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Genetic Monogamy in Long-Eared Owls ’ 854 SHORT COMMUNICATIONS DEL HOYO, J., A. ELLIOT, AND J. SARGATAL.1996. adox in the Buff-breasted Sandpiper. Am. Nat. Handbook of the birds of the world. Vol. 3. Hoa- 149:1051-1070. tzin to auks. Lynx Editions, Barcelona. LANK, D. B., C. M. SMITH,0. HANOTTE, T A. BURKE, DEWSBURY,D. A. 1982. Ejaculate cost and male AND E COOKE.1995. Genetic polymorphism for choice. Am. Nat. 119:601-610. alternativemating behaviourin iekking male Ruff, DUNNING,J. B., JR. 1993. CRC handbook of avian Philomachusougnax. Nature 378:59-62. body masses.CRC Press, Boca Raton, FL. MARTIN,P A., T.-J.-REIMERS,J. R. LODGE,AND E? J. EMLEN,S. T, l? H. WREGE,AND M. S. WEBSTER.1998. DZIUK. 1974. The effect of ratios and numbersof Cuckoldry as a cost of polyandry in the sex-role- spermatozoa mixed from two males on propor- reversed Wattled Jacana,Jacana jacana. Proc. R. tions of offspring. J. Reprod. Fert. 39:251-258. Sot. Lond. B 256:2359-2364. MILLER, A. P., AND J. V. BRISKIE.1995. Extra-pair pa- GAGE, M. J. G. 1994. Associationsbetween body size, ternity, sperm competition and the evol&ion-of mating pattern, testis size and sperm lengths testis size in birds. Behav. Ecol. Sociobiol. 36: acrossbutterflies. Proc. R. Sot. Lond. B. 258:247- 357-365. 254. OLSSON,M., T MADSEN,AND R. SHINE.1997. Is sperm really so cheap? Costs of reproduction in male GAGE, M. J. G. 1998. Mammalian sperm morphome- try. Proc. R. Sot. Lond. B. 265:97-103. adders, Vipera berus. Proc. R. Sot. Lond. B 264: GOMENDIO,M., ANDE. R. S. ROLDAN.1991. Sperm size 455-459. and sperm competition in mammals. Proc. R. Sot. PARKER,G. A. 1970. Sperm competition and its evo- lutionary consequencesin the insects. Biol. Rev. Lond. B 243:181-185. 45:525-567. HALLIDAY,T, ANDS. J. ARNOLD.1987. Multiple mat- PARKER,G. A. 1982. Why are there so many tiny ing by females: a perspective from quantitative sperm? Sperm compe&tion and the main&an& genetics.Anim. Behav. 35:939-941. of two sexes. J. theor. Biol. 96:281-294. HAR&SON,C. 1975. A field guide to the nests, eggs PIERCE,E. P, AND J. T LIFJELD.1998. High paternity and nestlings of British and European birds. Col- without paternity-assurancebehavior in the Purple lins, London. Sandpiper, a species with high paternal invest- HARRSION,C. 1978. A field guide to the nests, eggs ment. Auk 115:602-612. and nestlings of North American birds. Collins, PITELKA,E A., R. T. HOLMES,AND S. E MCLEAN. 1974. London. Ecology and evolution of social organization in HARVEY,P. H., AND M. D. PAGEL.1991. The compar- arctic sandpipers.Am. Zool. 14:185-204. ative method in evolutionary biology. Oxford PURVIS.A.. ANDA. RAMBAUT.1994. Comoarativeanal- Univ. Press, Oxford. ysis by independent contrasts(CAIk), version 2. HEG, D., B. J. ENS, T. BURKE,L. JENKINS,AND J. I? Oxford Univ., Oxford. KRUIJT. 1993. Why does the typically monoga- SZEKELY,T, ANDJ. D. REYNOLDS. 1995. Evolutionary mous Oystercatcher(Haematopus ostralegus) en- transitionsin parental care in shorebirds.Proc. R. gage in extra-pair copulations? Behaviour 126: Sot. Lond. B 262~57-64. 247-289. - - VAN RHIJN, J. G. 1991. The Ruff: individuality in a LANCTOT, R. B., K. T. SCRIBNER, B. KEMPENAERS, AND gregarious wading bird. T & A.D. Poyser, Lon- P J. WEATHERHEAD.1997. Lekking without a par- don. The Condor 101:854-X59 0 The Cooper Ornithological Society1999 GENETIC MONOGAMY IN LONG-EARED OWLS ’ JEFFREY S. MARKS Montana CooperativeWildlife ResearchUnit, Universityof Montana, Missoula, MT 59812, e-mail: [email protected] JANISL. DICKINSON AND JOSEPH HAYDOCK HastingsNatural History Reservationand Museum of VertebrateZoology, Universityof California, 38601 East Carmel Valley Road, Cannel Valley, CA 93924 Abstract. We used DNA fingerprinting to study Owls (Asio otus). We detected no extra-pair fertiliza- genetic parentagein socially monogamousLong-eared tions (EPFs) in 59 nestlings from 12 nests. One of these nestswas solitary, but the other 11 had from one to five pairs of owls nesting simultaneouslywithin 30 I Received 5 May 1999. Accepted 26 July 1999. to 250 m. Thus, despite the presumablyhigh potential SHORT COMMUNICATIONS 855 for extra-pair matings, the Long-eared Owls that we hypothesis that EPFs occur in Long-eared Owls. We studied were genetically monogamous. In addition, also use band sharing coefficients from adults to de- based on low band sharing among adults, we found no termine whether nesting aggregationsare composedof evidence that nesting aggregationswere composed of close relatives. close relatives. Genetic monogamy appearsto be the rule for socially monogamousraptors. We suggestthat METHODS the high rate of male parental effort in raptors selects STUDY AREA AND FIELD METHODS againstEPFs becausefemales that engage in extra-pair activities risk losing parental investment by males The study areas were in Lake and Ravalli Counties, whose confidence in paternity is reduced owing to the western Montana, in isolated groves of quaking aspens behavior of their mates. (Populus tremuloides), black hawthorns (Crataegus douglasii), and willows (Salix sp.). The Paul Smith Key words: Asio otus, DNAfingerprinting, genetic study area (Lake County) had six Long-eared Owl monogamy, Long-eared Owl. nests in 1997 and five in 1998. The Victor study area (Ravalli Countv) was 150 km south of Paul Smith and Studies of mating systems based on genetic markers had four nests& 1998. The Airport study area was 14 have shown that cooulationsoutside the oair bond are km east of Paul Smith and had one nest in 1997 and common in many bird species (Westneat et al. 1990). two close-nestingpairs in 1998. Indeed, genetic monogamy seems to be the exception We captured adults at night in mist nets placed in rather than the rule in socially monogamouspasserines natural flight paths to the nest or set in front of flight- (Gowaty 1996). Among socially monogamous non- less young that had recently left the nest. Thus, we passerines,however, extra-pair fertilizations (EPFs) are were reasonablycertain that these owls were the social less common. A recent review showed that EPFs have parentsat the nests where they were captured.We de- occurred in 86% (42/49 species) of passerinesthat termined the sex of capturedadults based on presence have been studied but in only 48% (11/23 species)of or absenceof an incubationpatch. Adults were banded non-passerines(Westneat and Sherman 1997). On both with U.S. Fish and Wildlife Service metal leg bands. empirical and theoretical grounds, the occurrence of Young were banded at three to four weeks of age, just EPFs in birds is influenced by the density and syn- before or shortly after they left the nest. chrony of breeding pairs (Siutchbury and Morton 1995, Johnson and Burlev 1998). In addition. the BLOODSAMPLING AND DNA FINGERPRINTING amount of parental care provided’ by males m& be To determine whether nesting density is associated positively c&related with- their certainty of pateinity with EPFs in Long-eared Owls, we conductedparen- (Mflller and Birkhead 1993). tal-exclusion analyses using multilocus DNA finger- Because Long-eared Owls (Asio otus) sometimes printing. This permitted us to evaluatebackground lev- nest close to one another (Marks et al. 1994), they els of genetic variation in the populationof Long-eared representan important test case for the hypothesisthat Owls in western Montana and to perform paternity- genetic parentageis associatedwith breeding density. exclusion analysis for 59 nestlings from 12 nests dur- Male parental effort is extremely high in all speciesof ing two years of study. Blood samples(100 pL) were owls becausemales provide nearly all of the food to taken from the brachial vein of adults and nestlingsat the female and nestlings from courtship until at least the time of banding. Samples were placed in Queen’s mid-way through the brood-rearing period (Marks et lysis buffer (Seutin et al. 1991) in the field and then al. 1999), although they do not incubate or brood. tiansferred to the lab, where they were frozen at Thus, one might expect owls to be genetically monog- -20°C. We digested 5 to 10 UP of nuclear DNA with amous independentof proximate factors that could in- HaeIII and eviluated the quaiiry and concentrationof fluence mating systems(breeding density and breeding DNA in the restriction digests on a 1% agarose test synchrony). gel stainedwith ethidium bromide. Samplescontaining Investigators have used DNA fingerprinting to as- 5 kg of digested DNA in Ficoll loading buffer were sess relatednessand genetic parentagein three previ- loaded on 27-30 cm agarosegels (1%) aid run along- ous studies of owls. Galeotti et al. (1997) compared side a I-kb ladder at 40 V for three davs. The social genetic similarity within and between communal win- parents were run in lanes adjacent to <heir offspring ter roosts of Long-eared Owls and concluded that (Fig. 1). In addition, we ran sets of breeding adults (n roosts were not composed of close relatives. Lawless = 22 dyads) to obtain the backgroundlevel of band et al. (1997) found no evidence of EPFs in Eastern sharing within the population. DNA in the gel was Screech-Owls (Otus asio). which are highlv territorial denaturedin 0.5 M NaOWl.5 M NaCl and transferred and nest solit&y. In cbntrast, Johnso; (1997b) re- in fresh denaturing solution to a charged nylon filter ported a low level of EPFs (2 of 31 nestlings) in a (Zetaprobe). Filters were UV-crosslinked and hvbrid- colony of Burrowing Owls (Athene cunicularia), ized overnight at 63-65°C in DEPC-treated 1X-SSC, which nest below ground and do not defend large nest- 1% SDS. 1% Blotto. and then washed at 65°C in 0.75- ing territories.
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