USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.33breed on the Faral - PDF document

USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.33breed on the Farallon Islands in the Pacific Ocean (CommonMurre, Pigeon Guillemot, CassinÕs Auklet, Rhinoceros Auklet,and Tufted Puffin) are presented by Ainley (1990), Ainleyand others (1990a, b, c) and Boekelheide and others (1990).Four inshore fish feeding alcids of the northern Pacific Ocean(KittlitzÕs Murrelet, Pigeon Guillemot, Spectacled Guillemot,and Marbled Murrelet) are reviewed by Ewins and others(1993) (also see Marshall 1988a for a review of the MarbledMurrelet). The Ancient Murrelet, another inhabitant of thenorthern Pacific Ocean, has been reviewed by Gaston (1992).Alcids that nest in small, loosely-aggregated colonies, asisolated pairs, or in areas less accessible to researchers, havenot been well studied. For instance, the reproductive biologyof CraveriÕs and KittlitzÕs murrelets and Spectacled Guillemotsis largely unknown. Although Marbled Murrelets have receivedconsiderable attention during the last two decades, thereproductive ecology of this species is not well understood.Unlike many other alcids, Marbled Murrelets do not nest inconspicuous colonies on cliffs, in rock crevices, or in burrowsin the ground. Instead, this species nests on the alpine tundraor in the upper canopy of old-growth coniferous trees (Hamerand Nelson, this volume b; Marshall 1988a). Additionally,Marbled Murrelets are secretive around their nests and activeduring low light periods at dawn and dusk. Consequently,few nests have been located and observed, and few quantitativedata have been collected.This paper summarizes the reproductive ecology of theauk family and specifically compares Marbled Murrelets tothe other alcids. Such a comparison may allow for a betterunderstanding of the reproductive strategy of Marbledconservation and management of this species.Nest Sites and ColonialityThe nest sites of all alcids have been described, althoughfew nests of some species have been located (e.g., KittlitzÕsand Marbled murrelets). Murres and Razorbills nest primarilyon cliff ledges or in crevices or caves. The nests of Commonand Thick-billed murres are in the open whereas those ofRazorbills are typically partially or fully enclosed (Byrd andothers 1993; Harris and Birkhead 1985). Puffins andRhinoceros Auklets nest in burrows they excavate. Addition-ally, nests of these species are found in rock crevices (Tuftedand Horned, on the level ground of forested islands (RhinocerosAuklet), and among boulders and rocks of islands lackingsoft substrate for burrowing (Atlantic Puffin) (Byrd andothers 1993; Hatch and Hatch 1983). The guillemots nest incracks and crevices of cliffs, among stones or boulders, in Research Wildlife Biologists, Oregon Cooperative Wildlife Research Current address: Postdoctoral Research Associate, Pacific Northwest Plautus impennis]). Reviews of four auks thatChapter 3Comparative Reproductive Ecology of the Auks(Family Alcidae) with Emphasis on the Marbled Murrelet several thousand pairs. CraveriÕs Murrelets probably nest inBreeding Site, Nest Site, and Mate FidelityDe Santo and NelsonChapter 3Reproductive Ecology of Auks Table 1ÑExtant members of the family Alcidae Common name(s)Scientific nameDovekie (Little Auk)...............................................Razorbill (Razorbill Auk)........................................Common Murre (Common Guillemot)...................Thick-billed Murre (BrunnichÕs Guillemot)............Black Guillemot......................................................Spectacled Guillemot (Sooty Guillemot)................Pigeon Guillemot.....................................................Marbled Murrelet....................................................KittlitzÕs Murrelet....................................................XantusÕ Murrelet.....................................................CraveriÕs Murrelet...................................................Ancient Murrelet.....................................................Japanese Murrelet (Crested Murrelet).....................Crested Auklet.........................................................Least Auklet............................................................Whiskered Auklet....................................................CassinÕs Auklet........................................................Parakeet Auklet.......................................................Rhinoceros Auklet (Horn-billed Puffin)..................Tufted Puffin...........................................................Horned Puffin..........................................................Atlantic Puffin......................................................... aNomenclature according to American OrnithologistsÕ Union (1983) USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.35 SpeciesBreeding siteMate fidelityDegree of coloniality yesyessmall to large colonies, scattered pairsyesprobablysmall to large coloniesyesprobablysmall to large coloniesprobably?small to large coloniesyesyessmall to large colonies, loose aggregations, scattered pairs??small to medium colonies, solitarilyyesprobablysmall, loose aggregations, medium colonies, isolated pairsprobably?probably in loose aggregations; probablysolitarilypossibly?solitarilyyesyessmall to large coloniesprobablyprobablyprobably in loose aggregations and scattered pairsyespossiblysmall to large colonies??small to medium coloniesyesyessmall to large coloniesyesyessmall to large colonies??small to medium coloniesyesprobablysmall to large coloniesyes?small, loose to large coloniesyes?small to large colonies, solitarilyyes?small to large coloniesyes?large coloniesyesyessmall to large colonies De Santo and NelsonChapter 3Reproductive Ecology of Auks USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. as cited by Harris and Birkhead 1985). Murray and others(1983) observed that 64 percent of XantusÕ Murrelets retainedthe same nest sites for two years, and Roby and Brink (1986)found that 91 percent of Least Auklets used the same nestentrance in two consecutive years.At least six alcids show mate fidelity (table 2). Divorcerates have been reported to be approximately 24 percent forCrested Auklets (Jones 1993a) and approximately 7 percentfor Black Guillemots and Atlantic Puffins. These figureswere confirmed by Harris and Birkhead (1985). Emslie andothers (1992) have shown that mate retention has a positiveinfluence on reproduction of CassinÕs Auklets; both fledgingand breeding success were higher for pairs that practicedmate retention.No studies have been conducted on individually marked,breeding Marbled Murrelets, but indirect evidence suggeststhat they show both mate and breeding site fidelity. Murreletsare primarily observed in groups of two throughout the year,and many groups include a male and female (Carter 1984,Sealy 1975c). Strong and others (1993) observed at-sea groupsof murrelets in spring and summer and reported that of 4918groups, 55 percent consisted of pairs. The possibility existsthat these twosomes were mated pairs, although withoutobservations of marked birds this is speculative. MarbledMurrelet activity has been documented in the same foreststands for periods up to 18 years (Divoky and Horton, thisvolume), and murrelet nests have been found in the sametrees (Nelson and others 1994; Nelson and Peck, in press;Singer and others, in press), and on the same general locationof tundra (Simons 1980), in consecutive years. Theseobservations suggest breeding site fidelity. Reuse of nestshas recently been documented for the ground nesting KittlitzÕsMurrelet, a close relative of the Marbled Murrelet (Naslundand others 1994).Adult Life History CharacteristicsHistorically, the Great Auk, which became extinct inthe 1800s, was the largest member of the Alcidae, ca. 5 kg(Harris and Birkhead 1985). At present, the murres are thelargest alcids (ca. 1 kg). Fifteen alcids are small by comparison,having body masses less than half that of the murres (table3). The Marbled Murrelet has a mass of 220 g, approximately22 percent that of the murres.Adult annual survival has been estimated for ten species(table 3). The lowest estimates of this population parameterare 75 percent reported for both the Least Auklet, the smallestalcid (ca. 85 g), and 77 percent for the Ancient Murrelet, arelatively small alcid (ca. 200 g), (fig. 1, r2 = 0.45, )De Santo and NelsonChapter 3Reproductive Ecology of Auks De Santo and NelsonChapter 3Reproductive Ecology of Auks SpeciesMean bodyAnnualLongevityAge firstBreeding frequencySurvival of (yr) 164???1/season620906, 6, 304-61/season10048920, 26, 324-61/season9419122??4068412, 202-8+most annually and 1/season490???50 pct. annually48780-909, 11, 14+3-41/season and probably not every year221????224????167?9?most annually and probably 1/season151????2067753-41/season183????27286?possibly 31/season86754.5 predicted3most annually and 1/season121???1/season177865, 10, 202-41-2/season297???1/season533?6, 7?probably 1/season773?6??612????4609413, 203-8+ (most at 5) (1978), USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. murrelets typically have clutch sizes ) = 0.10, De Santo and NelsonChapter 3Reproductive Ecology of Auks De Santo and NelsonChapter 3Reproductive Ecology of Auks SpeciesMean eggEgg mass as pct.Clutch size range mass (g)adult body mass(average)31 (calculated)19190141ca. 11012110010-1215012-131-2 (1.83)56111-2 (1.60)53111-2 (1.76)351813415137221-2 (1.70)35231-2 (1.88)46221-2 (1.99)36221-2 (1.80)36141182212716138ca. 14178ca. 15193121ca. 60ca. 10161ca. 131 Reviewed by Harris and Birkhead (1985), Evans (1981)2Reviewed by Harris and Birkhead (1985)3Reviewed by Harris and Birkhead (1985)4Reviewed by Harris and Birkhead (1985)5Reviewed by Harris and Birkhead (1985); Cairns (1987), Divoky and others (1974)6Reviewed by Ewins and others (1993); Kitaysky (1994), Thorensen (1984)7Reviewed by Ewins (1993), Ewins and others (1993); Kuletz (1983), Nelson (1987)8Hirsch and others (1981), Nelson and Hamer (this volume a), Sealy (1974, 1975b), Simons (1980)9Reviewed by Day and others (1983); Sealy (1975b)10Murray and others (1983)11DeWeese and Anderson (1976), Schšnwetter (1963)12Reviewed by Gaston (1992); Gaston (1990), Gaston and Jones (1989), Jones (1992), Sealy (1975b, 1976), Vermeerand Lemon (1986)13Ono (1993), Ono and Nakamura (1993), Schšnwetter (1963)14Reviewed by Jones (1993a); BŽdard (1969b)15Jones (1993b), Piatt and others (1990), Roby and Brink (1986)16Freethy (1987), Williams and others (1994)17Ainley and others (1990a), Manuwal (1979), Vermeer and Lemon (1986)18Sealy (1972), BŽdard (1969b)19Ainley and others (1990c), Freethy (1987), Sealy (1972), Wilson and Manuwal (1986)20Reviewed by Boone (1986); Ainley and others (1990c), Sealy (1972)21Freethy (1987), Sealy (1972)22Reviewed by Harris and Birkhead (1985) USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. SpeciesIncubatingIncubationMean duration ofEgg neglect shift (hours)incubation (days)both1229?both12-2436?both12-2433probably notboth12-2432very infrequentlybothca. 1-329yes??ca. 28?both2-4 but up to 1728yesbothca. 2427-30 (probable range)yes for several hrs to 1 day????both24-144, most at 7234yes for 1-19 daysboth???both48-12034yes for 1-3 daysboth24-72?yes for 5 daysboth?35possiblyboth2432yesboth24ca. 35?both2439very infrequentlyboth?35?both2445yes for 1-3 daysboth?44???41?both2-50range 35-45yes frequently De Santo and NelsonChapter 3Reproductive Ecology of Auks De Santo and NelsonChapter 3Reproductive Ecology of Auks SpeciesMean hatchingMean fledgingJuvenile 6577?789332798830738534666827???7067?6745????33?????91�90ca. 505076?6366?8281?86100?75806565??8169?6370?7670?72730.4-13.3 observed, aIncludes replacement eggs for Common Murre, Razorbill, Thickbilled Murre, and Pigeon Guillemot, and possiblyfor Black Guillemot, and Atlantic and Horned puffins; does not include second broodsbFledging is defined as departure from the nest to the ocean1Reviewed by Harris and Birkhead (1985); Evans (1981), Stempniewicz (1981)2Reviewed by Harris and Birkhead (1985), Hudson (1985)3Reviewed by Harris and Birkhead (1985), Hudson (1985); Ainley (1990), Boekelheide and others (1990), Murphy(1994); Hatch and Hatch (1990b); also see Byrd and others (1993)4Reviewed by Harris and Birkhead (1985), Hudson (1985); Hatch and Hatch (1990b); also see Byrd and others (1993)5Reviewed by Harris and Birkhead (1985), Hudson (1985); Cairns (1981), Divoky (1994, pers. comm.)6No information located7Reviewed by Ewins and others (1993); Ainley and others (1990b), Kuletz (1983), Nelson (1987); also see summaryby Ewins (1993)8Nelson and Hamer (this volume b)9No information located10Drost (1994), Murray and others (1983)11No information located12Gaston (1990, 1992), Rodway and others (1988), Vermeer and Lemon (1986)13Ono (1993), Ono and Nakamura (1993)14Knudtson and Byrd (1982), Piatt and others (1990), Sealy (1982); also see Jones (1993a)15Jones (1992), Piatt and others (1990), Roby and Brink (1986), Sealy (1982); also see Jones (1993b)16Knudtson and Byrd (1982), Williams and others (1994)17Ainley and others (1990a), Manuwal (1979), Thorensen (1964), Vermeer and Cullen (1982), Vermeer and LemonSealy (1984)19Vermeer and Cullen (1979), Watanuki (1987), Wilson and Manuwal (1986)20Reviewed by Byrd and others (1993)21Reviewed by Byrd and others (1993)22Reviewed by Barrett and Rikardsen (1992), Harris and Birkhead (1985), Hudson (1985); Barrett and Rikardsen (1992),Nettleship (1972) USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. and Birkhead 1985 for review], Pigeon Guillemots [Ainleyand others 1990b], CassinÕs Auklets [Ainley and others 1990a;Manuwal 1979], Horned Puffins [Wehle 1983]) layreplacement eggs. Egg replacement in murres has beenreported to be between 15 and 43 percent (reviewed byBoekelheide and others 1990; Byrd and others 1993). Tenpercent of CassinÕs Auklet pairs replaced naturally lost eggs,and 54 percent replaced eggs removed by researchers(Manuwal 1979). Hatching and fledging success ofreplacement clutches was often lower than first clutches(Ainley and others 1990a; Byrd and others 1993; Manuwal1979; Murphy 1994). The incidence of egg replacement islow for Least and Crested auklets (Piatt and others 1990)and XantusÕ Murrelets (Murray and others 1983) andapparently does not occur in Ancient Murrelets (Sealy 1976).CassinÕs Auklet is the only alcid known to lay a secondclutch following the rearing of their first brood (Ainley andothers 1990a; Manuwal 1979). Hatching and fledging successof second clutches were usually lower than those of firstclutches (Ainley and others 1990a). It is not known if MarbledMurrelets lay replacement eggs or if they attempt to raisemore than one brood per season.Figure 5ÑRelationship between mean adult mass and mean hatchlingmass for 18 alcids (see tables 3 and 7 for values).Figure 4ÑRelationship between mean egg mass and mean hatchlingmass for 18 alcids (see tables 4 and 7 for values).Development and Survival of the Young = 0.98, ) = 0.91, )De Santo and NelsonChapter 3Reproductive Ecology of Auks USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.43Table 7ÑCondition of alcid chicks at hatching and age at which homeothermy (uniform body temperature maintained nearly SpeciesDevelopmentalPlumageMean bodyPct. adultAge (days) ofstage at hatchingmass (g)mass athomeothermy semi-precocialdowny21132-5intermediatedownyca. 609-109-10intermediatedowny55-95 (range)6-1010intermediatedownyca. 6579-10semi-precocialdownyca. 40ca. 101-4semi-precocialdowny40 (n=1)8?semi-precocialdowny3881semi-precocialdowny3315probably 1-2semi-precocialdowny???precocialdowny2415probably 1-2precocialdowny???precocialdowny31132precocialdowny???semi-precocialdownyca. 2510probably 4-5semi-precocialdowny1112-14probably 5semi-precocialdowny1311probably by 7semi-precocialdowny19113-4semi-precocialdowny???semi-precocialdowny5710?semi-precocialdowny648?semi-precocialdowny5910?semi-precocialdowny47116-7 De Santo and NelsonChapter 3Reproductive Ecology of Auks USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. SpeciesBroodingPeriod ofFeeding parentTime at which young parentbrooding (days)reach independenceboth2-7both at nest; probably neither at seaat fledgingboth5-10both at nest; male at seaseveral weeks following fledgingbothuntil fledgingboth at nest; male at sea70-85 daysbothuntil fledgingboth at nest; male at sea?both3-5both at nest; neither at seaat fledging????bothat least 3both at nest; neither at seaat fledgingboth0.5-3.0both at nest; probably neither at seaat fledging??both at nest?both1-2neither at nest; both at sea???both at sea?both2neither at nest; both at sea42-56 days??neither at nest; both at sea?both7both at nest; neither at seaat fledgingboth7both at nest; neither at seaat fledging?probably 7both at nestprobably at fledgingboth3-5both at nest; neither at seaat fledgingboth?both at nest?both4both at nestprobably at fledging????????both9both at nest; neither at seaat fledging and others (1992, in press)De Santo and NelsonChapter 3Reproductive Ecology of Auks De Santo and NelsonChapter 3Reproductive Ecology of Auks SpeciesMean fledgingMean body massMean pct. age (days)at fledging (g)adult mass2712067-8018ca. 17020-3021170-270 (range)18-2822180193735686ca. 33545 (1 obs.)1113846095probably 27-401496729possibly 90 (1 obs.)possibly 401-224142-4??22612-131-2??33ca. 24580-1002987104probably 39-4210692431539035235795232961494906938400654627169 1Reviewed by Gaston (1985), Harris and Birkhead (1985); Evans (1981)2Reviewed by Gaston (1985), Harris and Birkhead (1985); Lloyd (1979)3Reviewed by Gaston (1985), Harris and Birkhead (1985); Hatch and Hatch (1990a)4Reviewed by Gaston (1985), Harris and Birkhead (1985), Hudson (1985); Birkhead and Nettleship (1981), Hatch andHatch (1990a)5Reviewed by Gaston (1985), Harris and Birkhead (1985), Hudson (1985); Cairns (1981, 1987)6Kitaysky (1994), Kondratyev (1994), Thorensen (1984)7Reviewed by Ewins (1993), Gaston (1985); Ainley and others (1990b), Drent and others (1964), Kuletz (1983)8Reviewed by Gaston (1985); Hirsch and others (1981), Nelson and Hamer (this volume a), Nelson and Hardin (1993a),Nelson and Peck (in press), Sealy (1974, 1975a), Simons (1980), Singer and others (1992, in press)9Day and others (1983), Naslund and others (1994)10Reviewed by Gaston (1985); Murray and others (1983)11Reviewed by DeWeese and Anderson (1976), Gaston (1985)12Gaston (1992), Jones and Falls (1987), Sealy (1976), Vermeer and Lemon (1986)13Reviewed by Gaston (1985), Ono and Nakamura (1993)14Reviewed by Gaston (1985); Jones (1993a), Piatt and others (1990)15Reviewed by Gaston (1985); Piatt and others (1990), Roby and Brink (1986)16Reviewed by Byrd and Williams (1993); Williams and others (1994)17Reviewed by Gaston (1985); Ainley and others (1990a), Manuwal (1979), Thorensen (1964), Vermeer and Cullen(1982), Vermeer and Lemon (1986)18Sealy and BŽdard (1973)19Reviewed by Byrd and others (1993), Gaston (1985); Ainley and others (1990c), Leschner (1976), Vermeer (1980),Vermeer and Cullen (1979), Wilson and Manuwal (1986); also see Bertram (1988)20Reviewed by Gaston (1985); Ainley and others (1990c), Boone (1986), Vermeer and Cullen (1979), Wehle (1980)21Reviewed by Gaston (1985), Wehle (1983); Sealy (1973c)22Reviewed by Gaston (1985), Harris and Birkhead (1985); Barrett and Rikardsen (1992), Harris and Hislop (1978),Nettleship (1972) USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. = 0.68, )developmental modes is not significant [ = 0.19, )and others, this volume; Varoujean and Williams, this volume).De Santo and NelsonChapter 3Reproductive Ecology of Auks De Santo and NelsonChapter 3Reproductive Ecology of Auks USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.