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Cavity-Adoption and the Evolution of Coloniality

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Cavity-Adoption and the Evolution of Coloniality The Condor 104:240±247 q The Cooper Ornithological Society 2002 CAVITY ADOPTION AND THE EVOLUTION OF COLONIALITY IN CAVITY-NESTING BIRDS JESSICA R. EBERHARD1 Smithsonian Tropical Research Institute, Apdo. 2072, Balboa, Republic of Panama Abstract. Among cavity-nesting birds, a distinction can be made between excavators, which dig their own cavities, and cavity-adopters, which nest in pre-existing cavities. His- torically, these two types of species have been grouped together as ``cavity-nesters,'' but it has become clear that the two nesting habits are associated with very different suites of life- history characters. This paper tests the hypothesis that cavity-adopters differ from excavators and other nest-building species in their propensity to evolve colonial breeding. Because of their dependence on pre-existing cavities, cavity-adopters presumably have less control than excavators over the location of their nests, and this could limit their ability to nest near conspeci®cs. A literature survey of the nesting behavior of 842 species in 17 bird families shows that coloniality almost never occurs in species that are obligate cavity-adopters. A phylogeny-based comparative analysis of nesting behavior in the Anseriformes indicates that in this group, colonial breeding has evolved less frequently in lineages of cavity-adopt- ers than would be expected by chance. Together, this evidence supports the hypothesis that colonial breeding systems are unlikely to evolve in lineages of cavity-adopters. Key words: Anseriformes, birds, cavity nesting, colonial nesting, comparative method. EvolucioÂn de la Colonialidad en Aves que Nidi®can en Cavidades Resumen. Las aves que nidi®can en cavidades pueden dividirse en dos grupos: las es- pecies excavadoras, que construyen sus propias cavidades, y las no-excavadoras, que nidi- ®can en cavidades pre-existentes. HistoÂricamente, estos dos tipos de especies han sido agru- padas colectivamente como aves que anidan en cavidades, pero estudios recientes demues- tran que los dos tipos de nidi®cacioÂn estaÂn asociados con diferentes caracteres de sus his- torias de vida. En este estudio se pone a prueba la hipoÂtesis que especies no-excavadoras di®eren de las excavadoras y de otras especies que construyen nidos en cuanto a su pro- pensioÂn a evolucionar haÂbitos de reproduccioÂn colonial. Debido a que dependen de cavi- dades pre-existentes, las especies no-excavadoras presumiblemente tienen menos control que las excavadoras sobre la ubicacioÂn de sus nidos, y esto podrõÂa limitar su habilidad para nidi®car cerca de otros miembros de su especie. Una revisioÂn bibliogra®ca sobre el com- portamiento de nidi®cacioÂn de 842 especies pertenecientes a 17 familias demuestra que casi nunca se observa colonialidad en especies que obligatoriamente utilizan cavidades pre- existentes. Un anaÂlisis comparativo utilizando una ®logenia y datos del comportamiento de nidi®cacioÂn de miembros del orden Anseriformes indica que la colonialidad ha evolucionado con menor frecuencia de lo esperado al azar en linajes de especies no-excavadoras que utilizan cavidades. En conjunto, estos datos apoyan la hipoÂtesis que los sistemas de nidi®- cacioÂn colonial tienen baja probabilidad de evolucionar en linajes de no-excavadoras que anidan en cavidades. INTRODUCTION banks, or in termitaria; cup and domed nests built with materials such as twigs, grass, mud, The great diversity of nests used by birds in- and spider silk; and elaborately woven nests like cludes the huge mounds of soil and decaying those built by weaverbirds, orioles, and oropen- vegetation used by megapodes; cavities in trees dolas (Collias and Collias 1984). Among cavity- or among rocks; burrows in the ground, in nesters, a distinction can be made between ex- cavators (which dig their own cavities) and cav- ity-adopters (which nest in pre-existing cavities). Manuscript received 11 June 2001; accepted 11 Jan- These two types of cavity-nesters have histori- uary 2002. cally been grouped together as having a single 1 Present address: Macaulay Library of Natural Sounds, Cornell Laboratory of Ornithology, 159 Sap- nest type, but it has become clear that the dis- sucker Woods Rd., Ithaca, NY 14850. E-mail: jre24@ tinction is an important one to make when study- cornell.edu ing the evolution of cavity-nesting lineages. [240] EVOLUTION OF COLONIALITY IN CAVITY-NESTERS 241 Excavators differ from cavity-adopters in a of feeding territories, with aquatic habitat, and number of life-history traits, including nesting with nest exposure to predators (Rolland et al. success, clutch size, duration of incubation and 1998). There are several possible advantages to nestling periods, annual productivity, and adult nesting in colonies, including access to and se- survival (Martin and Li 1992). In a study of life- lection of favorable breeding sites, enhanced history traits of North American Passeriformes foraging ef®ciency, improved detection of and and Piciformes, Martin (1995) found that vari- defense against predators, and access to breed- ation in fecundity and adult survival is better ing partners (Siegel-Causey and Kharitonov explained by nest sites, and their associated nest 1990). Breeding pairs probably use the presence predation, than by food limitation. In fact, when or reproductive success of conspeci®cs as cues adult survivorship and annual fecundity are to indirectly assess the commodities necessary compared among species representing an array to breed (e.g., food, safety, health, mates, breed- of nest types, excavators rank among the highest ing sites), and then choose breeding sites that in survivorship and lowest in fecundity, while are near those of successful conspeci®cs (Dan- cavity-adopters generally have low survivorship chin and Wagner 1997). However, even if the and high fecundity (Martin 1995). In the case of above factors favor gregarious nesting, colonia- cavity-adopters, larger clutch sizes may have re- lity can only result if nesting substrates are sulted from the historically limited availability available in large enough patches to accommo- of nest sites rather than from nest failure rates date large groups. Because pre-existing cavities or food availability. Therefore, a consequence of suitable for breeding are not likely to be dense, the dependence on existing cavities may be se- obligate cavity adoption might be expected to lection for increased reproductive effort when a preclude the evolution of group nesting even if nest site is obtained (Martin and Li 1992, Martin selection favored group nesting. 1995). This paper examines the hypotheses that A recent paper by Collias (1997) argued that among cavity-nesting birds, excavators and cav- the diversity of nests built by passerine birds is ity-adopters differ in their propensity to evolve a key to explaining the adaptive radiation of this colonial breeding, and that obligate cavity adop- avian group during the late Tertiary. Collias em- tion tends to block the evolution of coloniality. phasized the importance of the ¯exibility in nest- I review data on the nest types and nest disper- site choice that accompanies the ability to build sion of several avian groups that include large a variety of nest types, and linked this ¯exibility numbers of cavity-nesting species, to examine with the radiation of passerines into new niches. the evidence that coloniality is unusual in cavi- Another context in which ¯exibility in nest-site ty-adopting species. Recognizing the importance choice might be important is in the evolution of of taking phylogenetic relationships into account colonial nesting (e.g., Eberhard 1998), particu- when testing this kind of evolutionary hypoth- larly among cavity-nesters, in which the evolu- esis, I also present a phylogeny-based compar- tion of social behavior may be affected by ative analysis of nesting behavior in the Anse- whether a species is an excavator or a cavity- riformes that tests the hypothesis that coloniality adopter. For coloniality to evolve, one funda- is unlikely to evolve among non-excavating cav- mental condition must be met: breeding pairs ity-adopters. must be able to nest in proximity to other breed- ing pairs. Because they depend on pre-existing METHODS cavities, cavity-adopters have much less control I obtained information on nesting behavior from over the location of their nests than do excava- the literature, and followed Sibley and Monroe's tors, and the breeding densities of obligate cav- (1990) taxonomic classi®cation of birds in tal- ity-adopters can be limited by the availability of lying the data. Rather than attempting to com- nest sites (Brawn and Balda 1988). Flexibility plete an exhaustive survey, I chose avian groups and control over nest placement could allow gre- on the basis of two criteria: that they contain a garious nesting by making it easier for breeding substantial number of cavity-nesting species, pairs to build nests near conspeci®c pairs. and that published information on their nesting A comparative analysis of coloniality in birds behavior be readily available. For each species has shown that coloniality is a relatively labile in the groups examined, I collected the follow- trait, and that it is correlated with the absence ing data: nest type (open nest or cavity nest), 242 JESSICA R. EBERHARD type of cavity-nester (excavator or cavity-adopt- MINSTATE and MAXSTATE functions; see er), and breeding dispersion (solitary or colo- Maddison and Maddison 1992). The observed nial). In species that were variable in a given numbers of gains and losses in coloniality were character, the predominant character state was obtained by mapping the dependent variable used (e.g., solitary nesters that occasionally form (nest dispersion) on the resolved tree; when nec- loose groups were coded as solitary). However, essary, ambiguity was resolved using the DEL- in cases where two character states appeared to TRAN option. For each set of calculations, I re- be equally common in a species, the character corded the probability of having as many or was coded as polymorphic. Species for which more than the observed number of gains in co- data on both nest type and nesting dispersion loniality, and as many or more than zero losses.
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