Os escorpiões são únicos dentre os artrópodes terrestres em muitos aspectos da biologia reprodutiva. A corte em escorpiões envolve sequências complexas de comportamentos ritualizados; é dividida em três fases: iniciação, dança e transferência de espermatozoides. Cerca de 50 espécies de escorpiões dentre as 1600 atualmente reconhecidas tiveram a corte descrita. Do mesmo modo, são poucos os estudos com tamanho de prole e investimento reprodutivo em escorpiões. Até o momento, a descrição da corte em Tityus bahiensis foi realizada com base em poucas observações incompletas. Os objetivos deste trabalho foram: estudar a corte em T. bahiensis para reconhecer os repertórios comportamentais e padrões de comportamento para espécie, através de um etograma e de um fluxograma; investigar prováveis funções das categorias comportamentais; abordar aspectos da época reprodutiva; e estudar o tamanho de prole, tempo de gestação e investimento reprodutivo. Dezenove cortes, de 202 pareamentos realizados, foram utilizadas para elaboração do etograma e do fluxograma. As sequências de comportamentos foram analisadas com o programa JWacher sup TM/sup e compiladas através de um script em Perl. Para o estudo do tamanho de prole, data de nascimento e período de gestação, foram compilados dados de 76 fêmeas. Desses nascimentos, 12 foram provenientes de cortes observadas em laboratório. A corte em T. bahiensis é complexa e está organizada de modo que a realização de uma fase prepara e condiciona o aparecimento da seguinte. A fase de iniciação tem funções como a procura por parceiros sexuais e o reconhecimento específico, sexual e da predisposição dos parceiros à corte. A fase de dança é dividida em dois módulos. No módulo I, o macho estimula e conduz a fêmea a uma superfície adequada para deposição do espermatóforo, preferencialmente uma casca de árvore (X2=24,314; g.l.=5; p<0,001). O módulo II apresenta comportamentos frequentes e repetitivos, com a função de estimular a fêmea e preparála para a fase seguinte. A fase de transferência de espermatozoides é rápida e estereotipada. Tem a função de inseminar e estimular a fêmea para uma cópula bemsucedida. Durante a cópula, o macho executa cortejo copulatório (tateamento com pernas e roçar com quelíceras). A fase de pós-transferência segue um padrão, mas apresenta poucas categorias, sendo que a maioria tem a provável função de romper o flagelo do espermatóforo inserido no gonóporo do macho; o macho pode consumir o espermatóforo e não ocorre canibalismo sexual. As fêmeas de T. bahiensis são capazes de controlar diferentes fases da corte obrigando os machos a utilizar comportamentos de estímulo, ao invés de coerção, para serem aceitos. A exibição de cortejo copulatório pelos machos é um forte indicativo da existência de seleção críptica feminina em T. bahiensis, sendo uma estratégia reprodutiva para estimular a fêmea a aceitar os espermatozoides do macho durante a aquisição do esperma. Apesar da época reprodutiva compreender o ano todo, os meses de maior atividade sexual são novembro a abril. O tamanho de prole na primeira parição é 2-25 filhotes (n=76). Além disso, T. bahiensis é capaz de produzir 1-4 proles com uma inseminação. O tempo de gestação é 2,5-12,8 meses e, apesar da capacidade de parir o ano todo, a maioria das parições ocorreu nos meses quentes, sobretudo, entre novembro e janeiro (X2=164,912; g.l.=11; p<0,001). Adicionalmente, o tamanho de prole é correlacionado com o tamanho corpóreo materno (rs=0,593; p=0,042; n=12). Assim, fêmeas maiores produzem mais filhotes e, portanto, apresentam maior investimento reprodutivo. / Scorpions are unique among terrestrial arthropods in many reproductive biology traits. Courtship involves a complex series of ritualized behaviors; it is divided into three phases: initiation, Promenade à deux, and sperm transfer. Courtship behavior has been described for about 50 of the 1600 extant scorpion species. Likewise, litter size and reproductive investment in scorpions are poorly known. Previous studies on Tityus bahiensis described courtship based on few and incomplete observations. The aims of this work were to study courtship behavior in T. bahiensis in order to recognize behavior repertories and behavior patterns for the species by making an ethogram and fluxogram; investigate probable functions of behavior categories; address aspects of the reproductive season; and study litter size, gestation period, and reproductive investment. Nineteen courtships, out of 202 interactions, were used to make the ethogram and the fluxogram. Behavioral sequences were analyzed with JWacherTM and compiled by a Perl script. For the study of litter size, date of birth, and gestation period, data of 76 females were compiled. Of these, 12 births were obtained from courtship observed in the laboratory. Courtship in T. bahiensis is complex and it is organized in a way that the execution of one phase prepares and regulates the appearance of the next. Initiation phase has functions such as the search for mating partner and species, sexual, and predisposition to court recognition. Promenade à deux phase is divided into two modules. In module I, male stimulates and leads female to a suitable surface for spermatophore deposition, preferentially a bark (X2=24,314; g.l.=5; p<0,001). Module II is characterized by frequent and repetitive behaviors, with the function of stimulating the female and preparing her to the next phase. Sperm transfer phase is rapid and stereotyped. Its function is to inseminate and stimulate the female for a successful mate. During mate, male executes copulatory courtship (feeling and kissing). Post transfer phase has a pattern, but with feel categories; most of them has the probable function of breaking the spermatophore flagellum inserted in the male gonopore; male may consume spermatophore, and there is no mating cannibalism. The females of T. bahiensis are capable of controlling different courtship phases, engaging males to use stimulating behaviors, rather than coercion, to be accepted. The exhibition of copulatory courtship by males strongly indicates the existence of cryptic female choice in T. bahiensis, being a reproductive strategy to stimulate the female to accept male spermatozoids during sperm uptake. Although reproductive season takes place all year long, the months in which reproductive activity is greater are throughout November to April. Litter size at the first parturition is 2-25 offspring (n=76). Moreover, T. bahiensis is capable of producing 1-4 broods with a single insemination. Gestation period is 2,5-12,8 months, and, although the capacity of giving birth throughout the year, most of the parturition occurred in the warm season, specially throughout November to January (X2=164,912; g.l.=11; p<0,001). In addition, litter size is correlated to female body size (rs=0,593; p=0,042; n=12). Thus, larger females produce more offspring and invest more into reproduction.

Roadways are a pervasive feature of northeastern landscapes and can be a significant source of mortality for turtles. Until recently, little has been known about the design requirements for successful under-road passages for turtles and other wildlife to move safely between bisected habitat patches. At outdoor laboratories, using a factorial experimental design, we examined movements in response to varying light levels, and barrier opacity for painted turtles (Chrysemys picta, n=833), Blanding’s turtles (Emydoidea blandingii, n=49), and spotted turtles (Clemmys guttata, n=49). Additionally, we examined tunnel size, tunnel entrance design, and artificial lighting for painted turtles only. All three species responded poorly to a 0% available light treatment. As the amount of natural light transmitted through the tops of tunnels increased, successful completion of the trials increased. Furthermore, turtles generally moved at a slower rate when traveling along a translucent barrier, compared to an opaque one. Our results indicate the importance of designing road passage structures for freshwater turtles that provide adequate tunnel lighting in combination with specific entrance designs that meet the goals of the project.

Whereas circadian rhythms in humans and many other organisms are closely linked to the solar day and even slight deviations could cause significant health problems, several spider species thrive despite extremely shortened or extended circadian clocks. These naturally occurring clocks influence a variety of behaviors, which may help spiders maintain a precarious balance between their conflicting roles as predator and prey. The southern black widow, Latrodectus mactans, is primarily nocturnal in its locomotor activity, but this activity appears only loosely regulated by its circadian clock. This study attempts to determine how internal circadian oscillators affect other aspects of black widow behavior, since preliminary data suggest that they also possess potentially irregular circadian patterns. The behavioral patterns of twenty female black widows were recorded over a span of several weeks. Approximately 3,400 hours of footage in standard (12-hour light-dark cycles) and constant conditions (complete darkness) were analyzed and specific behaviors were recorded using the software BORIS. This study focused on three specific sets of behavior: foraging versus retreat patterns, general activity levels throughout a 24-hr period, and waste disposal. Predation stimuli were also introduced during each cycle set to determine how the widows’ responses were affected by their internal circadian oscillators. Free-run behaviors (behavior no longer entrained to the standard 24hr cycle) observed during the dark-dark cycles differed from the behavior observed during light-dark cycles in both frequency and duration of actions, particularly those related to foraging or web work. Likewise, certain behaviors and prey avoidance techniques observed during light-cycle periods were not observed during constant conditions. This experiment examines several previously unstudied black widow behaviors for generating a better understanding on how they act in natural conditions and to determine how their actions may be influenced by their highly unusual circadian rhythms. Ultimately, this experiment will contribute to a larger, ongoing study investigating circadian-controlled behaviors and rhythms in spiders.

Behavior may be dramatically influenced by changing environments, and differences in light intensity environments may have important behavioral consequences. One approach to understanding changes in behavior is by studying behavioral syndromes, suites of correlated behaviors reflecting between individual consistencies in behavior expressed within a behavioral situation (e.g., correlations between antipredator behaviors in different habitats), or across behavioral contexts (e.g., correlations among feeding, antipredator, or mating behavior) (Sih et al. 2004a). Guppies (Poecilia reticulata) provide a great model system to study behavior. Guppies are small, freshwater tropical fish that inhabit still pools in swift-flowing streams, and the backwaters of small rivers in mountain forest areas of Trinidad (Houde 1997; Magurran 2005). In this study I attempted to answer the following questions using three low predation populations of guppies : 1) Do guppies display a behavioral syndrome for aggression and/or courtship across light situations?; 2) Are there mean level changes in aggression or courtship across light situations?; 3) Are mean level changes influenced by differing social environments that include or lack sexually receptive females?; 4) Are there correlations between behavior and the androgen hormones testosterone and 11-ketotestosterone?; 5) Are there correlations between behavior and the stress hormone cortisol?; 6) Are there mean level changes in hormone release rates across light environments?; and 7) Is flexibility in hormone release rates influenced by social environments that include or lack sexually receptive females? Guppies exhibit behavioral syndromes for both aggression and courtship. Furthermore, guppies exhibit behavioral flexibility for both aggression and courtship, but only in social environments that include sexually receptive females. I found no correlations between behavior and androgen hormones. I also did not find any correlations between behavior and the stress hormone cortisol. Furthermore, I did not find any mean level changes in hormone release rates across light environments. Interestingly, cortisol levels were higher in social environments in which sexually receptive females were absent. Many studies have looked at how the environment influences courtship behavior in guppies, especially employing high predation populations, but few studies have examined aggressive behavior or behavior in general with low predation populations. Furthermore, few studies have determined the role social environments play, and how hormones may interact with behavior. This study is important because it helps illuminate how low predation populations deal with changes in light intensity environments, and adds to what we understand about guppy behavior in general.

Parasites often have profound effects on the survival and evolution of their hosts, and hence on the structure and health of entire ecosystems. Yet basic questions, such as the degree of virulence of a given parasite on its host, and factors influencing which hosts in a population are at the greatest risk of infection, are vexingly difficult to resolve. The western blacklegged tick-western fence lizard (Ixodes pacificus-Sceloporus occidentalis) system is important, primarily because I. pacificus, a vector of the Lyme disease spirochete Borrelia burgdorferi, is dependent on S. occidentalis for blood meals in its subadult stages, and this lizard possesses an innate immune response that removes the Lyme disease pathogen from attached ticks. My study focused on two aspects of the I. pacificus-S. occidentalis interaction. In Chapter 1, I investigated factors correlating with the intensity of I. pacificus infestations on S. occidentalis. Infection intensity (parasites per host) is often highly variable within a host population, though certain individuals, such as males, tend to be more heavily infected. Previous work in the I. pacificus-S. occidentalis system suggests that differences in behavior, such as the frequency of territorial patrols, may contribute to variation in tick intensity among lizards. I therefore hypothesized that lizard traits that correlate with dominance would also correlate with infestation intensity. Specifically, I predicted that larger and more colorful males would have higher infestation intensities than less impressive animals. In this chapter, I also focused on site selection by ticks infesting S. occidentalis. Skin folds on the necks of these lizards (nuchal pockets) may function to divert ectoparasites away from eyes, ears, and other potentially vulnerable structures. I therefore also looked for factors correlating with tick attachment in these pockets. I sampled ticks on adult male S. occidentalis in the spring and summer, which is the seasonal peak for both S. occidentalis territorial behavior and subadult I. pacificus abundance. After determining the site of infestation and intensity of ticks on these lizards, I re-infested lizards with laboratory-reared I. pacificus larvae, and again quantified tick intensity and attachment location. Contrary to expectation, no host traits correlated with tick intensity among ticks naturally infesting lizards, and lab-reared larval intensity was negatively correlated with lizard body size. As expected, ticks acquired by lizards naturally concentrated inside nuchal pockets, and I also observed this site preference among ticks in lab-based experimental infestations. Although the general pattern, lab-reared ticks were more varied in the sites on which they fed. There was a negative correlation between infestation intensity and the proportion of ticks attached in nuchal pockets. Unsurprisingly, the most reliable predictor of tick intensity and site selection was the season. In Chapter 2, I explored how tick attachment affects male S. occidentalis contest behavior. I. pacificus infestation has been shown to have negative physiological impacts on S. occidentalis, but mechanisms linking physiological changes to ultimate fitness consequences have been largely underexplored. I hypothesized that tick infestation reduces male S. occidentalis fighting ability by reducing O2 carrying ­­capacity­, or by obstructing or damaging vulnerable structures on their hosts. I held fifty half-hour trials between pairs of size- and ventral badge-matched male S. occidentalis, with one male in each pair infested with lab-reared I. pacificus larvae. I found that tick infestation negatively correlated with aggressive behavior in these staged contests. In support of reduced O2 capacity as the mechanism of reduced aggression, my ecologically relevant infestation intensities seemed to cause significant declines in hematocrit among experimentally infested lizards relative to controls. However, the site at which ticks attached did not significantly correlate with the aggressiveness of their lizard hosts. This is one of only a handful of studies to address the direct effect of I. pacificus on S. occidentalis. My study demonstrates that tick infestation can be detrimental to the fitness of their lizard hosts even without the transmission of pathogens. This insight may prove informative in future work on the ecology of I. pacificus-borne diseases in the western United States. This study is also one of only a few to use parasite infection to induce an asymmetry in fighting ability in intrasexual contests.

Salinization has been identified as an increasing threat to freshwater mussel diversity in recent years. Native mussels have been observed to display reduced salinity tolerance in comparison to some invasive bivalve species, but methods by which organismal tolerance is achieved are not well understood. This study was designed to compare the behavioral and physiological responses of the native Lampsilis ovata to that of the invasive Corbicula fluminea. Lampsilis were found to exhibit strong behavioral avoidance to salinity exposure, whereas Corbicula displayed very weak avoidance to comparable salinity concentrations followed by indications of osmotic conformation through physiological mechanisms. Prolonged valve closure in Lampsilis could translate to adverse consequences related to feeding, waste removal, and energetics. Alternatively, while physiological osmotic conformation in Corbicula is associated with increased energetic costs, it allows continued respiration and feeding. These differences could convey a competitive advantage with the increasing prevalence and severity of freshwater salinization events.

Aspectos do repertório comportamental e de estruturas resultantes de comportamento foram levantados para aranhas da infraordem Mygalomorphae. Os principais objetivos são buscar subsídios em informações provindas de caracteres comportamentais para auxiliar a elucidação da história evolutiva do grupo e entender como estes evoluíram dentro do grupo. Foi realizado um levantamento de informações sobre refúgios nas migalomorfas, a partir das quais foram delimitados 8 caracteres. Os caracteres foram incorporados a uma matriz de dados morfológicos provinda do trabalho de Goloboff e foi realizada uma análise de evidência total. A árvore resultante do consenso estrito da análise de evidência total mostrou-se melhor resolvida do que a árvore obtida apenas com os dados morfológicos. Foi realizada a otimização para a verificação da evolução dos padrões de teias dentro do grupo. Nossa análise mostra a existência de um padrão geral na evolução destes caracteres, com muitos deles apresentando fortes correlações. A construção de buracos está presente no ancestral comum entre migalomorfas e Mesothelae e aparece correlacionada com refúgios de alta conexão com o substrato. A hipótese de Coyle de que as tiras de seda teriam originado os lençóis de captura não é corroborada por nosso estudo. Ao contrário do que sugerem alguns autores, as estruturas finais resultantes de comportamentos podem ser uma fonte confiável de caracteres para o estudo da filogenia dos indivíduos que as produzem, além de possibilitar um melhor entendimento da evolução dos comportamentos que as originam. Também foi realizada uma análise comparativa do comportamento de autolimpeza em dois gêneros da família Theraphosidae. Observações deste comportamento resultaram na elaboração de um catálogo comportamental composto por 11 categorias. Análise das sequências comportamentais demonstra que este é um comportamento bastante fixo, com realização repetitiva de algumas sequências por longos períodos de tempo, não existindo um padrão geral que permita detectar início, meio e fim de uma sequência. São sugeridas algumas funções para este comportamento nas caranguejeiras, como a limpeza de estruturas sensoriais e/ou a proteção contra parasitas. A autolimpeza observada em indivíduos de outras famílias de Mygalomorphae mostrou a realização das mesmas categorias de modo semelhante. A manutenção destas categorias em famílias que divergiram há milhões de anos pode indicar um alto valor adaptativo das mesmas. A partir destes resultados, sugerimos que o tanto o comportamento quanto as estruturas finais são boas fontes de dados para análises filogenéticas. / Aspects of behavior and end-products of behavior were studied on Mygalomorph spiders. The main goal is to understand the evolution of these behavioral characters and to discover the evolutionary history of the group. As a result of an extensive review of the literature, 8 web characters were delimited in mygalomorphs. The characters were incorporated to a morphological matrix used by Goloboff and a total evidence analysis was performed. The resulting tree was better resolved than the strictly morphological one. Web related characters show a clear evolutionary signal, and some of them evolve in a correlated fashion. Burrow construction is an ancestral feature of spiders and appears correlated with silk-lining behavior. Coyles hypothesis that sheetwebs derive from silk lines is not supported by our analysis. Contrarily to some authorss suggestions, in our study the end-products proved to be a reliable source of characters for phylogenetic reconstructions, besides making possible a better understanding of the evolution of the behaviors that give rise to them. We performed a comparative analysis of grooming behavior in two genera of the family Theraphosidae. A behavioral catalog including 11 categories was concluded. Analysis of the behavioral sequences shows that grooming behavior is stereotyped, with some sequences being repeated for long periods of time, without a general pattern that let us detect the beginning, the middle or the end of a sequence. Spiders from distant Mygalomorph families perform the same repertoire of grooming behaviors, performed in similar ways. The maintenance of this ancestral repertoire in families that are separated for millions of years points to its strong adaptive value. Grooming could be used to clean some sensorial structures and/or to protect the spiders against parasites. From these results, we suggest that not only behavior, but also the structures resulting from it, are a good source of data for phylogenetic analysis.

Aspectos do repertório comportamental e de estruturas resultantes de comportamento foram levantados para aranhas da infraordem Mygalomorphae. Os principais objetivos são buscar subsídios em informações provindas de caracteres comportamentais para auxiliar a elucidação da história evolutiva do grupo e entender como estes evoluíram dentro do grupo. Foi realizado um levantamento de informações sobre refúgios nas migalomorfas, a partir das quais foram delimitados 8 caracteres. Os caracteres foram incorporados a uma matriz de dados morfológicos provinda do trabalho de Goloboff e foi realizada uma análise de evidência total. A árvore resultante do consenso estrito da análise de evidência total mostrou-se melhor resolvida do que a árvore obtida apenas com os dados morfológicos. Foi realizada a otimização para a verificação da evolução dos padrões de teias dentro do grupo. Nossa análise mostra a existência de um padrão geral na evolução destes caracteres, com muitos deles apresentando fortes correlações. A construção de buracos está presente no ancestral comum entre migalomorfas e Mesothelae e aparece correlacionada com refúgios de alta conexão com o substrato. A hipótese de Coyle de que as tiras de seda teriam originado os lençóis de captura não é corroborada por nosso estudo. Ao contrário do que sugerem alguns autores, as estruturas finais resultantes de comportamentos podem ser uma fonte confiável de caracteres para o estudo da filogenia dos indivíduos que as produzem, além de possibilitar um melhor entendimento da evolução dos comportamentos que as originam. Também foi realizada uma análise comparativa do comportamento de autolimpeza em dois gêneros da família Theraphosidae. Observações deste comportamento resultaram na elaboração de um catálogo comportamental composto por 11 categorias. Análise das sequências comportamentais demonstra que este é um comportamento bastante fixo, com realização repetitiva de algumas sequências por longos períodos de tempo, não existindo um padrão geral que permita detectar início, meio e fim de uma sequência. São sugeridas algumas funções para este comportamento nas caranguejeiras, como a limpeza de estruturas sensoriais e/ou a proteção contra parasitas. A autolimpeza observada em indivíduos de outras famílias de Mygalomorphae mostrou a realização das mesmas categorias de modo semelhante. A manutenção destas categorias em famílias que divergiram há milhões de anos pode indicar um alto valor adaptativo das mesmas. A partir destes resultados, sugerimos que o tanto o comportamento quanto as estruturas finais são boas fontes de dados para análises filogenéticas. / Aspects of behavior and end-products of behavior were studied on Mygalomorph spiders. The main goal is to understand the evolution of these behavioral characters and to discover the evolutionary history of the group. As a result of an extensive review of the literature, 8 web characters were delimited in mygalomorphs. The characters were incorporated to a morphological matrix used by Goloboff and a total evidence analysis was performed. The resulting tree was better resolved than the strictly morphological one. Web related characters show a clear evolutionary signal, and some of them evolve in a correlated fashion. Burrow construction is an ancestral feature of spiders and appears correlated with silk-lining behavior. Coyles hypothesis that sheetwebs derive from silk lines is not supported by our analysis. Contrarily to some authorss suggestions, in our study the end-products proved to be a reliable source of characters for phylogenetic reconstructions, besides making possible a better understanding of the evolution of the behaviors that give rise to them. We performed a comparative analysis of grooming behavior in two genera of the family Theraphosidae. A behavioral catalog including 11 categories was concluded. Analysis of the behavioral sequences shows that grooming behavior is stereotyped, with some sequences being repeated for long periods of time, without a general pattern that let us detect the beginning, the middle or the end of a sequence. Spiders from distant Mygalomorph families perform the same repertoire of grooming behaviors, performed in similar ways. The maintenance of this ancestral repertoire in families that are separated for millions of years points to its strong adaptive value. Grooming could be used to clean some sensorial structures and/or to protect the spiders against parasites. From these results, we suggest that not only behavior, but also the structures resulting from it, are a good source of data for phylogenetic analysis.

<p>Plants often grow in communities closely surrounded by neighbouring plants. Plants can actively and intensely compete for resources and also anticipate competition by sensing environmental cues from the presence and identity of neighbours. Moreover, it’s been proposed that the evolution of both increased and decreased competitive ability may serve as a mechanism for invasiveness. However, still little is known about how plants integrate competitive responses when sensing multiples cues of competition and which individual competitive traits respond to the identity of competitors. In addition, whether and why the evolution of competitive traits may contribute to the ability of introduced species to become invasive is also poorly understood.</p> <p>Here I present a body of work that examined the competitive responses of a native and an invasive plant species to cues of competition and the identity of neighbours. I also examined how experimental manipulation of pot volume, to control belowground resources, affects plant growth and allocation. In one study I tested the competitive responses of the North American native, <em>Impatiens pallida</em>, to cues signalling the presence of neighbours above and belowground simultaneously in competitive environments composed of either siblings or strangers. I demonstrate that<em> I. pallida</em> can recognize siblings and shows more aggressive competitive behaviours towards strangers than kin.</p> <p>In two other studies, I compared the competitive responses of the invasive and native ecotypes of <em>Alliaria petiolata</em> to changes in density, as well as to the presence and identity of neighbours. I found that invasive ecotypes produced less competitive phenotypes especially under high density. Moreover, I found that invasive ecotypes performed better when sharing rooting space with neighbours that were siblings.</p> <p>Taken together, these results demonstrate the ability of these plant species to respond to the identity of neighbours and provide strong evidence in support of the evolution of reduced competitive ability hypothesis in invasive plant species potentially mediated by the action of kin selection in invasive ecotypes.</p> / Doctor of Philosophy (PhD)

Badges of status are supposed to have insignificant production costs, so use costs are thought to be most important in ensuring signal reliability. Use costs arise from the use of the status signal in social interactions. Social experiences that arise from the use of inappropriate signals in social interactions may drive mechanisms that result in reduced fitness for inappropriate signalers. The role of social control, probing and familiarity in producing use costs was explored. There was no evidence that social control by dominants produced a cost for cheaters and no evidence that social control by subordinates produced a cost for inappropriate signaling by Trojans. Probing produced a cost for cheating when resource value was high but not when resource value was low. Familiarity had some effect on the cost and benefit of cheating but in patterns that were not predicted. Familiarity both eliminated a benefit of cheating and reduced a cost of cheating; therefore it is uncertain how familiarity affects honest signaling. The status signal of the receiver had no effect on the cost or benefits of cheating, and there was no evidence of punishment. Social experiences have the potential to affect signal development to produce a correspondence between signal and status. The effects of social experience on signal production were examined and there was little evidence that social experience influenced bib development. Neither aggression expressed nor aggression received was not predictive of bib size. Additionally, tests on the different measures of winning experience produced conflicting conclusions regarding the relationship between winning experience and bib development.