Mating System of the Dusky Antbird, a Tropical Passerine, As Assessed by Dna Fingerprinting

Home , Antbird, Cercomacra, Dusky antbird

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The Condor 99.5 12-S 14 0 The Cooper Ornithological Society 1997

MATING SYSTEM OF THE DUSKY ANTBIRD, A TROPICAL PASSERINE, AS ASSESSED BY DNA FINGERPRINTING

ROBERT C. FLEISCHER, CHERYL L. TARR, EUGENE S. MORTON, ALEXANDRA SANGMEISTER AND KIM C. DERRICKSON National Zoological Park, Smithsonian Institution, Washington, DC 20008

Abstract. We studied the genetic mating systemof PanamaProvince, Republic of Panamaduring the non- the Dusky Antbird (Cercomacra tyrannina) in Panama breeding (dry) season from 1991 through 1995, and using multilocusDNA fingerprinting.We found no ev- during the breeding seasonin 1993 and 1994. Dusky idence of extra-pair fertilization (EPF) in 15 offspring Antbirds were common in second-growth habitat, of nine families. We also found no evidence of intra- reaching densities of 3 per hectare (Morton et al., un- specific brood parasitism (ISBP) for 13 offspring in publ. data). Observationsand playbacksof Dusky Ant- eight families. bird song were used to establish the identities of ter- ritorial occupants(see Morton and Derrickson 1996 for Key words: Cercomacratyrannina, Dusky Antbird, DNA jngerprinting, extra-pair fertilizations, tropical more details on field methods). Once we established birds. that a pair occupied a territory, the area was system- atically searched for nests. Many empty nests were found, possibly becauseof prior predation. We found Assessmentsof mating systems based solely on be- that Dusky Antbirds, like most tropical passerinespe- havioral observations,even reasonably detailed ones, cies, have complete clutch sizes of two or fewer eggs. may not accurately represent genetic mating systems When a nest containing eggs or young was located, in natural populations of birds. This knowledge has we used song playbacks to lure adults into mist nets arisen largely from the recent application of powerful placed near the nest. Blood samples from adults and molecular methods, primarily multilocus DNA finger- nestlings were taken from the brachial artery (Wing- printing (see Fleischer 1996 for review). In songbirds, field and Famer 1976) and a volume of lysis buffer 70-80% of socially monogamousspecies studied us- (TE-SDS) approximatelyequal to the volume of whole ing DNA and/or protein methods have shown clear blood was added. Predationwas very high and all eggs evidence of extra-pair fertilization (EPF; Westneat left in the nest initially in our 1993 seasondisappeared 1990, Birkhead and Moller 1992, Stutchburyand Mor- within 2-3 days. This forced us to remove eggs from ton 1995). nests (after catching and bleeding parents) and to in- Most studiesof songbirdshave dealt with Temperate cubatethem long enough so that sufficient DNA could Zone breeders. The only studiesknown to us of trop- be obtained from the embryos. DMSO lysis buffer ical, socially monogamous birds (Telecky 1989, (Seutin et al. 1991) was used to store these tissue sam- Fleischer et al. 1994, Robertson and Kikkawa 1994) ples. Samples were transportedon ice and stored at have revealed high mate fidelity (0% EPF). We predict -20°C. that most tropical, behaviorally monogamous song Whole genomic DNA was isolated as in Fleischer birds also will exhibit genetic monogamy becausein et al. (1994) and subjectedto multilocus DNA finger- over 60% of such species (Morton 1979, 1980) pairs printing using the Jeffreys 33.15 and 33.6 probes (Jef- are sedentary year-roundon territories, maintain long- freys et al. 1985) and standardmethods (see Loew and term pair bonds, and nest asynchronouslywith other Fleischer 1996 for specific protocols). pairs (Stutchbury and Morton 1995). We present here Following the approachof Westneat(1990) the two resultsof multilocusDNA fingerprintingthat reveal no adults defending a nest were excluded from co-par- EPF or intra-specificbrood parasitism(ISBP) in a sam- entage if an offspring’s profile containedthree or more ple of clutchesof the Dusky Antbird (Cercomacra tyr- fragmentsthat could not be assignedto either putative annina), a tropical, sedentaryand behaviorally monog- parent (non-attributable bands). Band sharing coeffi- amous passerine. cients (S of Lynch 1990) were calculatedfrom a pres- METHODS AND MATERIALS ence/absencematrix of same-sized fragments for pu- Dusky Antbirds were marked with unique color band tative related and unrelated individuals. To assessthe combinationsin Parque National Soberania,Gamboa, possibility that our data could not reveal EPFs, we sub- stitutedthe nearestputative unrelatedadult for one parent (the sham) and then counted the number of non- i Received 29 August 1995. Accepted 15 January attributablefragments and calculatedS (Fleischer et al. 1997. 1994).

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methods to the assessmentof genetic mating sys- the evolution of vocal soundsused at close range. tems in vertebrates,p. 133-161. In J. D. Ferraris Proc. Int. Ornithol. Congr. 18:737-741. and S. R. Palumbi [eds.], Molecular zoology: ad- MORTON,E. S., AND K. C. DERRICKSON.1996. Song vances, strategiesand protocols.Wiley-Liss, New ranging by the Dusky Antbird, Cercomacra tyr- York. ann.ni ranging without song learning. Behav. FLEISCHER,R. C., C. L. TARR,AND T. K. PRATT.1994. Ecol. Sociobiol. 39: 195-201. Genetic structureand mating system in the Palila, ROBERTSON,B. C., AND J. K~KKAWA.1994. How do an endangered Hawaiian honeycreeper, as as- they do it?-Monogamy in Zosterops lateralis sessedby DNA fingerprinting. Mol. Ecol. 3:383- chlorocephala. J. Ornithol. 135:459. 392. SEUTIN,G., B. N. WHITE,AND I? 'I BOAG. 1991. Pres- JEFFREYS,A. J., V WILSON,AND S. L. THEIN. 1985. ervation of avian blood and tissue samples for Hypervariable ‘minisatellite’ regions in human DNA analyses.Can. J. Zool. 69:82-90. DNA. Nature 314:67-73. STUTCHBURY,B., ANDE. S. MORTON.1995. The effect of breeding synchrony of extra-pair mating sys- LOEW, S., AND R. C. FLEISCHER.1996. Multilocus tems in songbirds.Behaviour 132:675-690. DNA fingerprinting, p. 456-461. In J. D. Ferraris TELECKY,T 1989. The breeding biologv and mating and S. R. Palumbi [eds.], Molecular zoology: ad- __ system of the Common Myna (Acridotheres tris- vances, strategiesand protocols. Wiley-Liss, New tis). Ph.D. diss., Univ. Hawaii, Honolulu. York. WESTNEAT,D. E 1990. Genetic parentage in the In- LYNCH,M. 1990. The similarity index and DNA fin- digo Bunting: a study using DNA fingerprinting. gerprinting. Mol. Biol. Evol. 7:478-484. Behav. Ecol. Sociobiol. 27:67-76. MORTON,E. S. 1979. A comparative study of avian WESTNEATD. E, l? W. SHERMAN,AND M. L. MORTON. social systemsin northern Venezuelan habitats,p. 1990. The ecology and evolution of extra-pair 233-259. In J. Eisenberg [ed.], Vertebrateecology copulation in bird<: Current Omithol. 7:330-369. in the northern Neotropics. Smithson. Inst. Press, WINGFIELD.J. C., AND D. S. FARNER.1976. Avian en- Washington,DC. docrinology-field investigations and methods. MORTON,E. S. 1980. The ecological backgroundfor Condor 78:570-573.

The Condor 99~514-519 Q The CooperOmlthological Society 1997

ON THE BEHAVIOR OF THE BLACK SWIFT ’

MANUEL MAR~N Museum of Natural Science and Department of Zoology and Physiology, Louisiana State University, Baton Rouge, LA 70803 and Western Foundation of Vertebrate Zoology, Camarillo, CA 93012, e-mail: [email protected]

Abstract. The behavior of Black Swifts was stud- then most roostedon the cave walls. During incubation ied in southern California from 1990 to 1992. Four and early brooding, an adult always remained at the types of aerial interactions were distinguished: (a) nest, and food transfer between adults was observed group chase, (b) pair chase, (c) pair contact, and (d) during this period. touch and grasp. The latter two interactions can be Key words: behavior, aerial and hostile displays, intraspecificor interspecific.Aerial copulationwas not begging, roosting, Cypseloides niger. observed. Nestlings, from age 18 days onward, and adultsgave hostile or deterrencedisplays by wing-rais- ing. Begging by nestlings was silent but aggressive Swifts as a group are remarkably uniform in shape, toward the adult. Silent begging may be an antipre- which makes them difficult to identify, and their aerial dator strategyfor a speciesthat producesa single chick lifestyle makes them difficult to observe. As a result, per season. Nestlings have a far more conspicuous little is known about their behavior, even for common white facial marking than adults; this may function as species. This is especially true in the New World a target signal to guide food delivery in the dimly lit swifts. nesting conditions and as an aid for the adult to find The subfamily Cypseloidinae contains 12-l 3 spe- the nest when arriving late at night. Adults roostedon cies, most of which are tropical or semitropicalin dis- the nest for the first half of the nestling period and tribution. All are similar in body shape,and most have uniform dark plumage and facial markings, which in some are distinctive. All speciesin the subfamily nest I Received 8 March 1996. Accepted 29 November in sites behind waterfalls, in caves, or deep gorges,on 1996. sea cliffs, and in sea caves. Moisture, inaccessibility