La présent étude examine aspect de l'écologie chimique de la Pyrale du maïs (Ostrinia nubilalis, Hübner, Lepidoptera: Crambidae). L’isolement reproducteur des phérotypes et leur spécialisation à la plante hôte ont reçu une attention particulière. Toutes les expériences ont été conçues pour avoir des informations pertinents sur le plan écologique des signaux chimiques libérés ou perçu par la Pyrale du maïs pendant la période de reproduction. La plupart des travaux en cadre de cette thèse dans des conditions naturelles et en tenant compte de tous les traits comportementaux de la Pyrale. Premièrement, le paysage chimique du champ de maïs a été étudié pour connaître les signaux chimiques rencontrés par une femelle de Pyrale du maïs qui est à la recherche d’une plante hôte. Les mélanges des volatiles qui composent l'environnement olfactif du champ de maïs a été identifié et leurs rôles discutés selon la biologie et le comportement d’oviposition de la Pyrale du maïs.Deuxièmement, en tenant compte de la périodicité du comportement de la Pyrale du maïs nous avons étudié les modifications circadiennes des profils des volatiles libérés par les plantes individuelles du maïs et par le champ du maïs.Troisièmement, le fidélité d’hôte plantes de la Pyrale du maïs a été étudié. Les volatiles libérés par différentes plantes hôtes au moment de vol d’oviposition ont été identifiés et le comportement de ponte a été étudié dans le tunnel du vol.Quatrièmement, les signaux chimiques impliqués dans l'isolement reproducteur des phérotypes de la Pyrale du maïs a été étudié. Nous nous sommes concentrés sur la phéromone mâle produite par les pinceaux androconiaux et sur leur rôle dans le comportement de cour. / The present PhD study investigates the aspect of chemical ecology in the European corn borer (ECB, Ostrinia nubilalis Hübner, Lepidoptera: Crambidae): pherotypes reproductive isolation and host plant specialisation.All the experiences were designed to have ecologically relevant information about the chemical signals released or perceived by ECB moth during the reproduction period. Most of the work was conducted close to natural conditions and taking into account all the behavioural traits of ECB.First we investigated what can be the chemical landscape of maize field for host-seeking ECB moth. The whole volatile blends that make up the olfactory environment of maize field biotope was identified according to the ECB oviposition behaviour (Paper I). Secondly taking into account the diel periodicity of ECB behaviour we studied the diel changes of volatile profiles released from maize plant and from maize field (Paper II). Thirdly, we studied the process of host fidelity within the ECB host plants. The volatiles released from different host plants at the time of ECB oviposition flight were identified and the oviposition behaviour was studied in wind tunnel to evidence the host plant preferences of ovipositing females (Paper III). Fourthly, we address the question on assortative mating in ECB pherotypes and we investigated the chemical signals involved in the reproductive isolation in ECB pherotypes. We focused on the male pheromone produced by hairpencils and on courtship behaviour (Paper IV).

Genetic variation is essential for biological evolution, for maintaining viability of populations, and to ensure ecosystem resilience. Increased human exploitation and environmental change result in rapid loss of biological variation, including genetic diversity. Measures to halt this trend require that biological diversity is assessed and monitored. Assessment of biodiversity includes identifying patterns of distribution of genetic variation within individual species. This thesis focuses on spatial genetic structure and assessment of units for conservation in continuous environments without apparent migration barriers. Empirical data refer to Scandinavia and the model species are northern pike (Esox lucius), brown trout (Salmo trutta), and harbour porpoise (Phocoena phocoena). Questions regarding monitoring genetic diversity and releases of alien populations are also addressed.  The spatial genetic structure of the northern pike in the Baltic Sea is characterized by isolation by distance and continuous genetic change. Positive genetic correlation was found among pike within geographical distances of less than 150 km. This distance may be used to suggest management units in this area. For the brown trout, genetic monitoring identified two sympatric populations within a small mountain lake system. The situation is characterized by a clear genetic but no apparent phenotypic dichotomy. Scientific support for a genetically distinct Baltic harbour porpoise population is limited, and the spatial genetic structure of the harbour porpoise in Swedish waters needs to be clarified. Data for launching conservation genetic monitoring programs is available for only a few Swedish species. Millions of forest trees, fish, and birds are released annually in Sweden and the documentation on these releases is poor. To meet responsibilities of safeguarding biodiversity and surveying biological effects of releases, there is an urgent need for studies aimed at evaluating genetic diversity. / At the time of the doctoral defense, the following paper was unpublished and had a status as follows: Paper 2: Manuscript.

The present PhD study investigates the aspect of chemical ecology in the European corn borer (ECB, Ostrinia nubilalis Hübner, Lepidoptera: Crambidae): pherotypes reproductive isolation and host plant specialisation.All the experiences were designed to have ecologically relevant information about the chemical signals released or perceived by ECB moth during the reproduction period. Most of the work was conducted close to natural conditions and taking into account all the behavioural traits of ECB.First we investigated what can be the chemical landscape of maize field for host-seeking ECB moth. The whole volatile blends that make up the olfactory environment of maize field biotope was identified according to the ECB oviposition behaviour (Paper I). Secondly taking into account the diel periodicity of ECB behaviour we studied the diel changes of volatile profiles released from maize plant and from maize field (Paper II). Thirdly, we studied the process of host fidelity within the ECB host plants. The volatiles released from different host plants at the time of ECB oviposition flight were identified and the oviposition behaviour was studied in wind tunnel to evidence the host plant preferences of ovipositing females (Paper III). Fourthly, we address the question on assortative mating in ECB pherotypes and we investigated the chemical signals involved in the reproductive isolation in ECB pherotypes. We focused on the male pheromone produced by hairpencils and on courtship behaviour (Paper IV).

Submitted by Renata Lopes (renatasil82@gmail.com) on 2017-10-09T20:17:53Z No. of bitstreams: 1 ludemilamartinsdesouza.pdf: 1476036 bytes, checksum: 13c93142a0ac36c1cce3f43d89f846ca (MD5) / Approved for entry into archive by Adriana Oliveira (adriana.oliveira@ufjf.edu.br) on 2017-10-10T12:20:05Z (GMT) No. of bitstreams: 1 ludemilamartinsdesouza.pdf: 1476036 bytes, checksum: 13c93142a0ac36c1cce3f43d89f846ca (MD5) / Made available in DSpace on 2017-10-10T12:20:05Z (GMT). No. of bitstreams: 1 ludemilamartinsdesouza.pdf: 1476036 bytes, checksum: 13c93142a0ac36c1cce3f43d89f846ca (MD5) Previous issue date: 2017-04-27 / Dados sobre a ecologia alimentar de duas aves de sub-bosque foram coletados na Reserva Biológica Municipal do Poço D’anta, Juiz de Fora – MG, sudeste do Brasil. Conopophaga lineata e Conopophaga melanops são encontradas ocorrendo em simpátria em algumas localidades da Região Sudeste. Estudos com espécies aparentadas e simpátricas podem ser úteis no sentido de avaliar a influência do nicho sobre co ocorrência e co-existência. Durante as amostragens utilizando redes ornitológicas obtivemos 29 capturas de 25 indivíduos de C. lineata, e 28 capturas de 21 indivíduos de C. melanops. Na triagem das amostras fecais de C. lineata e C. melanops foram encontrados, respectivamente, 310 e 276 itens. C. lineata consumiu predominantemente Formicidae (53,5%), Coleoptera (6,8%). Entretanto, o índice alimentar (IA) de Coleoptera (7,47) ficou abaixo de outros grupos como Formicidae (68,2) e Araneae (8,80). C. melanops consumiu altas proporções de Formicidae (23,2%), Coloptera (12,3%), Orthoptera (11,6%) e Isoptera (11,2%), embora em termos de IA o predomínio tenha sido de Formicidae (24,66) e Coleoptera (20,14). A sobreposição de nicho entre as espécies foi maior que o esperado (observado = 0.723). Apesar das diferenças nas proporções dos tipos de artrópodes consumidos não foram suficientes para revelar separação de seus nichos alimentares. Embora se encontrem na literatura que C. lineata seleciona tanto tipo de presas quanto artrópodes maiores que 5 mm em proporções acima do esperado pelo acaso, assim como foi observado para C. melanops predação preferencial sobre alguns táxons. Os resultados encontrados demonstram que, em alguns casos, os padrões ecológicos das respostas das aves aos recursos encontrados no ambiente podem ser diferentes daqueles encontrados em outras regiões. / Data on the food ecology of two understory birds were collected at the Municipal Reserva Biológica Municipal do Poço D’anta, Juiz de Fora – MG, Southeastern Brazil. Conopophaga lineata and Conopophaga melanops are found occurring in sympatry in some localities of the Southeast Region. Studies with related and sympatric species may be useful in assessing the influence of the niche on co-occurrence and co-existence. During the samplings using ontological networks were carried out 29 catches of 25 individuals of C. lineata, and 28 catches of 21 individuals of C. melanops. In the sorting of the fecal samples of C. lineata and C. melanops were found, respectively, 310 and 276 items. C. lineata consumed predominantly Formicidae (53.5%), Coleoptera (6.8%). However, Coleoptera dietary index (AI) (7.47) was below other groups such as Formicidae (68.2) and Araneae (8.80). C. melanops consumed high proportions of Formicidae (23.2%), Coloptera (12.3%), Orthoptera (11.6%) and Isoptera (11.2%), although in terms of AI the predominance was Formicidae (24.66) and Coleoptera (20.14). The niche overlap between species was higher than expected (observed = 0.723). Although differences in the proportions of the types of arthropods consumed were not sufficient to reveal separation of their food niches. Although it is found in the literature that C. lineata selects both prey and arthropod species larger than 5 mm in proportions above that expected by chance, just as it was observed for C. melanops preferential predation on some taxa. The results show that, in some cases, the ecological patterns of bird responses to the resources found in the environment may be different from those found in other regions.

Comparative studies on endangered species and their more common congeners can shed light on the mechanism of species rarity, and enable conservationists to formulate effective management strategies. I compared the breeding and pollination systems of the endangered Geodorum eulophioides and its two more common sympatric congeners in subtropical China. Geodorum eulophioides and G. recurvum were self-compatible, both depending on Ceratina cognata for fruit production, while G. densiflorum can autonomously self. Although the floral visitation frequency of G. eulophioides was the highest among the three, its natural fruit set was similar to that of G. recurvum, but both lower than that of G. densiflorum. These results partially explain the difference in species abundance. Coupled with severe habitat loss and degradation, the extremely low pollinator visitation and natural fruit set of G. eulophioides calls for rapid establishment of ex-situ collection, in conjunction with improving in-situ habitat. Natural hybridization tendencies were also studied among species.

Interaction of genes and culture is crucial for human evolution. Human ethnic groups and subcultures frequently function as discrete units, and people clearly distinguish between in- group and out-group individuals on a cultural basis. This thesis aims to model the formation of distinct cultural clusters, cultural equivalents of distinct species. Historical development of theories of blending inheritance led to the formation of biometric parallels to Mendelism. Galton-Pearson model of nonparticulate inheritance with constant offspring variance, the most influential model of continuous inheritance ever formulated, was based on measurements of genetically transmitted traits. Ronald Fisher later demonstrated, that this type of inheritance directly stems from polygenic traits with additive genetic variance. Dan Sperber's metaphor of culture space allows integrating any continuous models of position inheritance into computer simulations of the evolution of culture. Most studies today, however, employ particulate models of cultural inheritance. The exceptional works of Cavalli-Sforza and Feldman pioneer the continuous models of cultural inheritance applying Galton-Pearson model to culture. Galton-Pearson inheritance is, unfortunately, not a very good model of cultural transmission. Parental...

Natural populations exhibit a non-random mating behavior and it is assumed that mate preferences causing the non-random mating behavior play a role in sympatric speciation. In my thesis, I have proposed a new model of a mate preference that is based on ecological performance and I have named it fitness-based mating. Individuals that express this mate preference choose primarily fit partners. Fitness-based mating is modelled for haploid, diallelic populations. Individuals are distributed across two niches, and genomes are simplified to two loci. The first locus is subject to natural selection, and the second-locus genotype gives the strength of the mate preference. The population is separated into females and males, among which only females exhibit the mate preference. Ecological selection acts on both sexes alike. With the model I have investigated how female choosiness based on direct advantages offered by their partners can cause and maintain a polymorphic population. Fitness-based mating is an evolutionary successful mating strategy. It spreads in a population due to its amplifying effect on the reproductive success and on the attractiveness of its carriers. A polymorphism arises naturally in the model. The emergence of a stable polymorphism of traits underlying ecological selection is of special interest, because a polymorphism can be a precursor of speciation.

La plasticité comportementale et les interactions entre organismes sont connues pour avoir une influence sur l’utilisation de l’espace, notamment la sélection de l’habitat de chasse. Dans l’Atlantique Nord-Est, les phoques gris et veaux marins vivent en sympatrie. Ces deux espèces sont réparties dans différentes colonies où les interactions entre organismes (liées à la taille de la colonie et à la présence simultanée ou non des deux espèces) ainsi que la disponibilité de l’habitat (et donc des proies) varient localement. L’objectif de cette thèse était d’étudier l’utilisation spatiale et la sélection d’habitat de chasse à l’échelle locale de ces deux espèces dans des situations contrastées de dynamique de colonies. Pour cela, plusieurs individus de différentes colonies situées en Irlande, en Écosse et en France ont été équipés de balises GPS/GSM. Les données récoltées ont permis de caractériser leurs trajets en mer ainsi que d’identifier les zones de chasse permettant par la suite la sélection d’habitat. Contrairement aux veaux marins, qui ont réalisé des déplacements restreints aux alentours des colonies, la plupart des phoques gris se sont déplacés entre différentes colonies. Ainsi, l’étude de la sélection d’habitat à l’échelle individuelle pour les individus se déplaçant entre différentes colonies a mis en évidence l’influence de la plasticité comportementale et suggéré celle de la personnalité de l’animal. A l’échelle de la colonie, chez les deux espèces, les résultats ont mis en évidence l’influence de la disponibilité de l’habitat (et indirectement de la disponibilité des proies) ainsi que des interactions intra-spécifiques (liée à l’effet de densité dépendance) sur l’utilisation spatiale et la sélection de l’habitat de chasse. Cependant, la sélection de l’habitat de chasse était principalement liée à la distance au reposoir et la bathymétrie. Ces deux paramètres sont liés aux caractères de chasseur à place centrale et benthique des phoques. Cette étude a également mis en évidence, pour les sites où les deux espèces étaient présentes, une certaine ségrégation spatiale et des différences de sélection d’habitat entre les deux espèces. / The behavioural plasticity and the interactions between organisms are known to influence foraging habitat selection and spatial usage. In the Northeast Atlantic, greyand harbour seals live in sympatry. These two species are spread into different colonies where interactions between organisms (linked to the colony size et and the presence of one or both species) and habitat availability (linked to the prey distribution) vary locally. The objective of this PhD was to study the spatial usage and foraging habitat selection of these two species at the local scale, in contrasted situations of population dynamics. Several individuals were tagged with GPS/GSM tags in different colonies located in Ireland, Scotland and France. Telemetry data was used to characterize the seals’ trips at sea and to identify their foraging areas in order to model foraging habitat selection. While harbour seals only performed trips restricted around their haulout sites, grey seals moved between colonies. Studying habitat selection at the individual scale, for individuals moving between colonies, highlighted the influence of behavioural plasticity and suggested the influence of individual personality. At the colony scale, for both species, results highlighted the influence of habitat availability (indirectly linked to prey availability) and intra-specific interactions (linked to density dependence effects) on spatial usage and foraging habitat selection. However, the foraging habitat selection was mainly explained by the distance from the last haulout and the bathymetry. These two parameters are linked to the central place forager and benthic feeder characteristics of the seals. Furthermore, this study also highlighted, for study sites where both species occur, a spatial segregation and differences in foraging habitat selection between grey and harbourseals.

Submitted by Marlene Santos (marlene.bc.ufg@gmail.com) on 2014-09-17T21:01:01Z No. of bitstreams: 2 José Neiva Mesquita Neto.pdf: 2799737 bytes, checksum: 590e98d41066ac6babc47eeb04637796 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) / Rejected by Luciana Ferreira (lucgeral@gmail.com), reason: Marlene, olhe nas orientações a questão da citação, a qual deve ser de acordo com a NBR 6023: ALCÂNTARA, Guizelle Aparecida de. Caracterização farmacognostica e atividade antimicrobiana da folha e casca do caule da myrciarostratadc.(myrtaceae). 2012. 41 f. Dissertação (Mestrado em Ciências Farmacêuticas) - Universidade Federal de Goiás, Goiânia, 2012. on 2014-09-18T12:33:01Z (GMT) / Submitted by Marlene Santos (marlene.bc.ufg@gmail.com) on 2014-09-18T19:01:02Z No. of bitstreams: 2 license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) José Neiva Mesquita Neto.pdf: 2799737 bytes, checksum: 590e98d41066ac6babc47eeb04637796 (MD5) / Approved for entry into archive by Luciana Ferreira (lucgeral@gmail.com) on 2014-09-19T11:32:20Z (GMT) No. of bitstreams: 2 license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) José Neiva Mesquita Neto.pdf: 2799737 bytes, checksum: 590e98d41066ac6babc47eeb04637796 (MD5) / Made available in DSpace on 2014-09-19T11:32:20Z (GMT). No. of bitstreams: 2 license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) José Neiva Mesquita Neto.pdf: 2799737 bytes, checksum: 590e98d41066ac6babc47eeb04637796 (MD5) Previous issue date: 2013-09-19 / Conselho Nacional de Pesquisa e Desenvolvimento Científico e Tecnológico - CNPq / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / Plant-pollinator interactions in syncronopatric species of Psychotria (Rubiaceae). Psychotria species are commonly found in the understory of forested areas in tropical regions, and are important components for the functioning of these ecosystems. Many species of the genus are sympatric and bloom during the same period, being considered potentially syncronopatrics. For this reason, this genus has been considered as a good model to infer general patterns and mechanisms of speciation in the tropics. In the first paper of this dissertation, the interactions between potentially sympatric species of Psychotria and their pollinators were analyzed in order to verify the possible existence of sharing, specialization or generalization of pollinators and plants in the analyzed system. Then, data of secondary studies that contained identifying pollinators of Psychotria were analyzed From these data, we generated graphs and analyzes of interaction networks and niche overlap. Altogether, nine species of Psychotria and 25 species of pollinators of occurrence in the Atlantic Forest were included in the analyzes. The plant with the most generalist network is Psychotria tenuinervis and the pollinators with the highest proportion of links and consequently with more important role in the network belonged orders Lepidoptera and Hymenoptera. Psychotria species showed niche overlap in pollination and pollinator sharing. The period of greatest intensity of flowering coincided with the rainy season in the Atlantic Forest, with up to seven species coflowering. Because Psychotria species occur in sympatry and have flowering overlap and asymmetric interactions with flower visitors, it is possible that they are involved in a process of facilitation in pollination. The second paper aimed to investigate if four sympatric populations of Psychotria show pollinators sharing and pollination niche overlap . Another objective was to determine if their flowering season are synchronics. We found a dual synchrony in this system, the first in the intensity of flowering among the populations of Psychotria and the second in the increase of the abundance of flowers and pollinators. The species of Psychotria showed sharing fundamental niche of pollination and positive impact on reproductive success. However, we note that the sharing of pollinators can not be generalized to the entire system, but to the modules or pairs of species. Thus, each species of Psychotria has one or more species of preferential pollinator for sharing and that this relationship is not always reciprocal. This reinforces that the interactions among the plant species are asymmetric. / Interação planta-polinizador em espécies sincronopátricas de Psychotria (Rubiaceae). Psychotria contribui de forma importante para a diversidade florística tropical e são comumente encontradas em sub-bosque de áreas florestadas em regiões tropicais, sendo importantes componentes para o funcionamento destes ecossistemas. Muitas espécies do gênero são simpátricas e florescem em um mesmo período do ano, sendo consideradas potencialmente sincronopátricas. Por esta razão, este gênero tem sido considerado como um bom modelo para inferir padrões e mecanismos gerais de especiação nos trópicos. No primeiro artigo, as interações entre espécies potencialmente simpátricas de Psychotria e seus polinizadores foram analisadas com o objetivo de verificar a possível existência de partilha, de especialização ou generalização de polinizadores e plantas no sistema analisado. Para isso, foram incluídas informações secundárias de estudos que continham a identificação em nível de espécie de polinizadores de Psychotria. A partir desses dados, foram gerados grafos e realizadas análises de redes de interação e sobreposição de nicho. Ao todo, foram incluídas nas análises nove espécies de Psychotria com registro de ocorrência para a Mata Atlântica, sendo que elas interagiram com 25 espécies de polinizadores. A planta mais generalista da rede foi Psychotria tenuinervis e os polinizadores que concentraram maior número de links e consequentemente com papel mais importante na rede pertenceram as ordens Lepidoptera e Hymenoptera. As espécies de Psychotria apresentaram sobreposição de nicho na polinização com partilha de polinizadores. O período de maior intensidade de floração coincidiu com o período chuvoso na Mata Atlântica, com até sete espécies apresentando coflorescimento. Assim, pode-se inferir que a polinização em Psychotria demanda polinizadores generalistas. Pelo fato das espécies de Psychotria ocorrerem em simpatria, possuírem floração sobreposta e interações assimétricas com visitantes florais, é possível que estejam envolvidas em um processo de facilitação na polinização. Já o segundo artigo objetivou averiguar se quatro populações simpátricas de Psychotria, em cofloração, apresentaram partilha de polinizadores e sobreposição de nicho na polinização. Além disso, objetivou verificar se existe sincronia entre as populações vegetais e destas com comunidade de polinizadores. Foi encontrada uma dupla sincronia nesse sistema, sendo a primeira na intensidade de floração entres as populações de Psychotria e a segunda no aumento da abundância de flores e de polinizadores. As espécies de Psychotria apresentaram partilha fundamental do nicho da polinização e com reflexo positivo no sucesso reprodutivo. No entanto, verificamos que a partilha de polinizadores não pode ser generalizada a todo o sistema, mas sim a módulos ou pares de espécies. Dessa forma, cada espécie de Psychotria possui uma ou mais espécies preferencias para partilha de polinizadores e que nem sempre essa relação é reciproca. Isso reforça que as interações entre as plantas são assimétricas.

When species compete for resources, in a stable homogeneous environment, there are two possible outcomes. The first is that one species will out-compete the other and exclude it from the environment. This is known as the competitive exclusion principle. The second is that both species will manage to coexist. Coexistence can only occur if the species’ niches are differentiated such that interspecific competition is minimised, or eliminated. This outcome is known as resource partitioning. Two closely related Australian skink species of the Egernia genus, Egernia multiscutata and Egernia whitii, are abundant and sympatric on Wedge Island in South Australia’s Spencer Gulf. The species are morphologically very similar and appear to have very similar life histories and habitat requirements. Ostensibly, they would compete for limiting resources in this environment. This thesis is the first investigation into resource partitioning in this previously unstudied model organism. I report the results of multi-faceted investigations into the coexistence of the skinks, E. multiscutata and E. whitii on Wedge Island and the evidence for, and mechanisms of, any facultative resource partitioning between them. Study methods involved a transect survey of most of Wedge Island to determine the species’ distributions and any evidence for resource partitioning; a morphological comparison to investigate any potential competitive advantages of either species; a habitat choice experiment to establish retreat-site preferences in the absence of interspecific interference; and, a series of staged dyadic encounter experiments to investigate interspecific competitive interactions. Resource partitioning was evidenced by differential distributions of the species among substrates containing the elements required for permanent refuge shelters. This partitioning was not mediated by avoidance of particular substrates but by the presence of the opponent species, combined with attraction to suitable substrates. Asymmetries in some morphological characters were found to confer a potential competitive advantage to E. multiscutata in agonistic encounters with E. whitii. Both species were found to have the same refuge site preferences when interference competition was experimentally removed. This result was not concordant with observed resource partitioning in the field and suggests that the habitat choices of both species are modified by the presence of the opponent species. Analyses of staged dyadic encounter experiments showed that E. multiscutata was more likely to gain greater access to a contested habitat resource and more likely to exclude E. whitii from the resource than vice-versa. Nevertheless, the outcome of competitive interactions was not completely deterministic and there was some tolerance of co-habitation. E. multiscutata’s competitive advantage was attributable largely to its greater mass and head dimensions relative to snout to vent length. However, differential behavioural responses to the threat of larger opponent size also played an important part in resource partitioning between the species.