Differential allocation of reproductive effort to territorial establishment and maintenance by male yellow warblers (Dendroica petechia)

Summary We tested the hypothesis that the basis of the variation in reproductive strategy among male yellow warblers (Dendroica petechia) is a tradeoff in the allocation of reproductive effort to intrasexual competition (territorial effort) and to parental care (parental effort). Since a negative correlation between level of parental effort and amount of brown streaking on the breast (plumage score) has already been demonstrated in this species, this study looked for evidence of a positive correlation of territorial effort with plumage score and a negative correlation with parental effort. Analysis of the spatial and temporal use of territories, prior to egg-laying, by males with different plumage scores supported the prediction that brighter (higher plumage score) males allocate more effort to territorial establishment and maintenance (Fig. 1). Male plumage score was positively and highly significantly correlated with six different measures of the relative amount of time spent, rate of energy expended, and risk of injury assumed by each male during territorial activities. The level of territorial effort was also found to be negatively correlated with the level of parental effort expended by the same males later in the season, confirming that there is a tradeoff in allocation of reproductive effort to these two major components. Further analysis revealed that territory quality was positively correlated with male plumage score (Figs. 3 and 4), while average nestling growth rate and other indices of reproductive success were not (Fig. 6). These results suggest that increased territorial effort by brighter males enables them to occupy higher quality territories where high levels of parental effort are not necessary to maintain high levels of reproductive success. Since reproductive success was not correlated with plumage score, this study further supports the differential allocation hypothesis that different males are using alternative strategies for the allocation of reproductive effort.     

Landscape Patterns of Florida Scrub Jay Habitat Use and Demographic Success

Remote sensing and geographical information systems are used to analyze landscapes important to species of conservation concern. The accuracy of the methods depends on how closely habitat mapping classes are linked to population demography. Habitat use by Florida Scrub Jays ( Aphelocoma c. coemlescens ) was quantified using circular plots. Habitat variation was mapped using high-resolution aerial photography on a site where all Florida Scrub Jays were color-banded. Nest site selection, nest success, yearling production, and breeder survival were measured within Florida Scrub Jay territories. Habitat use was lowest in areas without scrub oaks or areas within 136 m from forests. Open oak, dominated by scrub oaks and open sandy areas, had the highest use and nest success among habitats. Open oak occurred as narrow patches ( <20 m wide) in landscapes dominated by matrix habitat (palmetto-lyonia and swale marshes). Most wide patches (>50 m) of open oak were potential population sources, where reproduction exceeded mortality. Areas with patches of open oak of less than 1 ha were usually population sinks. Open oak occurred as less flammable patches in a landscape subject to frequent fires. Population sources varied temporally and spatially with fires and site potential to support scrub oaks (soils). Analyses of landscape patterns and dynamics indicated that habitat mapping should not only include patches of currently optimal habitat but should also include landscapes associated with open oak. The influences of landscape patterns on habitat use, reproductive success, survival, and territory size can be quantified at different scales starting with attributes associated with habitat patches, nest sites, and territories. Potential mapping errors occur, however, when habitats are used to quantify the areal extent of sources and sinks and similar population attributes important for species persistence.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

Sexual selection and variation in reproductive strategy in male yellow warblers (Dendroica petechia)

Summary Male yellow warblers (Dendroica petechia) exhibit extensive variation in the amount and conspicuousness of the sexually distinctive brown streaking on the breast. We investigated this intraspecific variation in degree of sexual dichromatism to see if male plumage rank (essentially the amount of brown streaking) is correlated with the amount of reproductive effort allocated to parental investment, as sexual selection theory would predict. As predicted, the average rate of feeding visits to the nest by males was negatively correlated with the plumage rank of the male in 1982 (Fig. 1) and 1983 (Fig. 2), and varied little between years or among study areas. Within one study area, the higher nest visit rate of dull (low plumage rank) males was correlated with higher nestling growth rates. But in the other two study areas, where bright (high plumage rank) males predominated, nestlings of bright males tended to have the highest growth rates despite minimal nest visit rates. Thus, overall there was no correlation between male plumage rank and nestling growth rate, a potential index of relative reproductive success, in either year (Figs. 3 and 4). This overall equal nest success likely helps to maintain the behavioral and plumage polymorphism. Since we could find no evidence that the high nestling growth rates of bright males were due to an adjustment of female parental effort to compensate for low male parental effort, there must be some other benefit of being bright which contributes to nest success. To address this, we used the relationship between parental feeding rates and nestling growth rates for each nest as an index of relative territory quality. This analysis suggests that bright males generally occupy the best territories available, possibly because they are more aggressive in obtaining and defending them than dull males. Dull males appear to be relegated to poorer territories, where an increase in male parental contribution would be necessary to achieve high nestling growth rates. We propose that different males are using alternative but evolutionarily stable reproductive strategies, in a way that is consistent with the predictions of sexual selection theory. Allocation of limited reproductive effort to parental investment decreases as sexual dichromatism increases, which probably reflects an increased investment in intermale competition for the best territories.     

Delayed maturation in plumage colour: Evidence for the female-mimicry hypothesis in the kestrel

Summary In many sexually dichromatic species, young males have female-like plumage during their first potential breeding year. The female-mimicry hypothesis (FMH) supposes that by possessing female-like plumage young males deceive older conspicuous males into believing that they are females, thus reducing competition from adult males. The status-signalling hypothesis (SSH) supposes that adult males can distinguish sex, but postulates that young males reduce competition from adult males by reliably signaling low status with their dull plumage. We tested these hypotheses in the European kestrel (Falco tinnunculus). Female-like young males settled to breed closer to adult males than did other adult males (Figs. 1a, b). By settling near adult males, young males seemed to increase their chance of mating with adult females. Adult female-young male pairs had better reproductive success than yearling-yearling pairs. These results suggest that there is an adaptive value in possessing a female-like plumage colour in the breeding season. To test the FMH, we measured sexual preference of adult males when adult females and young males were simultaneously shown in an aviary. Adult males were unable to recognize sex, because in half the cases they preferred young males (Fig. 3). However, when adult males and females were shown simultaneously, males preferred females (Fig. 2). Our results support the FMH rather than the SSH, because young males successfully deceived older males by their plumage.     

Nonideal breeding habitat selection: a mismatch between preference and fitness

The selection of breeding sites in heterogeneous habitats should ideally be based on cues closely reflecting habitat quality and thus predicting realized individual fitness. Using long-term population data and data on territory establishment of male Northern Wheatears (Oenanthe oenanthe), we examined whether territory characteristics linked to individual fitness (reproductive performance and survival) also were linked to territory preference. Breeding territories varied in their physical characteristics and their potential effects on reproductive performance, and this variation among territories was correlated from one year to the next. Of all measured territory characteristics (from the focal and the previous year) only territory field layer height predicted individual fitness, i.e., reproductive performance was higher in territories with permanently short rather than growing field layers. Territory preference, instead, was only linked to the size of territory aggregations, i.e., males settled earlier at territory sites sharing borders with several adjacent sites than at those with few or no adjacent sites. This mismatch between territory characteristics linked to fitness and those linked to territory preference was not explained by site fidelity or compensated for by the different fitness components measured. Because the results were not in agreement with an ecological trap scenario, where poor habitats are preferred over high-quality habitats, our results suggest a more general case of nonideal habitat selection. Whereas nonideal selection with respect to territory field layer height may be explained by its poor temporal predictability within the breeding season, the preference for territory aggregations is still open to alternative adaptive explanations. Our study suggests that nonideal habitat selection should be investigated by direct estimates of preferences (e.g., order of territory establishment) and their links to habitat characteristics and fitness components. Furthermore, we suggest that the probability of establishing a territory needs to be included as a factor influencing patterns of habitat selection.     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

Female pied flycatchers Ficedula hypoleuca choose male characteristics in homogeneous habitats

Summary The paper reports the results of a 2-year study of pairing success of male pied flycatchers in a homogeneous habitat. A handicapping experiment was carried out in which certain wing and tail feathers were removed from randomly selected males. Handicapped males had reduced pairing success, they lost weight, and they sang less frequently than control males. Male pairing success was positively correlated with the darkness of the plumage, body-size, and previous breeding experience. Earlier studies on the same species have failed to detect any relationships between pairing success and male characteristics, possibly because of habitat heterogeneity and variation in nest site quality. The evolutionary basis for female choice of male characteristics is discussed. There are reports that males with attractive traits (e.g. black plumage) provide a high quality of parental care. However, the fact that male pairing success was related to male conspicuousness makes it difficult to discriminate between active and passive female choice.     

Elaborately ornamented males avoid costly parental care in the house finch (<Emphasis Type="Italic">Carpodacus mexicanus</Emphasis>): a proximate perspective

Selection should favor flexibility in reproductive tactics when the combination of sexual traits and reproductive behaviors that achieve the highest fitness differs between males within a population. Understanding the functional significance of variation in male reproductive tactics can provide insight into their evolution. Male house finches (Carpodacus mexicanus) in a Montana population display continuous variation in parental tactics: males with more elaborated (redder) plumage color provide little or no parental care compared to less elaborated (dull) males. Here, we first determined whether elevation of prolactin (a pituitary hormone) was related to variation in male parental tactics and, second, we used the relationship between prolactin levels and parental behavior to investigate why redder males avoid a high investment in parental care. We found that prolactin elevation was closely associated with paternal care. In addition, males with redder plumage color had low prolactin levels, whereas dull males, which provision twice as frequently, had high levels of prolactin. We also found that male condition was unrelated to plumage color but negatively related to prolactin levels. These results suggest that the low provisioning of redder males was not due to physiological constraints, but instead reflected a tactic to avoid the costs associated with parental care. The condition benefits accrued by redder males may explain their higher post-breeding survival compared to dull males. Moreover, dull males were previously shown to have higher pairing success than redder males, suggesting that the relationship between male plumage color and parental care may reflect individually optimized parental tactics.Communicated by H. Kokko     

Effects of Clearcutting on Habitat Use and Reproductive Success of the Ovenbird in Forested Landscapes

We studied Ovenbirds ( Seiurus aurocapillus ) in northern New Hampshire during 1992 and 1993 to determine whether edge-related changes in habitat use and reproductive success reported in fragmented landscapes exist in predominantly forested landscapes. Six study plots were placed adjacent to four recent clearcuts (2.1–5 ha) and extended 400 m into the forest interior. Nests, territories, and territorial males obtaining mates were equally distributed in edge (0–200 m) and interior (201–400 m) areas. Nest survival was higher in the forest interior in 1992 and for 1992 and 1993 combined. The proportion of pairs fledging ≥ 1 young, fledgling weight, and fledgling wing-chord did not differ between edge and interior in either year. Number of young fledged per pair was slightly lower in edge areas, but these differences were not significant. We conclude that clearcutting in extensively forested landscapes can affect Ovenbird reproductive success. Nevertheless, the effect on Ovenbird populations is moderated by the abundance of mature forest cover in the region and by the tendency of Ovenbirds to renest after initial nest failure.     

Negative Effects of Habitat Loss on Survival of Migrant Warblers in a Forest Mosaic

Abstract: Habitat loss and fragmentation in forested landscapes often negatively affect animal abundance; however, whether these factors also affect fitness is not well known. We hypothesized that observed decreases in bird occurrence and abundance in landscapes with harvested forests are associated with reduced apparent survival of adults. We defined apparent survival as an estimate of survival that accounts for an imperfect resighting probability, but not permanent emigration (i.e., dispersal). We examined the association between spatially extensive habitat loss and apparent survival of males of 2 Neotropical migrant species, Blackburnian Warbler (Dendroica fusca) and Black‐Throated Green Warbler (D. virens), over 7 years in the Greater Fundy Ecosystem, New Brunswick, Canada. We estimated apparent survival among and within breeding seasons. We quantified amount of habitat in the context of individual species. In this landscape, boundaries between land‐cover types are gradual rather than clearly identifiable and abrupt. Estimated apparent within‐season survival of both species decreased as a function of amount of habitat within a 2000‐m radius; survival was approximately 12 times (95% CI 3.43–14) greater in landscapes with 85% habitat than in landscapes with 10% habitat. Apparent annual survival also decreased as a function of amount of habitat within a 100‐m radius. Over the range of habitat amount, apparent annual survival decreased 15% (95% CI 7–29%) as the amount of habitat decreased. Our results suggest that reduced species occurrence in landscapes with low proportions of habitat is due partly to lower apparent survival at these sites. This mechanism operates both directly (i.e., via effects on mortality or dispersal during breeding) and possibly through indirect effects during the nonbreeding season. Habitat loss was associated not only with a lower number of individuals, but also with lower survival of those individuals.     

Territory dynamics in a sage sparrow population: are shifts in site use adaptive?

Summary In 1981–1985, we studied territory dynamics in a southeastern Idaho sage sparrow population to evaluate the hypothesis that territory shifts represent adaptive adjustments in site use. We predicted that shifts should (1) result in changes in territory characteristics, (2) be influenced by previous reproductive success and result in greater success, and (3) decline in magnitude for individual males over time. Habitat features of territories changed little as a result of shifts, but territory size increased. Correspondingly, habitat features of territories were unrelated to reproductive success, whereas territory size was positively related. The magnitude of territory shifts was negatively correlated with preshift fledging success, and after shifts, males experienced greater average reproductive success than before. Successive territory shifts by individual males became progressively smaller. Thus, territory shifting by sage sparrows seems to be adaptive behavior aimed primarily at increasing territory size.     

Territorial resource valuation deviates from habitat quality in an ecologically trapped,long-distance migratory bird

Preference measures are widely used in habitat selection studies to test an animal’s choice relative to particular habitat features, but most measures are subject to criticism as they fail to indicate the underlying behavioral motivation. Order of settlement on breeding sites has been proposed as an effective measure in migratory organisms, as it conceptually approaches a choice experiment. We tested the assumption that early red-backed shrikes (Lanius collurio) are more willing to defend their territorial resource than individuals arriving later. We earlier showed that shrikes arriving first settled in forest plantations that resulted in lower reproductive success compared to territories on farmland, suggesting an ecological trap. Therefore, individuals are expected to place higher value on the lower quality sites in forests. Within the context of resource valuation theory in animal contests, we used a simulated territorial intrusion experiment to measure territorial defense and to evaluate the perceived value of the territory during the settlement phase in both habitat types. Males arriving early were much more motivated to defend their territory than late birds. After correction for the disparity in the timing of arrival between habitat types, shrikes also more vigorously defended their territories in the forest habitat associated with the lowest reproductive returns. Although some resource valuation mechanisms remain unclear, our results show that early and late-arriving individuals strongly differ in behavioral motivation to hold their territorial resources. This study also demonstrates for the first time that organisms may exhibit a higher degree of territorial aggressiveness in a lower quality habitat.     

Use of fecundity measured directly throughout the breeding season to test a source-sink demographic model

Populations of landbirds (bird species that occupy terrestrial habitats for most of their life cycle) are declining throughout North America (north of Mexico) and Europe, yet little is known about how demography is driving this trend. A recent model of 5 geographically separated populations of Cerulean Warblers (Dendroica cerulea) that was based on within-season sampling of nest survival and fledgling success shows that all populations are sinks (annual reproduction is consistently less than annual adult mortality). I tested this indirect model by directly measuring fecundity (number of female fledglings/female) during the breeding season for 2 years in a Cerulean Warbler population occupying a mature forest in southwestern Michigan (U.S.A.) I determined territories of male birds on the basis of male plumage characters and phases of the nesting cycle (2007) and on uniquely color-banded males (2008). I transferred locations of identified males to topographic maps. I counted all fledglings in territories from May to July each year. The model I tested may apply only to single-brooded species; therefore, I searched the literature to estimate the percentage of single-brooded species in North America. The breeding season of Cerulean Warblers was short- nearly all nests were initiated from mid-May to late June. Nest predation and brood parasitism were primary and rare causes of nest failure, respectively. Significantly fewer Cerulean Warblers fledged from parasitized than from nonparasitized nests. Fledgling survival required to maintain the population size was well above previously published values for Neotropical migrants. Single-brooded species comprise 62% of North American breeding bird species for which the number of broods per year is known; I believe my results may apply to these species. The consistency between identification of populations as sources or sinks on the basis of either model estimates or direct measurements suggests that a demographic model relying on within-season sampling of fecundity is adequate to determine population status of single-brooded avian populations. In addition, on the basis of results of previous studies, annual adult survival rate of the Cerulean Warbler is typical of parulid warblers that are not declining. Thus, low fecundity, here determined with different quantitative methods, can drive status of landbird species with high-observed survival.     

Ideal dominance distributions: a test using red-winged blackbirds (Agelaius phoeniceus)

Summary The ideal dominance distribution model predicts that competition between individuals of a species for territories will result in socially dominant individuals acquiring territories in higher quality habitat than their subordinates. Although the dispersion and relative reproductive success of male red-winged blackbirds (Agelaius phoeniceus) across habitats in eastern Ontario appears to conform to the ideal dominance distribution model, data from a study of three captive groups comprised of males from both high (marsh) and low (upland field) quality habitats failed to support the prediction that males from marsh habitat are dominant to those from upland habitat. Contrary to the prediction males from uplands were generally dominant to males from marshes. We found a significant positive correlation between dominance and both increased epaulet size and increased body size. Controlling for these positive effects, upland males remained generally dominant to marsh males. Measurements of independent samples of males from both habitats indicated that the overall distribution of males does not conform to ideal dominance. We suggest that the strong between-year territory fidelity shown by male red-winged blackbirds and chance events when they initially acquire territories may contribute to this lack of conformity.     

Plumage characteristics, reproductive investment and assortative mating in tree swallows Tachycineta bicolor

Elaborate ornamental plumage has been associated with various measures of individual quality in many species of birds. Male plumage characteristics, which have been relatively well studied, have been shown to reflect past reproductive investment, as well as the potential for reproductive investment in the current breeding attempt. In contrast, the signalling functions of female traits remain largely unexplored. In this study, we investigated the relationship between plumage attributes of breeding adult tree swallows and past reproductive investment, current reproductive investment and social mate pairing strategy. Both males and older females possess metallic green to metallic blue iridescent plumage on their dorsal surface, making this a suitable species for this type of investigation. We did not find any effects of past reproductive investment and success on the plumage attributes of returning breeders. In contrast, female plumage hue covaried with fledging success, and female plumage brightness was positively associated with mean clutch egg mass. In addition, we found that social pairs mated assortatively with respect to plumage brightness. We argue that since plumage characteristics vary with age in both male and female tree swallows, plumage attributes in this species are indicative of breeding experience and may be honest signals of quality. Positive assortative pairing could be the result of mutual mate choice or intra-sexual competition for nest sites by both males and females.     

Life and Death in the Fast Lane: Demographic Consequences of Road Mortality in the Florida Scrub-Jay

Abstract: We examined the demographic consequences of road mortality in the cooperatively breeding Florida Scrub-Jay (Aphelocoma coerulescens ), a threatened species restricted to the oak scrub of peninsular Florida. Between May 1986 and July 1995 we monitored the survival and reproductive success of a color-banded population of jays along a two-lane highway at Archbold Biological Station. Annual mortality of breeding adults was 0.38 on road territories, significantly higher than the rate of 0.23 for breeders on nonroad territories. High mortality on road territories appeared to be a direct result of automobile traffic per se and not a consequence of road-induced changes in habitat characteristics. Mortality was especially high for immigrants without previous experience living along the road: in their first two years as breeders on road territories, naive immigrants experienced annual mortality of 0.50 and 0.45. From year 3 onward, however, annual mortality dropped to 0.29, not significantly different from the rate for birds on nonroad territories. This experience-dependent decline in road mortality could be caused either by surviving jays learning to avoid automobiles or by selective mortality operating through time (demographic heterogeneity). Proximity to the road had no effect on nesting success beyond its indirect effects on breeder experience and group size. Because the mortality of 30- to 90-day-old fledglings was significantly higher on road territories than on nonroad territories, however, breeder mortality greatly exceeded production of yearlings on road territories. Roadside territories therefore are sinks that can maintain populations of Florida Scrub-Jays only via immigration. Because Florida Scrub-Jays do not avoid roadside habitats and may even be attracted to them, road mortality presents a difficult challenge for the management and conservation of this threatened and declining species.     

Field-testing the protandry-based mating system for the lizard, Anolis carolinesis: does the model organism have the right model?

Using the lizard Anolis carolinensis as its subject, an early and enduring model of psychobiology was presented in which neuroendocrines and social behavior interact to coordinate reproduction between the sexes. The adaptive context for the model was protandry, here defined as a "male-first" emergence pattern from winter dormancy to the onset of breeding. In the protandry scenario, males emerge, become reproductive, and contest for territories. Then, pre-reproductive females emerge to settle on male territories, whereby ovarian recrudescence is facilitated by the behavior of courting males. A collateral inference of the protandry sequence is that females are choosing their future mates from among local males and their territories (i.e., intersexual selection). The model and its assumptions, though never validated by field data, have served as the paradigm for many laboratory experiments on A. carolinensis. Recently, however, field studies described an intrasexually selected mating system for A. carolinensis, without evidence of direct female choice. Differences between the selective inferences from the two perspectives led us to test in the field a number of protandry-based characteristics previously ascribed to A. carolinensis. We monitored free-ranging adults across the winter-to-breeding period using data on sex ratios, social behavior, gut contents, sex steroids, and gonadal condition. We found no evidence of protandry, and little support for any of the associated assumptions. Instead, the sexes broadly overlapped in their temporal transition from winter retreats to an active status in the habitat, with average male reproductive development about 1-2 weeks in advance of females. We replace the traditional protandry-based A. carolinensis paradigm with a realistic onset sequence into the breeding season and a new model for the species' mating system. Our study underscores the need for field validation when laboratory-generated data are fitted to adaptive paradigms.     

Sex-specific differences in site fidelity and the cost of dispersal in yellow-headed blackbirds

Male migratory birds tend to be more faithful than females to previous breeding sites, suggesting sex differences in costs or benefits of dispersal. In Illinois, greater site fidelity by male yellow-headed blackbirds (Xanthocephalus xanthocephalus) was associated with reduced reproductive success the following year for dispersers relative to non-dispersers. Dispersing females suffered no reduction in reproductive success the following year. Males that attracted few social mates, and thus had low reproductive success, were more likely to disperse, whereas females dispersed in response to low-patch reproductive success, regardless of their individual performance. Males that dispersed appeared to be successful acquiring territories because none was observed as a floater. The rate of dispersal by males in this low-density population was greater than in more dense populations where dispersing males may be less successful at acquiring territories. Despite success at obtaining territories, males that dispersed acquired territories on the periphery of wetlands where fewer females nested, resulting in lower reproductive success. In the second year after dispersing, however, males moved onto more central territories where they acquired larger harems. Thus, dispersal by males may be a long-term strategy requiring at least 2 years for benefits to be realized. Long-term success was enhanced because dispersing males moved to wetlands on which reproductive success was higher than on the wetlands they left. In addition to demonstrating that both individual and patch reproductive success affect dispersal decisions, these data indicate that when evaluating costs and benefits of dispersal, researchers should use a time frame beyond 1 year.     

Satellites and sneakers: opportunistic male breeding tactics in pupfish (Cyprinodon pecosensis)

Summary Manipulations of population density, availability of oviposition substrate, and size of breeding habitat affected the proportion of pupfish (Cyprinodon pecosensis) males adopting territorial and satellite breeding tactics. Satellites occupied stations above breeding substrates defended by territorial males. Both territorial and satellite males developed a bright blue breeding coloration. A third breeding tactic, sneak-spawning, was occasionally observed. Sneak-spawning males retained the cryptic female coloration and occasionally spawned on territories. The lower reproductive success of satellites and sneak-spawning males suggested that both are conditional breeding tactics adopted by competitively inferior males. Satellites were present in all treatments that favored a territorial breeding system, but were absent when the breeding system was a dominance hierarchy. Satellites were associated with territorial males that had higher reproductive success and larger territories than males without satellites. Satellites functioned as parasites rather than as mutualists, since they disrupted spawnings and stole copulations from territorial males. The occurrence and frequency of the conditional breeding behaviours in pupfish represent facultative responses of males to changes in the intensity of competition for breeding sites and females.