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1.  Virginity and the clutch size behavior of a parasitoid wasp where mothers mate their sons 
Theoretical and empirical research on the evolution of clutch size has proved to be an extremely productive area of evolutionary biology. A general prediction is that individuals should produce a smaller number of offspring when resources are more limited, such as when multiple individuals compete for the same resources for their development. However, we expect that the opposite prediction arises with virgin females of haplodiploid species, which are subject to extreme local mate competition. We test the key assumption and predictions of this theory with the parasitoid wasp Melittobia australica. Our data demonstrate that there is a trade-off between the size of the first and subsequent clutches and that virgin females adjust their production of sons according to the mating status (mated or not) of cofounding females. We also found that mated females facultatively change their offspring sex ratio in response to the mating status of cofoundresses. We discuss the potential mechanisms used to recognize the mating status and the implications of our results in the context of the extremely female-biased sex ratios observed across Melittobia species..
PMCID: PMC3947723  PMID: 24619355
clutch size; constrained female; local mate competition; Melittobia australica; reproductive strategy; sex allocation
2.  Terminal investment and senescence in rhesus macaques (Macaca mulatta) on Cayo Santiago 
Behavioral Ecology  2010;21(5):972-978.
Long-lived iteroparous species often show aging-related changes in reproduction that may be explained by 2 non-mutually exclusive hypotheses. The terminal investment hypothesis predicts increased female reproductive effort toward the end of the life span, as individuals have little to gain by reserving effort for the future. The senescence hypothesis predicts decreased female reproductive output toward the end of the life span due to an age-related decline in body condition. Nonhuman primates are ideal organisms for testing these hypotheses, as they are long lived and produce altricial offspring heavily dependent on maternal investment. In this study, we integrated 50 years of continuous demographic records for the Cayo Santiago rhesus macaque (Macaca mulatta) population with new morphometric and behavioral data to test the senescence and terminal investment hypotheses. We examined relationships between maternal age and activity, mother and infant body condition, interbirth intervals, measures of behavioral investment in offspring, and offspring survival and fitness to test for age-associated declines in reproduction that would indicate senescence, and for age-associated increases in maternal effort that would indicate terminal investment. Compared with younger mothers, older mothers had lower body mass indices and were less active, had longer interbirth intervals, and spent more time in contact with infants, but had infants of lower masses and survival rates. Taken together, our results provide strong evidence for the occurrence of reproductive senescence in free-ranging female rhesus macaques but are also consistent with some of the predictions of the terminal investment hypothesis.
PMCID: PMC2920293  PMID: 22475990
aging; life history; maternal investment; offspring fitness; reproductive senescence; rhesus macaques
3.  A behavioral mechanism underlying ecological divergence in the malaria mosquito Anopheles gambiae 
Behavioral Ecology  2010;21(5):1087-1092.
Disruptive selection mediated by predation on aquatic immature stages has been proposed as a major force driving ecological divergence and fostering speciation between the M and S molecular forms of the African malaria mosquito, Anopheles gambiae. In the dry savannahs of West Africa where both molecular forms co-occur, the S form thrives in temporary pools filled with rainwater, whereas the M form preferentially breeds in permanent freshwater habitats where predator pressure is higher. Here, we explored the proximal mechanisms by which predation may contribute to habitat segregation between molecular forms using progeny of female mosquitoes captured in Burkina Faso. We show that the S form suffers higher predation rates than the M form when simultaneously exposed to the widespread predator, Anisops jaczewskii in an experimental arena. Furthermore, behavioral plasticity induced by exposure to the predator was observed in the M form, but not in the S form, and may partially explain its habitat use and ecological divergence from the S form. We discuss the role of adaptive phenotypic plasticity in allowing successful colonization of a new ecological niche by the M form and highlight further research areas that need to be addressed for a better understanding of the ultimate mechanisms underlying ecological speciation in this pest of major medical importance.
PMCID: PMC2920295  PMID: 22476108
adaptation; Anopheles gambiae; behavior; habitat divergence; mosquito; notonectidae; phenotypic plasticity, predation; speciation
4.  Color signal information content and the eye of the beholder: a case study in the rhesus macaque 
Behavioral Ecology  2010;21(4):739-746.
Animal coloration has provided many classical examples of both natural and sexual selection. Methods to study color signals range from human assessment to models of receiver vision, with objective measurements commonly involving spectrometry or digital photography. However, signal assessment by a receiver is not objective but linked to receiver perception. Here, we use standardized digital photographs of female rhesus macaque (Macaca mulatta) face and hindquarter regions, combined with estimates of the timing of the female fertile phase, to assess how color varies with respect to this timing. We compare objective color measures (camera sensor responses) with models of rhesus vision (retinal receptor stimulation and visual discriminability). Due to differences in spectral separation between camera sensors and rhesus receptors, camera measures overestimated color variation and underestimated luminance variation compared with rhesus macaques. Consequently, objective digital camera measurements can produce statistically significant relationships that are probably undetectable to rhesus macaques, and hence biologically irrelevant, while missing variation in the measure that may be relevant. Discrimination modeling provided results that were most meaningful (as they were directly related to receiver perception) and were easiest to relate to underlying physiology. Further, this gave new insight into the function of such signals, revealing perceptually salient signal luminance changes outside of the fertile phase that could potentially enhance paternity confusion. Our study demonstrates how, even for species with similar visual systems to humans, models of vision may provide more accurate and meaningful information on the form and function of visual signals than objective color measures do.
PMCID: PMC2892627  PMID: 22475874
color signaling; communication; receiver perception; visual discrimination threshold modeling
5.  Feeding decisions of eastern bluebirds are situationally influenced by fledgling plumage color 
Behavioral Ecology  2010;21(3):456-464.
The relative amount of resources that avian parents provide to individual offspring within a brood represents a strategy that can have large effects on reproductive success. We tested whether parental feeding decisions of eastern bluebirds Sialia sialis are influenced by offspring plumage color by presenting pairs of differently colored fledglings side by side and observing how they were provisioned by parents. After a control period, we manipulated blue plumage color so that one sibling in each trial became relatively dark and one became relatively bright. During neither the control nor the experimental periods did either parent consistently feed naturally brighter or experimentally brightened sons more than drab sons. Under specific circumstances, however, both parents directed a higher proportion of their feeding attempts to more brightly colored sons. Paternal feeding attempts to brighter offspring during both the control and experimental periods increased in relation to the brightness of these fledglings relative to their brothers. Maternal feeding decision, on the other hand, were influenced by numerous variables during control and experimental periods including the date of the trial, the difference in mass between fledglings, the feeding behavior of fathers during the trial, the relative investment by fathers during the nestling stage, and the amount of UV chroma in fledgling plumage. Taken together, these results suggest that equal provisioning of offspring is the strategy most commonly adopted by eastern bluebirds but more brightly colored offspring will be fed preferentially when resources for offspring are limited.
PMCID: PMC2854528  PMID: 22476433
color; juvenal plumage; ornaments; parent–offspring interactions; plumage; relative parental investment; Sialia sialis
6.  Female Lincoln's sparrows modulate their behavior in response to variation in male song quality 
Behavioral Ecology  2010;21(3):562-569.
Sexually reproducing organisms should mate with the highest quality individuals that they can. When female songbirds choose a mate, they are thought to use several aspects of male song that reflect his quality. Under resource-limited environmental conditions, male Lincoln's sparrows (Melospiza lincolnii) vary among one another in several aspects of song quality, including song length, song complexity, and trill performance. In a 2-pronged approach, we tested whether variation in song quality of male Lincoln's sparrows influences the behavior of females that are in a reproductive-like state. Over two trials, we exposed females to songs from the high and low ends of the distribution of naturally occurring song quality variation and found a higher level of behavioral activity in females exposed to high-quality songs, especially when they had first been exposed to low-quality songs. We also examined female phonotaxis toward antiphonally played songs with experimentally elevated and reduced trill performance and found that females moved preferentially toward the songs with elevated trill performance. Contrary to most studies investigating the behavioral responses of wild, female songbirds to variation in male song, we obtained our results without administering exogenous estradiol, which can artificially perturb the female's physiology. Our results demonstrate that the behavior of female Lincoln's sparrows is modulated by the quality of male songs to which they are exposed and that trill performance plays a significant role in this behavioral modulation. Furthermore, as the order of song quality presentation matters, it appears that recent song experience also influences female behavior.
PMCID: PMC2854529  PMID: 22476505
bird song; mate choice; Melospiza lincolnii; phonotaxis; sexual selection; song playback; trill performance; vocal performance
7.  Songbird chemosignals: volatile compounds in preen gland secretions vary among individuals, sexes, and populations 
Behavioral Ecology  2010;21(3):608-614.
Chemical signaling has been documented in many animals, but its potential importance in avian species, particularly songbirds, has received far less attention. We tested whether volatile compounds in the preen oil of a songbird (Junco hyemalis) contain reliable information about individual identity, sex, or population of origin by repeated sampling from captive male and female juncos originating from 2 recently diverged junco populations in southern California. One of the populations recently colonized an urban environment; the other resides in a species-typical montane environment. The birds were field-caught as juveniles, housed under identical conditions, and fed the same diet for 10 months prior to sampling. We used capillary gas chromatography–mass spectrometry to quantify the relative abundance of 19 volatile compounds previously shown to vary seasonally in this species. We found individual repeatability as well as significant sex and population differences in volatile profiles. The persistence of population differences in a common environment suggests that preen oil chemistry likely has a genetic basis and may thus evolve rapidly in response to environmental change. These finding suggest that songbird preen oil odors have the potential to function as chemosignals associated with mate recognition or reproductive isolation.
PMCID: PMC2854530  PMID: 22475692
birds; chemical communication; Junco hyemalis; olfaction; pheromones
8.  Examination of prior contest experience and the retention of winner and loser effects 
Behavioral Ecology  2010;21(2):404-409.
In many animal taxa, prior contest experience affects future performance such that winning increases the chances of winning in the future (winner effect) and losing increases the chances of losing in the future (loser effect). It is, however, not clear whether this pattern typically arises from experience effects on actual or perceived fighting ability (or both). In this study, we looked at winner and loser effects in the jumping spider Phidippus clarus. We assigned winning or losing experience to spiders and tested them against opponents of similar fighting ability in subsequent contests at 1-, 2-, 5-, and 24-h intervals. We examined the strength of winner and loser effects, how long effects persist, as well as how experience affected perceived and actual fighting ability. Our results demonstrate that winner and loser effects are of approximately the same magnitude, although loser effects last longer than winner effects. Our results also demonstrate that previous experience alters actual fighting ability because both the assessment and escalation periods were affected by experience. We suggest that the retention time of experience effects depends on expected encounter rates as well as other behavioral and ecological factors. In systems with short breeding seasons and/or rapidly fluctuating populations, context-dependent retention of experience effects may allow males to track their status relative to the fluctuating fighting ability of local competitors without paying the costs necessary to recall or assess individual competitors.
PMCID: PMC2821427  PMID: 22476369
contest experience; fighting ability; male–male competition; perceived RHP; Phidippus clarus; winner and loser effect
9.  Spatial autocorrelation: an overlooked concept in behavioral ecology 
Behavioral Ecology  2010;21(5):902-905.
PMCID: PMC2920294  PMID: 22476031
GIS; Moran's I; spatial analysis

Results 1-9 (9)