Sociality is widespread across the animal kingdom and explanations for its incidence and persistence are numerous. Whilst various drivers of sociality have been identified and tested, controversies remain and we are still far from a complete understanding of the mechanisms underlying social structure. Here I use a combination of field observations on a free-living population of feral horses Equus caballus and laboratory behavioural experiments on the Pacific beetle roach Diploptera punctata to investigate the drivers of sociality in these species. I explore four key aspects of sociality: the influences of kinship on sociality and social development, the strength and persistence of social bonds, the relationship between inbreeding avoidance and dispersal and the potential influence of individuals on social structure. Whilst kinship is a major driver of social structure in most mammalian species, I present evidence in Chapter Three that horse society is not structured by levels of kinship; however, in Chapter Five, I show that kinship levels to potential mates are significant in female dispersal choices in this species. In Chapter Eight, I provide evidence for significant effects of kinship to companions upon social and physical development in D. punctata, indicating a clear potential benefit of kin-based associations. The stability of social bonds can have substantial effects upon social structure. In Chapter Three, I show that the bonds between female horses show significant stability and are formed independently to kinship levels, a rare result in a non-primate species. I also provide evidence consistent with the hypothesis that these bonds are driven by male harassment. Similarly, in D. punctata, I find in Chapter Eight that female clustering occurs within resting aggregations and that the most likely explanation is the avoidance of male harassment. I therefore propose that this driver of female sociality may be a highly prevalent force structuring animal societies. Inbreeding depression has been demonstrated in a variety of species and contexts. Here I show in Chapter Five that in horses, female dispersal is likely to be influenced by kinship levels with potential mates. In Chapter Four, I then show that more heterozygous males have a higher reproductive success, most likely due to their ability to utilise a larger home range. Finally, local population structure can be highly influenced by individual association choices and behaviour. In Chapter Two, I show that in horses, mothers may allow their sons to postpone dispersal by the maintenance of stronger mother-son bonds, permitting an extended period of social learning. In Chapter Seven, I demonstrate that consistent inter-individual variation in personality traits exists in D. punctata which is stable across life stages, despite age effects on the strength of boldness. This is a source of variation which may be extremely important for decision-making social groups. My main conclusion from this work is that male harassment is often a key driver of sociality which may frequently be overlooked. I also demonstrate that the effects of kinship are far-ranging but not omnipresent. This thesis therefore makes a major contribution to our understanding of the mechanisms underlying animal sociality and presents clear potential avenues for future research.

Submitted by Gabriela Lopes (gmachadolopesufpel@gmail.com) on 2017-07-11T18:17:37Z No. of bitstreams: 2 license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Dissertação Simone Andzeiewski.pdf: 1716530 bytes, checksum: 2a84b68dc4fb16f321fff1dc454467b1 (MD5) / Approved for entry into archive by Aline Batista (alinehb.ufpel@gmail.com) on 2017-07-17T20:38:01Z (GMT) No. of bitstreams: 2 Dissertação Simone Andzeiewski.pdf: 1716530 bytes, checksum: 2a84b68dc4fb16f321fff1dc454467b1 (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) / Made available in DSpace on 2017-07-17T20:38:01Z (GMT). No. of bitstreams: 2 Dissertação Simone Andzeiewski.pdf: 1716530 bytes, checksum: 2a84b68dc4fb16f321fff1dc454467b1 (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Previous issue date: 2017-02-24 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / A pérola-da-terra Eurhizococcus brasiliensis (Wille, 1922) (Hemiptera: Margarodidae) é uma cochonilha subterrânea que ataca raízes de plantas cultivadas e silvestres. A espécie é considerada a principal praga da videira no Brasil. No primeiro ínstar, a dispersão da cochonilha é realizada com o auxílio de formigas doceiras principalmente a Linepithema micans (Forel, 1908) (Hymenoptera: Formicidae) que se associa à cochonilha na busca de excrementos açucarados. Nesse trabalho, foi avaliado o efeito de iscas tóxicas a base de hidrametilnona no controle de L. micans, e consequentemente, na infestação de E. brasiliensis em novos plantios de videira. Inicialmente foi realizado um experimento em vasos em casa de vegetação em outubro de 2014 com a finalidade de selecionar formulações de iscas tóxicas à base de hidrametilnona (“grânulo pequeno”, “grânulo grande”, tipo “cereal” e gel) para o controle de L. micans. As iscas tóxicas foram oferecidas ad libitum em porta-iscas com substituição semanal. As avaliações foram realizadas semanalmente registrando o número de formigas forrageando a cada 10 minutos durante uma hora sobre uma solução aquosa de açúcar invertido 70%. A melhor formulação de isca tóxica selecionada em casa de vegetação foi avaliada a campo em dois experimentos, realizados em áreas naturalmente infestadas pela pérola-da-terra e por L. micans. No primeiro, foram efetuadas aplicações semanais de 450 g/ha da isca tóxica em Flores da Cunha, RS, e no segundo, foi avaliado uma e três aplicações semanais da mesma dose em Caxias do Sul, RS. Mudas do porta enxerto Paulsen 1103 foram plantadas em agosto de 2014 para o experimento em Flores da Cunha e em novembro de 2015 no experimento de Caxias do Sul, respectivamente. O monitoramento populacional das formigas foi feito semanalmente através de armadilhas subterrâneas do tipo “pitfall” utilizando mel e sardinha embebidos em algodão. A avaliação dos experimentos foi realizada em maio de 2015 (Flores da Cunha) e 2016 (Caxias do Sul), contando o número de cochonilhas presentes nas raízes. No experimento em casa de vegetação, a isca tóxica a base de hidrametilnona formulada em “grânulo pequeno” e “grânulo grande” controlaram a população de formigas nas primeiras quatro semanas após o fornecimento da isca. A isca tóxica formulada em gel também proporcionou mortalidade de 100% da população de formigas, no entanto seu controle foi mais lento, quando comparado as outras formulações. A isca “cereal” não proporcionou controle das formigas. A isca tóxica formulada em “grânulo pequeno” selecionada como a mais eficaz em casa de vegetação, fornecida semanalmente no campo, controlou a população de L. micans em 100% reduzindo a infestação da 8 pérola-da-terra em 99,9%. Quando o número de aplicações da isca foi reduzido para 1 a 3, houve um controle de 100% na população de formigas no tratamento que recebeu três aplicações até a quarta semana após o fornecimento da isca, porém as colônias se restabeleceram ao longo do experimento e os tratamentos (uma e três vezes) não foram eficazes no controle das espécies. Conclui-se que a formulação granulada à base de hidrametilnona “grânulo pequeno” é eficiente no controle de L micans em casa de vegetação e em aplicações semanais no campo. Aplicações de uma e três vezes da isca tóxica não proporcionaram controle satisfatório das espécies sendo necessário ajustar o número de tratamentos para um controle eficaz e econômico de E. brasiliensis e L. micans. / In Brazil, Eurhizococcus brasiliensis (Wille) (Hemiptera: Margarodidae) is considered to be the main pest of vine plants, attacking the roots of cultivated and wild plants. Linepithema micans (Forel, 1908) (Hymenoptera: Formicidae) is the primary agent responsible for spreading the Eurhizococcus brasiliensis, which are attracted by the honeydew excreted by the scale insect. In this work, the effect of a hydramethylnonbased granular toxic bait on the control of L. micans and E. brasiliensis in grapevines was evaluated on young grapevines. An experiment was conducted in a greenhouse in October 2014 and planted in individual pots were used to select different formulations of toxic baits based on hydramethylnon (small granule, large granule, cereal and gel) for the control of L. micans. The toxic baits were provided ad libitum in bait holders, with baits being replaced weekly. Weekly evaluations were carried out during a period of 10 weeks, with the number of foraging ants being counted every 10 minutes during 1 hour, in an aqueous solution of 70% inverted sugar. The best formulation of toxic bait selected in greenhouse was evaluated in the field in two experiments, carried out in areas naturally infested withe the scale and L. micans. In the first experiment 450 g/ha/week of the toxic bait were applied in Flores da Cunha, RS, and in the second, one and three weekly applications of the same dose were evaluated in Caxias do Sul, RS. In August 2014, seedlings from the Paulsen 1103 rootstock were planted in Flores da Cunha and in November 2015 in Caxias do Sul. The ants was monitored weekly in-ground pitfall traps using a honey and water solution (70%) and sardine conserved in oil absorbed in cotton wool as food attractants.The evaluation of the experiments was carried out in May of 2015 (Flores da Cunha) and 2016 (Caxias do Sul), counting the number of scale insects present in the roots.The experiment conducted in a greenhouse, the hydramethylnon based toxic bait in the formulation small granule and large granule controlled the ant population in the first 4 weeks of application.The gel formulation of the hydramethylnon toxic bait also provided 100% mortality of the population of ants; however, its control was slower when compared with the other two formulations and the cereal formulation did not differ from the control. The small granule toxic bait, applied weekly, effectively reduced the infestation of L. micans, controlling 100% of the population of ants and reducing the infestation of E. brasiliensis by 99.9%. When the number of bait applications was reduced to 1 to 3, there was a 100% reduction in the ants in the treatment that received three applications until the fourth week after application of the bait, but the colonies were re-establishment.The small granule toxic bait, applied weekly, effectively reduced the infestation of L. micans in greenhouse and field.The reduction of the number of applications (one and three times) did not provide satisfactory control and it will be 10 necessary to adjust the number of treatments for an efficient and economical control of the two species.

A biodiversidade dos rios brasileiros encontra-se sob severa ameaça, em razão da fragmentação e perda de hábitat que os barramentos representam. Até o presente, os aproveitamentos hidrelétricos têm seu licenciamento ambiental condicionado a análises de impacto locais, sem avaliação dos impactos cumulativos de múltiplas barragens em uma bacia hidrográfica, com relação à conservação da diversidade da ictiofauna. Estudos anteriores para subsídio de licenciamento de múltiplas barragens na bacia Taquari-Antas (RS) propuseram cenários alternativos de conservação (número e posição de barragens), quando ainda não havia métricas de conectividade dendrítica desenvolvidas. Entre os problemas identificados naqueles estudos figuram a quantificação dos efeitos cumulativos das barreiras sobre a conectividade da bacia, o estabelecimento de áreas prioritárias para conservação e a influência do número e da posição das barreiras na conservação da biodiversidade aquática em uma bacia. Avaliamos a aplicação de conectividade para mensuração de impactos cumulativos de fragmentação de bacias hidrográficas através de índices de conectividade dendrítica recentemente propostos. Propomos um método genérico e replicável para analisar quantitativamente os efeitos de sucessivos barramentos em relação à conectividade dos hábitats aquáticos em processos de migração e dispersão de peixes em bacias hidrográficas. Utilizamos simulações de cenários de múltiplos barramentos para a avaliação. Propomos também uma sistematização para a simulação de múltiplos barramentos. Além disso, hierarquizamos áreas para conservação por conectividade, aplicamos as novas métricas de conectividade a estudos anteriores e demonstramos casos de cenários de alta conectividade com outras implicações na conservação. Discutimos o amadurecimento do método para aplicação em licenciamento ambiental e planejamento de conservação, bem como limitações atuais e perspectivas para trabalhos futuros. / Biodiversity of Brazilian rivers is nowadays seriously threatened due to fragmentation and habitat loss that impoundments represent. Up to now, hydroelectric power plants have their environmental licensing processes conditioned to local impact analysis, with no evaluation of cumulative impacts of multiple dams in a watershed landscape on the conservation of the integrity of ichthyofauna. Former studies for supporting environmental licensing processes of multiple hydroelectric dams in the Taquaria-Antas basin (RS, Brasil) proposed alternative conservation scenarios (number and position of barriers), in a time when there were no dendritic connectivity metrics developed. Among the problems identified at that time, we point quantification of cumulative effects of multiple barriers on the drainage connectivity, detecting prioritary areas for conservation and detecting the influence of the number and position of the barriers in the catchment for best conservation of aquatic biodiversity. We evaluated the application of connectivity for quantifying the impacts of fragmentation in hydrographic basins through recently proposed indexes. We proposed a replicable and generic method for quantifying the effects of successive impoundments in relation to aquatic habitat connectivity in ecological processes of migration and dispersal of fishes in hydrographic basins. We used multiple barriers scenarios simulation in order to perform the assessments. We also propose a way of systematizing multiple barriers simulations. Moreover, we rank areas for conservation by connectivity, apply the new connectivity metrics on former studies and demonstrate cases of high connectivity scenarios with other implications on conservation. We discuss maturing the method for application on environmental licensing and conservation planning as well as current limitations and perspectives for future studies.

Tracking seed dispersal using traditional, direct measurement approaches is difficult and generally underestimates dispersal distances. Variation in chloroplast haplotypes (cpDNA) offers a way to trace past seed dispersal and to make inferences about factors contributing to present patterns of dispersal. Although cpDNA generally has low levels of intraspecific variation, this can be overcome by assaying the whole chloroplast genome. Whole-genome sequencing is more expensive, but resources can be conserved by pooling samples. Unfortunately, haplotype associations among SNPs are lost in pooled samples and treating SNP frequencies as independent estimates of variation provides biased estimates of genetic distance. I have developed an application, CallHap, that uses a least-squares algorithm to evaluate the fit between observed and predicted SNP frequencies from pooled samples based on network topology, thus enabling pooling for chloroplast sequencing for large-scale studies of chloroplast genomic variation. This method was tested using artificially-constructed test networks and pools, and pooled samples of Lasthenia californica (California goldfields) from Whetstone Prairie, in Southern Oregon, USA. In test networks, CallHap reliably recovered network topologies and haplotype frequencies. Overall, the CallHap pipeline allows for the efficient use of resources for estimation of genetic distance for studies using non-recombining, whole-genome haplotypes, such as intra-specific variation in chloroplast, mitochondrial, bacterial, or viral DNA.

Rapid range expansion of newly invasive species provides a unique opportunity for studying patterns of dispersal and gene flow. In this thesis, I examined the effect of landscape features on gene flow in the invasive grass Brachypodium sylvaticum at the edge of its expanding range. I used genome-wide Single Nucleotide Polymorphism (SNP) surveys of individuals from 22 locations in the Clackamas Watershed in the Portland, Oregon metropolitan region to assess genetic diversity and structure, to identify putative source populations, and to conduct landscape genetic analyses. Resistance surfaces were created for each landscape feature, using ResistanceGA to optimize resistance parameters. My STRUCTURE analysis identified three distinct clusters, and diversity analyses support the existence of at least two local introductions. Multiple Regression on distance Matrices (MRM) showed no evidence that development, roads, canopy cover, or agriculture had a significant influence on genetic distance in B. sylvaticum. The effect of geographic distance was marginal and reflected geographic clustering. The model of rivers acting as a conduit explained a large portion of variation in genetic distance. Results indicate that rivers influence patterns of dispersal of B. sylvaticum by human recreational activity centering on use of rivers, and possibly due to movement of deer.

In urban riparian areas, vegetation composition may be affected by urban land use changes at both the stream reach and the watershed scale. Moreover, the mechanisms by which seeds disperse may be affected both by reduction in seed sources due to vegetation removal and by the urban stream syndrome that produces flashier hydrographs and incised channels. I hypothesized that vegetation communities with high cover of native and hydrophilic species would be found in watersheds with high forest cover, while more limited cover of these species would be found in highly developed watersheds. Additionally, to examine the dispersal mechanisms contributing to these patterns, I hypothesized that 1) more seeds would be deposited in riparian areas by water than by wind; 2) the number of seeds deposited by streams would decrease as watershed urbanization increased; and 3) seeds deposited in the most urbanized sites would be primarily from species with traits favoring deposition by water, including large seed size and presence of a dispersal appendage. To investigate relationships between urban land cover types and riparian vegetation, I surveyed 30 randomly-selected riparian forests in the Portland-Vancouver metro area and related vegetation assemblages to watershed land cover. Vegetation was mapped to the nearest 1cm along three transects in each site. Land cover was characterized both within a 500m buffer around each site, and within the entire watershed. Relationships between land cover and vegetation assemblages were investigated using nonmetric multidimensional scaling and classification trees. To investigate the effect of watershed urbanization intensity on riparian seed deposition, I collected seeds deposited in nine riparian sites along a gradient of watershed total impervious area (TIA). I used a stratified-random approach to select sites. In each site, wind-deposited seeds were collected in funnel traps three times, and water-deposited seeds were collected in turf traps four times, over a 15-month period, spanning both wet and dry seasons. Consistent with my first hypothesis, communities dominated by native understory species were found exclusively in watersheds that were at least 15% forested by evergreen canopy. These findings suggest that native understory communities can persist in urban areas if adequate surrounding forest cover is maintained. Regarding my second major hypothesis, significantly more seeds were deposited by water than by wind (p < 0.05; mean of 155 seeds per turf trap; mean of 30 seeds per funnel trap). For shrubs, for species primarily dispersed by animals, and for species under 15m tall, hydrochory significantly increased delivery to riparian areas over the background seed delivery rate measured in funnel traps. There was a significant reduction in the number of seeds deposited by streams as TIA increased (adjusted R² = 0.74; p < 0.01). Deposition of shade-tolerant seeds decreased significantly, while deposition of non-native seeds increased significantly (p < 0.05) with watershed TIA and with development within 500m from the site, likely due to alterations of seed source pools of these species. Findings indicate that in an urban setting, small streams have the capacity to act as dispersal vectors, connecting fragmented populations that may otherwise be seedlimited. Riparian forests with diverse understory assemblages maintained by ongoing seed deposition may persist in urban areas with sufficient watershed forest, as well as with low development cover, in both the whole watershed and the near-stream area. Total seed deposition by streams, as well as deposition of shade-tolerant species, can be expected to decrease with increased watershed development. Results suggest that passive approaches to restoration of riparian forest understories in urban watersheds will only likely be successful with sufficiently high forest cover and with restricted development.

The thesis describes several unknown aspects of the spruce bark beetle, Ips typographus (L.), and the double-spined bark beetle, Ips duplicatus (Sahlberg.), life-cycles and ecophysiology. The first study focuses on I. typographus and its dispersal under epidemic conditions in the National park Šumava and introduces a novel method of fluorescent marking and detection of captured specimens. The second study is focused on ability of I. typographus to establish so-called sister-broods, i.e. re-emergence of females that are capable to continue egg laying without a need to mate again. The importance of sister-broods becomes apparent mainly in recent hot and dry vegetation seasons, which is demonstrated by a comparison of recent and historical data. The third study focuses on temperature-dependent development of I. duplicatus under laboratory conditions by the means of sandwich method. Altogether, the studies underline practical importance of ecophysiological studies on bark beetles as one of the tools for their effective management.

Survival and establishment of mangrove propagules at higher tidal elevations beyond the landward margin of their distribution is a requirement for the continued existence of mangrove populations in response to rising sea-level. Despite the growing body of literature that discusses mangrove recruitment patterns, few studies have empirically examined establishment and post-establishment growth success of propagules at the higher intertidal positions into witch mangrove populations are migrating. Using an experimental field approach, this study compares establishment and post-establishment growth success of propagules at three positions across a tidal elevation gradient within a landward-transgressing mangrove population of SW Florida (USA). I observed black mangrove, Avicennia germinans, recruitment as adults of this taxon may occupy low to high tidal elevations of the intertidal zone in SW Florida, indicating that propagules are capable of successfully establishing and reaching reproductive age in novel environments. However, establishment and post-establishment growth of A. germinans in encroachment areas landward to that of lower intertidal positions has not been examined. To accomplish this, I began by monitoring movement patterns of marked A. germinans propagules released at three intertidal positions during a high spring tide to confirm that propagule dispersal to encroachment areas located at higher tidal elevations occurred at the selected field site. Propagule survival, establishment success, and post-establishment growth rate of seedlings was monitored during a reciprocal transplant study utilizing two of the three intertidal positions, one representing a lower intertidal area within the mangrove population’s niche and one representing a higher intertidal area beyond the population’s landward margin. Regardless of parental tree origin, A. germinans propagules had greater establishment success in the lower intertidal position. Likewise mean seedling height was consistently greater among established seedlings in the lower intertidal although the difference in mean seedling height between tidal locations decreased linearly over the monitoring period. Propagule mortality was greatest at the higher intertidal position (27.5% of tethered propagules died) when compared to that in the lower intertidal (0.07%). Interestingly, the tidal position of propagule origin significantly influenced survival only during the first 33 days of the reciprocal transplant experiment. After this time interval and establishment as a seedling, no mortality was observed in either treatment position for 125 days. Together, results show that intraspecific variation in A. germinans propagule establishment and post-establishment seedling growth exists in landward transgressing populations across intertidal positions. My findings indicate that abiotic conditions of the higher intertidal environments into which mangroves are migrating may be detrimental for early life stages of A. germinans but not seedlings. Combined, my results suggest that investigations into mangrove success at novel intertidal positions should focus on limitations at the propagule life stage as there was no indication that survivorship varied among tidal elevation once mangrove seedlings were established. Finally, assessing maternal reserves of dispersing propagules may provide additional insight into the importance of mangrove propagule origin on initial survival.

Connectivity is a major contributor to the overall dynamics of marine populations. However, it still remains challenging to describe connectivity on ecologically meaningful scales of time and space. This is a major impediment to evaluating the impacts of marine protected area with respect to fisheries management objectives. This dissertation brings together a wide array of spatial and connectivity information in the Gulf of Mexico (GOM) with the goal of 1) understanding the spatial distribution of fish populations and source-sink dynamics and 2) evaluating whether this information can be integrated, through a modeling framework, to identify closed areas that could be beneficial to fisheries management in the Gulf of Mexico. First, a generalized additive modelling (GAM) approach is used to describe the distribution of a large number of species groups (i.e. functional groups) across the Gulf of Mexico (GOM) using a large fisheries independent data set (SEAMAP) and climate scale (decades) oceanographic conditions. Next a numerical Lagrangian particle transport model was developed that incorporates two major connectivity processes; site specific larval production and oceanographic transport for an entire large marine ecosystem and over multiple years. The two components are then combined to develop larval dispersal patterns for the entire GOM and identify areas operating as larval sources and sinks. Last, this information is integrated into an end-to-end ecosystem model to evaluate effectiveness of closing source and sink areas for the management of reef fish fisheries. Closed area managemeny simlautions for reef fish indicated closing reef fish source areas, as opposed to sinks, in the GOM is most efficient method of increasing total biomass and yield. However, the impacts across individual functional groups were site specific. Ultimately, these simulations demonstrate the inclusion of connectivity information could improve fishery management objectives in an ecosystem context.

In nature bacteria predominantly live on surfaces, in matrix-encased communities called biofilms. Biofilm formation displays dynamic developmental patterns resembling those of multicellular organisms. Using cooperative traits such as cell-cell signaling, bacteria in biofilms form complex architectures, known as microcolonies, in which cells become highly differentiated from their planktonic counterparts. Microcolonies are generally highly tolerant to bactericides, rendering biofilms extremely difficult to eradicate. The aim of this study was to investigate the last, and least understood stage of biofilm development, which involves the coordinated dispersal of single cells that revert to a free-swimming planktonic phenotype and escape from the biofilm. Strategies to induce biofilm dispersal are of interest due to their potential to prevent biofilms and biofilm-related infections. In the model organism Pseudomonas aeruginosa, reproducible patterns of cell death and dispersal can occur within biofilm structures, leaving behind empty or hollow microcolonies. These events were previously linked with the appearance of oxidative and/or nitrosative stress in mature microcolonies. Here, the involvement of reactive oxygen and nitrogen intermediates in biofilm development and dispersal processes was investigated in both mono- and mixed-species biofilms. By using specific fluorescent dyes and P. aeruginosa mutant strains, nitric oxide (NO), a by-product of anaerobic respiration and an important messenger molecule in biological systems, was found to play a major role in P. aeruginosa biofilm dispersal. Further, the results demonstrated that exposure to physiological, non-toxic concentrations of NO (in the low nanomolar range) causes biofilm dispersal in P. aeruginosa and restores its vulnerability to conventional antimicrobials. By using microarray techniques, NO was shown to induce global changes in genetic expression, including enhanced metabolic activity and motility and decreased adhesion and virulence in P. aeruginosa biofilms. The regulatory pathway implicated c-di-GMP, a newly discovered messenger molecule involved in the transition from sessility to motility in many bacterial species. NO-mediated dispersal was also observed in other single- and multi-species biofilms of clinically and industrially relevant organisms. Hence, the combined exposure to NO and bactericides was identified as a potential novel strategy for the removal of microbial communities, providing a low cost and environmentally safe solution to biofilm control.