof dealing with the unusualcircumstances that have existedon Middleton Island sincethe earthquake.Possibly this is a contributing factor in their failure to increase their numbers. A few chickswere fed while in the pondsand surviveda week or more in fresh water, either maintainingweight or showinglimited gl*owth. An adult wasobserved delivering a meal to one suchchick. This