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Puede La Ley De Lanchester Ayudar a Explicar La Dominancia Interespecõâ®Ca En Manuscript Received 31 July 2003; Accepted 8 Jan- Aves? Uary 2004

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Puede La Ley De Lanchester Ayudar a Explicar La Dominancia Interespecõâ®Ca En Manuscript Received 31 July 2003; Accepted 8 Jan- Aves? Uary 2004 SHORT COMMUNICATIONS 395 ment composition in the American Gold®nch. SAFRAN, R. J., AND K. J. MCGRAW. 2004. Plumage col- Comparative Biochemistry and Physiology B 131: oration, not length or symmetry of tail-streamers, 261±269. is a sexually selected trait in North American Barn MEISTER, A. 1965. Biochemistry of the amino acids. Swallows. Behavioral Ecology, in press. 2nd ed. Academic Press, New York. SAINO, N., R. STRADI,P.NINNI,E.PINI, AND A. P. MILTENBERGER, R. J., R. L. MYNATT,B.D.BRUCE,W. MùLLER. 1999. Carotenoid plasma concentration, O. WILKISON,R.P.WOYCHIK, AND E. J. MICHAUD. immune pro®le, and plumage ornamentation of 1999. An agouti mutation lacking the basic do- male Barn Swallows (Hirundo rustica). American main induces yellow pigmentation but not obesity Naturalist 154:441±448. SIEFFERMAN, L. M., AND G. E. HILL. 2004. Structural in transgenic mice. Proceedings of the National and phaeomelanin colors indicate male quality in Academy of Sciences 96:8579±8584. the Eastern Bluebird (Sialia sialis). Behavioral MùLLER,A.P.,AND T. A. MOUSSEAU. 2001. Albinism Ecology 14:855±861. and phenotype of Barn Swallows (Hirundo rusti- STRADI, R. 1998. The colour of ¯ight. Solei Gruppo ca) from Chernobyl. Evolution 55:2097±2104. Editoriale Informatico, Milan, Italy. NEEDHAM, A. E. 1974. The signi®cance of zoochro- STRADI, R., G. CELENTANO, AND D. NAVA. 1995. Sep- mes. Springer-Verlag, New York. aration and identi®cation of carotenoids in bird's OEHME, H. 1969. Vergleichende untersuchungen uber plumage by high-performance liquid chromatog- die farbung der vogeliris. Biologisches Zentral- raphyÐdiode-array detection. Journal of Chro- blatt 88:3±35. matography B 670:337±348. È OLIPHANT, L. W. 1988. Cytology and pigments of non- VOLKER, O. 1934. Die Abhangigkeit der Lipochrom- melanophore chromatophores in the avian iris, p. bildung bei VoÈgeln von P¯anzlichen Carotinoi- 65±82. In J. T. Bagnara [ED.], Advances in pig- den. Journal fuÈr Ornithologie 82:439±450. ment cell research. Liss, New York. WAKAMATSU, K., AND S. ITO. 2002. Advanced chemi- cal methods in melanin determination. Pigment OLIPHANT,L.W.,AND J. HUDON. 1993. Pteridines as Cell Research 15:174±183. re¯ecting pigments and components of re¯ecting WAKAMATSU, K., S. ITO, AND J. L. REES. 2002. The organelles in vertebrates. Pigment Cell Research usefulness of 4-amino-3-hydroxyphenylalanine as 6:205±208. a speci®c marker of phaeomelanin. Pigment Cell OLIPHANT, L. W., J. HUDON, AND J. T. BAGNARA. 1992. Research 15:225±232. Pigment cell refugia in homeothermsÐthe unique WARHAM, J. 1975. The crested penguins, p. 189±269. evolutionary position of the iris. Pigment Cell Re- In B. Stonehouse [ED.], Biology of the penguins. search 5:367±371. University Park Press, Baltimore, MD. The Condor 106:395±400 q The Cooper Ornithological Society 2004 CAN LANCHESTER'S LAWS HELP EXPLAIN INTERSPECIFIC DOMINANCE IN BIRDS? ERIN L. SHELLEY,MAILE Y. U. TANAKA,ANNA R. RATNATHICAM AND DANIEL T. B LUMSTEIN1 Department of Organismic Biology, Ecology, and Evolution, University of California, 621 Young Drive South, Los Angeles, CA 90095-1606 Abstract. We studied the applicability of Lanches- more abundant. Further analysis of the most abundant ter's laws of combat to explain interspeci®c dominance species revealed that it was less likely to visit the lo- in birds. We focused on 10 species of Australian birds cust trap when larger, more dominant heterospeci®cs in the arid zone of New South Wales that foraged at were present. Body size, and not numerical superiority, an established locust trap. Consistent with the ``linear seems to be an important determinant in interspeci®c law,'' larger species usually dominated smaller species foraging decisions in birds. in one-on-one encounters. We found no support for the Key words: body size, foraging behavior, group ``N-square law,'' which predicted that large numbers size, interspeci®c competition, Lanchester's laws. of smaller species could dominate larger species when ¿Puede la Ley de Lanchester Ayudar a Explicar la Dominancia Interespecõ®ca en Manuscript received 31 July 2003; accepted 8 Jan- Aves? uary 2004. Resumen. Hemos estudiado la aplicabilidad de las 1 Corresponding author: E-mail: [email protected] leyes del combate de Lanchester en explicar la domi- 396 SHORT COMMUNICATIONS nancia interespecõ®ca en aves. Hemos focalizado este We observed the behavior of birds foraging for in- estudio en 10 especies de aves australianas de la zona sects at a locust trap in the arid zone of southeastern aÂrida de New South Wales, las cuales se alimentaron Australia. We studied interspeci®c interactions to de- en trampas de insectos establecidas para tal ®n. Con- termine the effect of (1) species type, (2) body size, sistente con la ``ley lineal,'' las especies de mayor ta- and (3) abundance on interspeci®c dominance rank. In manÄo usualmente dominaron a las especies maÂs pe- doing so, we evaluated Lanchester's law's ability to quenÄas en los encuentros uno a uno. No encontramos explain nonfatal aggression and dominance in this as- evidencia que apoye la ``ley cuadraÂtica,'' la cual pre- semblage. Although the interactions between these dice que un gran nuÂmero de especies de pequenÄo ta- species are not immediately lethal, the ephemeral and manÄo podrõÂan dominar a especies de tamanÄo mayor patchy nature of resources in the arid zone increase the cuando las primeras son maÂs abundantes. Posteriores importance of locating and obtaining food. Should in- anaÂlisis sobre la especie maÂs abundantes revelaron que terspeci®c interactions interfere with an individual's la probabilidad de visita a las trampas de insectos es ability to obtain food, we can envision a large ®tness menor cuando individuos heteroespecõ®cos maÂs gran- cost. des y maÂs dominantes estaÂn presentes. El tamanÄo cor- poral, y no la superioridad numeÂrica, parece ser un METHODS importante factor en las decisiones de forrajeo en las We observed birds foraging at an insect light trap set aves. up by the Australian Plague Locust Commission for monitoring and researching plague locusts on the Fow- ler's Gap, New South Wales sheep station (31859S, Competition for resources within multispecies systems 1428429E). This locust trap, though technically an ar- leads to the establishment of dominance hierarchies in ti®cial food source for birds in the area, was in oper- which certain species outcompete others (Wallace and ation from 1976±1988, and 1994 to present. The trap Temple 1987, Travaini et al. 1998, Sandlin 2000). In consisted of a 3.0-m-diameter circular metal tub posi- some cases, body size is the primary factor that deter- tioned 0.6 m above the ground and ®lled with 0.15 m mines interspeci®c rank (Kohda 1991, Daily and Ehr- of water. Trees and brush were present nearby. We po- lich 1994, Bassett 1995, McGlynn 2000). However, in sitioned ourselves on a platform 22.3 m from the locust other instances, the relative number of individuals, in- trap. Directly above the center of the tub, a bright light dependent of body size or behavior, may affect inter- was turned on nightly. Insects, most commonly moths speci®c dominance (Basset 1997, Burger and Gochfeld (Order Lepidoptera), were drawn to and disoriented by 1984, Creel 2001, Chapman and Kramer 1996, Mc- the light, whereupon they fell into the water and col- Glynn 2000). lected on the surface. Additionally, a large number of The study of human warfare has led to the devel- water insects such as water-boatmen and backswim- opment of many different military strategies. In 1916, mers (Hemiptera) and diving beetles (Coleoptera) in- F. W. Lanchester published his theory of combat, habited the trap. Although the relative abundance of which included two mathematical models which have insects in the trap seemed to vary each morning, a become known as the ``linear law'' and the ``N-square variety of insects were present each day. All observed law'' (Lanchester 1916). The linear law predicts that bird species foraged on these insects on multiple oc- in one-on-one combat, the stronger (i.e., larger) indi- casions, although not all were exclusively insectivo- vidual will win. The N-square law predicts that larger- rous (e.g., honeyeaters). numbered groups can overcome smaller-numbered During 53 hr of morning observations between 21 groups, with differences in individual body size being April and 3 May 2001, we continuously recorded ev- of little or no importance. Aside from military appli- ery dominant±subordinate interspeci®c interaction that cations, small businesses have used Lanchester's strat- occurred at the locust trap as a win or a loss. At the egies to take over markets dominated by larger busi- time of the interaction, the recorder noted the species ness (e.g., Yano 1995), a scenario which is at least of the winner and loser, the time of the interaction, and super®cially much like animals competing for access the number of other individuals of each species present to a limited resource. Recently, biologists have begun at the trap. Presence was prede®ned as being perched to test the applicability of these laws to a variety of atop or inside the main tub of the trap or the light animal species engaged in mortal combat. In 1993, canister, ¯ying in the immediate airspace, (within 0.3 Franks and Partridge were the ®rst to show that Lan- vertical m of the surface of the trap), or sitting atop a chester's models could explain the outcome of inter- small post connected to one edge of the tub. Domi- actions among slave-making and army ants. Additional nant-subordinate interactions consisted of pecking at, studies of interspeci®c competition among ants chasing, supplanting, or any other interactions in (McGlynn 1999, 2000) provide further support for the which the subordinate individual of one species moved linear and N-square laws.
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